Functional response in habitat selection and the tradeoffs between foraging niche components in a large herbivore

How herbivore behaviour is influenced by changes in resource levels is central for understanding trophic interactions. We examined whether foraging tradeoffs change with food levels by comparing habitat selection and space use within and between two neighbouring, predator‐free Svalbard reindeer populations. The populations faced different food levels due to contrasting grazing history. Summer resource selection in radiocollared females was assessed by a multi‐dimensional niche approach based on habitat variables obtained from a satellite image (e.g. the normalised difference vegetation index, NDVI) and a digital terrain model. The population at the overgrazed Brøggerhalvøya faced overall lower plant cover, biomass and primary productivity (i.e. lower NDVI) than the population at Sarsøyra. At Brøggerhalvøya, most reindeer selected for productive habitat when choosing home range and patches within the home range. In contrast, habitat selection at Sarsøyra was more affected by abiotic conditions such as moisture, which may influence plant quality. Here, reindeer used patches with even less biomass than the average reindeer at the poorer Brøggerhalvøya. Such a difference in habitat preference with different habitat availability (a functional response in habitat selection) probably reflected increased selection for high‐quality forage at the expense of high forage quantity at Sarsøyra. Accordingly, a negative relationship between habitat productivity and home range size was only present across individuals within Brøggerhalvøya, where forage quantity was the important foraging niche component. Individuals having poor (and large) home ranges apparently could not compensate for this by higher patch selectivity compared to individuals with richer home ranges. The results indicate changes in foraging tradeoffs at contrasting resource levels and that strong interactions occur between habitat selection, space use and the foraging niche structure in the absence of predation.     

Patch choice of avian herbivores along a migration trajectory–From Temperate to Arctic

Migratory waterfowl species seem to track temporal and spatial pulses of optimal forage availability on their way from temperate wintering to arctic breeding sites. In order to unravel the relative contribution of forage quality and forage biomass to foraging choices in avian herbivores, we experimentally manipulated biomass and quality of main forage plants through fertilisation and grazing exclusion at three sites along the flyway of barnacle geese, Branta leucopsis. Fertilisation increased the nitrogen content of the forage and grazing exclusion increased biomass levels. Manipulated plots were offered to wild geese in a random block experimental design and goose visitation was measured through dropping counts. At all sites there was a trend towards a higher preference of plots with increased quality and average biomass above plots with an average quality and increased biomass. Generally, geese preferred plots with highest standing crop of nitrogen. The numerical response of the geese to forage changes was supported by behavioural observations at the Baltic site. We conclude that for migrating barnacle geese the bottlenecks in the standing crop of nitrogen appear to lie in the limited biomass availability at the Baltic stopover site and the limited nutrient content of food in the Arctic breeding site, restricting the potential nutrient intake on these sites.     

Temporal scales of density‐dependent habitat selection in a large grazing herbivore

Habitat selection is a density‐dependent process, but little is known regarding how this relationship may vary across different temporal scales. Over long time scales, grazing shapes the structure, diversity and functioning of terrestrial ecosystems, and grazing‐induced changes in forage production over time are likely to affect the level of density dependence in habitat selection. In this fully‐replicated, landscape‐scale experiment, we investigated how density‐dependent habitat selection by a large grazing herbivore, sheep Ovis aries, develops over the time scale of a decade. We also address an often‐neglected challenge in habitat selection studies; namely, whether there is variation in use within a particular habitat or vegetation type and why. We found clear evidence of density dependence in habitat selection, with a wider use of habitats at high density. Despite a change in the standing biomass of high‐productivity vegetation at high herbivore density over the years, with herb biomass declining and graminoid biomass increasing, there was no clear evidence that these grazing‐induced changes in habitat over the years were strong enough to affect the level of density‐dependent habitat selection. The difference in selection for high versus low‐productivity habitats remained similar, despite annual fluctuations in the strength of selection. We found strong variation in selection within each vegetation type, even when vegetation types were mapped at a fine‐resolution scale. Our study shows that despite the interactive effects of herbivores and habitats, they are not always sufficiently strong enough to affect the level of density‐dependent habitat selection.     

