Grassland-herbivore interactions: How do grazers coexist?

We develop a new approach to modeling grazing systems that links foraging characteristics (intake and digestive constraints) with resource dynamics via the probability of encounter with different grass heights. Three complementary models are presented: the generation of a grass height structure through selective grazing; investigating the conditions for consumer coexistence; and, using a simplified resource structure, the consequences for consumer abundance. The main finding is that coexistence between grazers differing in body size is possible if a single-resource type becomes differentiated in its height structure. Large grazers can facilitate food availability for smaller species but with the latter being competitively dominant. The relative preference given to different resource partitions is important in determining the nature of population interactions. Large-body and small-