Differences between resource patches modify root herbivore effects on plants

While soil resource heterogeneity and root herbivory can have significant direct influences on plant growth, soil heterogeneity may also have indirect effects by influencing the foraging behavior of root herbivores. We used sand-filled greenhouse pots to assess root herbivore foraging behavior and potential interactions between patch quality, herbivore foraging, and plant biomass production (yield). Individual pots were divided into four quarters: one fertilized, and three unfertilized, two of which were planted with tree seedlings. Two treatments were used to create fertilized quarters: high-organic manure fertilizer, and slow-release mineral fertilizer. Seedlings of red maple (Acer rubrum L.) and Virginia pine (Pinus virginiana L.) were used to create two single-species and one mixed-species treatments. Root-feeding beetle larvae were added to the pots and allowed to forage freely for ∼8 weeks. At harvest, root herbivores in organic-fertilized pots were strongly attracted to fertilized quarters despite their relatively low-root biomass. Herbivore distribution was significantly different in mineral fertilized pots, where larvae were most abundant in planted quarters, which is also where most of the plant roots occurred. Whole pot plant yield was significantly reduced by larvae; this effect was stronger in the mineral fertilized pots than organic fertilized pots. While one of the plant species appeared more sensitive to herbivory, root herbivores had a greater influence on yield in mixed-species pots than in single-species pots. Overall, these results suggest that patch quality influences on herbivore foraging may indirectly alter yield and plant community composition. Responsible Editor: Angela Hodge.     

Small mammalian herbivore determines vegetation response to patchy nutrient inputs

Nutrient inputs to plant communities are often spatially heterogeneous, for example those deriving from the dung and urine of large grazing animals. The effect of such localised elevation of nutrients on plant growth and composition has been shown to be modified by the grazing of large herbivores. However, there has been little work on interactions between small mammalian herbivores and such patchy nutrient inputs, even though these interactions are potentially of major significance for plant performance and community structure.
We examined the effect of simulated cattle urine deposition on the vegetation structure, above-ground biomass and species composition of chalk grassland within enriched patches. Short-term exclosures were used to determine whether a small herbivore (rabbit) would preferentially graze the vegetation in enriched patches and what impact this interaction would have on the performance of plants in such patches. Rabbit grazing pressure determined whether nutrient inputs had a negative or positive effect on plant biomass. Nutrients increased plant biomass in the absence of grazing, but when exposed to grazing, plants in nutrient-rich patches had more biomass consumed by herbivores than neighbouring plants. Further, nutrients increased the relative palatability of a less preferred forage species ( Brachypodium pinnatum ), contributing to changes in plant community composition. We conclude that a small herbivore can drive plant responses to patchily distributed nutrients.     

Density dependent and temporal variability in habitat selection by a large herbivore; an experimental approach

Both density dependent and density independent processes such as climate affect population dynamics in large herbivores. Understanding herbivore foraging patterns is essential to identify the underlying mechanisms behind variation in vital rates. However, very little is known about how animals vary their selection of habitat temporally, alone or in interaction with density during summer. At the foraging scale, we tested using a fully replicated experiment whether domestic sheep Ovis aries stocked at high (80  per  km²) and low (25  per  km²) densities (spatial contrasts) varied their habitat selection temporally over a four year period. We predicted reduced selection of high productivity vegetation types with increasing density, and that seasonal and annual variation in climate would affect this density dependent selection pattern by increasing competition for high quality habitats in late grazing season and in years with poor vegetation development and over time related to vegetation responses to grazing. As predicted from the Ideal free distribution model, selection of high productivity habitat decreased at high density. There was also a marked temporal variation in habitat selection. Selection of the most productive vegetation types declined towards the end of each grazing season, but increased over years both at low and high sheep density. There was only weak evidence for interactions, as selection ratio of highly productive habitats tended to increase more over years at low density as compared to high density. Limited interactive effects of density and annual variation on habitat selection during summer may explain why similar interactions in vital rates have rarely been reported for summer seasons. Our results are consistent with the view that variation in habitat selection is a central mechanism for climate and density related variation in vital rates.     

Cattle select African savanna termite mound patches less when sharing habitat with wild herbivores

African savanna termite mounds function as nutrient‐rich foraging hotspots for different herbivore species, but little is known about their effects on the interaction between domestic and wild herbivores. Understanding such effects is important for better management of these herbivore guilds in landscapes where they share habitats. Working in a central Kenyan savanna ecosystem, we compared selection of termite mound patches by cattle between areas cattle accessed exclusively and areas they shared with wild herbivores. Termite mound selection index was significantly lower in the shared areas than in areas cattle accessed exclusively. Furthermore, cattle used termite mounds in proportion to their availability when they were the only herbivores present, but used them less than their availability when they shared foraging areas with wild herbivores. These patterns were associated with reduced herbage cover on termite mounds in the shared foraging areas, partly indicating that cattle and wild herbivores compete for termite mound forage. However, reduced selection of termite mound patches was also reinforced by higher leafiness of Brachiaria lachnantha (the principal cattle diet forage species) off termite mounds in shared than in unshared areas. Taken together, these findings suggest that during wet periods, cattle can overcome competition for termite mounds by taking advantage of wildlife‐mediated increased forage leafiness in the matrix surrounding termite mounds. However, this advantage is likely to dissipate during dry periods when forage conditions deteriorate across the landscape and the importance of termite mounds as nutrient hotspots increases for both cattle and wild herbivores. Therefore, we suggest that those managing for both livestock production and wildlife conservation in such savanna landscapes should adopt grazing strategies that could lessen competition for forage on termite mounds, such as strategically decreasing stock numbers during dry periods.     

Predicting Extreme Droughts in Savannah Africa: A Comparison of Proxy and Direct Measures in Detecting Biomass Fluctuations,Trends and Their Causes

We monitored pasture biomass on 20 permanent plots over 35 years to gauge the reliability of rainfall and NDVI as proxy measures of forage shortfalls in a savannah ecosystem. Both proxies are reliable indicators of pasture biomass at the onset of dry periods but fail to predict shortfalls in prolonged dry spells. In contrast, grazing pressure predicts pasture deficits with a high degree of accuracy. Large herbivores play a primary role in determining the severity of pasture deficits and variation across habitats. Grazing pressure also explains oscillations in plant biomass unrelated to rainfall. Plant biomass has declined steadily and biomass per unit of rainfall has fallen by a third, corresponding to a doubling in grazing intensity over the study period. The rising probability of forage deficits fits local pastoral perceptions of an increasing frequency of extreme shortfalls. The decline in forage is linked to sedentarization, range loss and herbivore compression into drought refuges, rather than climate change. The results show that the decline in rangeland productivity and increasing frequency of pasture shortfalls can be ameliorated by better husbandry practices and reinforces the need for ground monitoring to complement remote sensing in forecasting pasture shortfalls.     

·荒漠草地植物多样性对草食动物采食的响应机制

草食动物采食对草地植物多样性和生态系统功能的影响机制是放牧生态学研究的核心问题。该研究以内蒙古锡林郭勒盟苏尼特右旗荒漠草原的长期放牧控制实验为平台, 从既有草地植物多样性和动物偏食性两个层面系统地研究了荒漠草地植物多样性对草食动物采食的响应机制。结果显示: 1)荒漠草地植物对草食动物采食呈现4种响应模式: 放牧“隐没种”、放牧“敏感种”、放牧“无感种”、“绝对优势种”; 2)在群落尺度上, 物种多样性指数随放牧强度增加而减少, 与不放牧小区相比, 重度放牧(HG)与适度放牧(MG)小区植物多样性均下降, 且这一规律同样适用于功能群多样性, 灌木半灌木这一功能群内物种多样性对放牧干扰较敏感; 3)在草地既有植物的基础上, 以不放牧小区为参考系, 草食动物对植物功能群偏食性的排序为: 一二年生草本(AB) >多年生杂类草(PF) >灌木半灌木(SS) >多年生禾草(PG), 且偏食性物种主要分布于AB和PF中; 4)植物多样性与动物偏食性基本呈显著负相关关系(p < 0.05)。     

Ecosystem engineering varies spatially: a test of the vegetation modification paradigm for prairie dogs

Colonial, burrowing herbivores can be engineers of grassland and shrubland ecosystems worldwide. Spatial variation in landscapes suggests caution when extrapolating single‐place studies of single species, but lack of data and the need to generalize often leads to ‘model system’ thinking and application of results beyond appropriate statistical inference. Generalizations about the engineering effects of prairie dogs (Cynomys sp.) developed largely from intensive study at a single complex of black‐tailed prairie dogs C. ludovicianus in northern mixed prairie, but have been extrapolated to other ecoregions and prairie dog species in North America, and other colonial, burrowing herbivores. We tested the paradigm that prairie dogs decrease vegetation volume and the cover of grasses and tall shrubs, and increase bare ground and forb cover. We sampled vegetation on and off 279 colonies at 13 complexes of 3 prairie dog species widely distributed across 5 ecoregions in North America. The paradigm was generally supported at 7 black‐tailed prairie dog complexes in northern mixed prairie, where vegetation volume, grass cover, and tall shrub cover were lower, and bare ground and forb cover were higher, on colonies than at paired off‐colony sites. Outside the northern mixed prairie, all 3 prairie dog species consistently reduced vegetation volume, but their effects on cover of plant functional groups varied with prairie dog species and the grazing tolerance of dominant perennial grasses. White‐tailed prairie dogs C. leucurus in sagebrush steppe did not reduce shrub cover, whereas black‐tailed prairie dogs suppressed shrub cover at all complexes with tall shrubs in the surrounding habitat matrix. Black‐tailed prairie dogs in shortgrass steppe and Gunnison's prairie dogs C. gunnisoni in Colorado Plateau grassland both had relatively minor effects on grass cover, which may reflect the dominance of grazing‐tolerant shortgrasses at both complexes. Variation in modification of vegetation structure may be understood in terms of the responses of different dominant perennial grasses to intense defoliation and differences in foraging behavior among prairie dog species. Spatial variation in the engineering role of prairie dogs suggests spatial variation in their keystone role, and spatial variation in the roles of other ecosystem engineers. Thus, ecosystem engineering can have a spatial component not evident from single‐place studies.     

Invertebrate,not small vertebrate,herbivory interacts with nutrient availability to impact tallgrass prairie community composition and forb biomass

The effects of herbivores and their interactions with nutrient availability on primary production and plant community composition in grassland systems is expected to vary with herbivore type. We examined the effects of invertebrate and small vertebrate herbivores and their interactions with nutrient availability on grassland plant community composition and aboveground biomass in a tallgrass prairie ecosystem. The abundance of forbs relative to grasses increased with invertebrate herbivore removals. This increase in forb abundance led to a shift in community composition, where invertebrate removals resulted in greater plant species evenness as well as a divergence in composition among plots. In contrast, vertebrate herbivore removals did not affect plant community composition or aboveground biomass. Nutrient additions alone resulted in a decrease in plant species richness and an increase in the abundance of the dominant grass, but the dominant grass species did not greatly increase in abundance when nutrient additions were combined with invertebrate removals. Rather, several subdominant forbs came to dominate the plant community. Additionally, the combined nutrient addition and invertebrate herbivore removal treatment increased forb biomass, suggesting that invertebrate herbivores suppress the responses of forb species to chronic nutrient additions. Overall, the release of forbs from invertebrate herbivore pressure may result in large shifts in species composition, with consequences for aboveground biomass and forage quality due to altered grass:forb ratios in grassland systems.     

Biodiversity and reduced extinction risks in spatially isolated rodent populations

Ecologists have rarely explored the potential influence of local (alpha) biodiversity on the stability and local extinction of spatially isolated populations. Twenty years of annual counts of a small, grazing rodent (Utah prairie dogs, Cynomys parvidens ) from 20 different isolated local populations (colonies) in southern Utah, U.S.A. were analysed. These prairie dogs exhibited large fluctuations and repeated extinctions at individual colonies during the census period. Frequency of extinction at a colony declined dramatically as the number of locally occurring plant species increased. This pattern was not explained by differences among colonies in plant productivity, plant species composition, colony size, or variability in annual counts. Thus, lower extinction risk of consumer populations may be associated with greater resource diversity, and maintaining high local plant diversity may help sustain spatially isolated herbivore populations in fragmented habitats.     

Interactive Effects of Black-Tailed Prairie Dogs and Cattle on Shrub Encroachment in a Desert Grassland Ecosystem

The widespread encroachment of woody plants throughout the semi-arid grasslands in North America has largely resulted from overgrazing by domestic livestock, fire suppression, and loss of native large and small mammalian herbivores. Burrowing-herbivorous mammals, such as prairie dogs (Cynomys spp.), help control shrub encroachment through clipping of shrubs and consumption of their seedlings, but little is known about how this important ecological role interacts with and may be influenced by co-existing large herbivores, especially domestic livestock. Here, we established a long-term manipulative experiment using a 2 × 2 factorial design to assess the independent and interactive effects of black-tailed prairie dogs (Cynomys ludovicianus) and cattle (Bos taurus) on honey mesquite (Prosopis glandulosa) abundance and structure. We found that, after five years, mesquite abundance was three to five times greater in plots where prairie dogs were removed compared to plots where they occurred together or alone, respectively. While both prairie dogs and cattle reduced mesquite cover, the effect of prairie dogs on reducing mesquite abundance, cover, and height was significantly greater than that by cattle. Surprisingly, cattle grazing enhanced prairie dog abundance, which, in turn, magnified the effects of prairie dogs on mesquite shrubs. Mesquite canopy cover per hectare was three to five times greater where prairie dogs and cattle were absent compared to where they occurred together or by themselves; whereas, cumulative mesquite height was two times lower on sites where prairie dog and cattle occurred together compared to where they occurred alone or where neither occurred. Data from our experimental study demonstrate that prairie dogs and moderate grazing by cattle can suppress mesquite growth, and, when their populations are properly managed, they may interact synergistically to significantly limit mesquite encroachment in desert grasslands.     

Disturbance regimes and mountain plover habitat in shortgrass steppe: Large herbivore grazing does not substitute for prairie dog grazing or fire

Restoring historical disturbance regimes to enhance habitat for grassland birds can conflict with livestock production goals and has been controversial because of uncertainty in the frequency and pattern of different disturbances prior to European settlement. We studied nesting habitat for the mountain plover (Charadrius montanus) in relation to prescribed fire, grazing by large herbivores (cattle), and grazing by black-tailed prairie dogs (Cynomys ludovicianus) in the shortgrass steppe of northeastern Colorado. Breeding mountain plovers primarily occurred on black-tailed prairie dog colonies or areas burned during the previous dormant season. Vegetation surrounding mountain plover nests and foraging locations was characterized by a fine-scale mosaic of prostrate (<4 cm tall) vegetated patches interspersed with >35% bare soil in a given square meter, with this fine-scale pattern distributed over a broad (>100-m radius) area. Mountain plovers rarely occupied grassland lacking prairie dogs or recent fire, but those that did selected sites with similar vegetation height and bare soil exposure as sites on burns and prairie dog colonies. Vegetation structure at mountain plover-occupied sites was also similar to random sites on burns and prairie dog colonies, but differed substantially from sites managed only with cattle. Intensive cattle grazing at twice the recommended stocking rate during spring (Mar–May) or summer (May–Oct) for 6 years produced significantly less bare soil than burns and prairie dog colonies, particularly following years with average or above-average precipitation. Thus, intensive cattle grazing did not substitute for prairie dog grazing or fire in terms of effects on vegetation structure and mountain plover habitat. Both prescribed burning and increased size and distribution of black-tailed prairie dog colonies appear to be effective and complementary means to manage for mountain plover breeding habitat in shortgrass steppe. Provision of mountain plover habitat has tradeoffs with traditional management for livestock production. Thus, managers need to clearly define desired outcomes for management to provide multiple ecosystem goods and services. © 2012 The Wildlife Society.     

Facilitative and competitive effects of a large species with defensive traits on a grazing-adapted,small species in a long-term deer grazing habitat

Plants can adapt to grazing environments by developing defensive traits, such as spines and toxins, or having a small phenotype, such as short and prostrate growth forms. This study examined facilitative and competitive interactions between species with different types of grazing adaptation. We predicted that large species with defensive traits sometimes protect grazing-adapted species without defensive traits from herbivores, but competitively suppress them overall. We conducted an experiment using fences and removals of an unpalatable plant in the long-term deer grazing habitat of Nara Park in Nara, Japan. We evaluated the seasonal variations in the facilitative and competitive effects of a defensive perennial, Urtica thunbergiana, on the growth, survival, reproduction, and final fitness of a small palatable annual species, Persicaria longiseta, during a growing season. The populations of the two species in the park have adapted to the grazed habitat by increasing the density of stinging hairs (Urtica) and developing inherently short shoots and small leaves (Persicaria). We found that Urtica individuals had facilitative effects on the growth of Persicaria individuals under grazing during a few periods of the growing season, but had neutral effects on survival and plant fitness throughout the season. In the fenced plots, Urtica had negative effects on the growth, survival, and reproduction of Persicaria. These results suggest that the relative importance of the facilitative and competitive effects of Urtica on Persicaria fluctuated due to seasonal variations in grazing pressure and vegetative productivity. Although well-defended plants often facilitate less-defended species, we conclude that the facilitative effects of Urtica on Persicaria are limited in a plant community with a long history of intensive grazing.     

Insect herbivory in an experimental agroecosystem: the relative importance of habitat area, fragmentation, and the matrix

Habitat area, fragmentation, and the surrounding matrix influence levels of herbivory in various ecosystems, but the relative importance of these effects has rarely been assessed. We compared levels of herbivory and densities of dominant arthropod herbivores (the hemipteran insects Agallia constricta, Empoasca fabae, Therioaphis trifolii, Lygus lineolaris and Halticus bractatus ) among experimental plots that varied in the area and fragmentation of clover habitat and the composition of the matrix (bare ground or grass) surrounding clover habitat. To assess levels of herbivory, we compared clover biomass within herbivore exclosures to the biomass accessible to herbivores. The area and fragmentation of clover habitat, as well as matrix composition, significantly influenced the collective densities of herbivores, although each species exhibited unique responses to habitat structure. Herbivory was strongest in plots with large (64  m²) as compared to small (16  m²) amounts of clover habitat. The difference in clover biomass between the inside and outside of exclosures increased significantly with increasing density of Empoasca fabae but was unrelated to the densities of the other herbivores, suggesting that Empoasca fabae was an exceptionally important herbivore in this system. This study supports the view that herbivore densities and herbivory generally increase with increasing area of plant monocultures, but emphasizes that levels of herbivory may be driven primarily by one or a few key herbivore species.     

Pyric Herbivory and the Nexus Between Forage,Fire and Native and Introduced Large Grazing Herbivores in Australian Tropical Savannas

Earth’s tropical savannas typically support high biomass of diverse grazing herbivores that depend on a highly fluctuating resource: high-quality forage. An annual wet–dry cycle, fire and herbivory combine to influence forage quality and availability throughout the year. In the savannas of northern Australia, a depauperate suite of large native (marsupial) herbivores (wallaroos [Osphranter spp.] and the agile wallaby [Notamacropus agilis]) compete for resources with non-native large herbivores introduced in the late nineteenth century, particularly bovines (feral and managed cattle [Bos spp.] and feral water buffalo [Bubalus bubalis]) that now dominate the landscape. Anecdotal reports of recent population declines of large macropods and negative impacts of bovines highlight the need to better understand the complex relationship between forage, fire and abundance of native and introduced large herbivores. The pyric herbivory conceptual model, which posits complex feedbacks between fire and herbivory and was developed outside Australia, predicts that native and introduced large herbivores will both respond positively to post-fire forage production in Australian savannas where they co-occur. We used grazing exclosures, forage biomass and nutrient analyses and motion-sensor camera-trapping to evaluate the overall robustness of the pyric herbivory model in the Australian context, specifically whether forage quantity and quality are impacted by herbivory, season and fire activity, and which forage attributes most influence large grazing herbivore abundance. Forage quantity, as measured by live, dead and total herbaceous biomass and proportion of biomass alive, was higher inside herbivore exclosures, even at relatively low densities of herbivores. Forage quality, as measured by fibre content, was not affected by herbivory, however, crude protein content of live herbaceous biomass was greater outside herbivore exclosures. Recent fire was an important predictor of all measures of forage quantity and quality. Recent fire occurrence decreased overall quantity (biomass) but increased quality (decreased fibre content and increased crude protein content); late dry season fires resulted in forage with the highest crude protein content. The predictions of the pyric herbivory conceptual model are consistent with observations of the feeding behaviour of introduced bovines and some large macropods in northern Australian savannas, lending support to the global generality of pyric herbivory in fire-prone grassy biomes.

     

Overcompensation and grazing optimisation in a swan–pondweed system?

1. The general notion is that negative effects of vertebrate herbivores on water plants, which play a key role in freshwaters, prevail, and that positive feedbacks of herbivores on plants are insignificant. 2. The most likely systems to find such positive feedbacks are those in which herbivores exert strong feeding pressures on plants during part of the year. Previous theoretical work has suggested that compensatory production occurs when migratory Bewick's swans forage on tubers of fennel pondweed. As a corollary, the swans can exploit the tubers down to a level that maximises their tuber yield. 3. In order to test these hypotheses, I measured pondweed tuber biomass on three occasions per year (just before and after foraging, and just before tuber sprouting) in three consecutive years. The 17 sampling sites in the Lauwersmeer (the Netherlands) were classified according to their silt content and water depth. Within four silt‐depth classes, I predicted for each year tuber biomass production and, from that, the optimum foraging threshold that would result in the maximum tuber biomass yield. 4. Water depth did not affect tuber production, and silt content only did in one of the 3 years. In accordance with overcompensation predictions, tuber production was higher at plots with moderate foraging pressures than at plots with little or no grazing. However, the winter and summer conditions following the swan foraging had large unpredictable effects on tuber mortality and production. 5. These results indicate that overcompensation by fennel pondweed occurs and that Bewick's swans are generally able to profit from it, albeit without fine‐tuning of the foraging threshold to the yield.     

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.     

Vertebrate herbivores and ecosystem control: cascading effects of faeces on tundra ecosystems

We tested the hypothesis that large herbivores manipulate their own food supply by modifying soil nutrient availability. This was investigated experimentally the impact of faeces on grasses, mosses and soil biological properties in tundra ecosystems. For this, we increased the density of reindeer Rangifer tarandus platyrhynchus faeces and studied the response of a tundra system on Spitsbergen to this single faecal addition treatment for four subsequent years. From the third year onwards faecal addition had unambiguously enhanced the standing crop of grasses, as evidenced by an increase in both shoot density and mass per shoot. Although reindeer grazing across experimental plots was positively related to the abundance of grasses in anyone year, the increase in grass abundance in fouled plots failed to result in greater grazing pressure in those plots. Faecal addition enhanced soil microbial biomass C and N, particularly under wet conditions where faecal decay rates were greatest, whilst grasses appeared to benefit from faeces under dry conditions. Whilst growth of grasses and soil microbial biomass were stimulated by faecal addition, the depth of the extensive moss layer that is typical of tundra ecosystems was significantly reduced in fouled plots four years after faecal addition. The greatest reduction in moss depth occurred where fouling increased soil microbial biomass most, suggesting that enhanced decomposition of moss by a more abundant microbial community may have caused the reduced moss layer depth in fouled plots. Our field experiment demonstrates that by the production of faeces alone, vertebrate herbivores greatly impact on both above‐ and belowground components of tundra ecosystems and in doing so manipulate their own food supply. Our findings verify the assertion that grazing is of fundamental importance to tundra ecosystem productivity, and support the hypothesis that herbivory is instrumental in promoting grasses whilst suppressing mosses. The widely observed inverse relationship between grass and moss abundance in the field may therefore reflect the long history of plant‐herbivore interactions in tundra ecosystems.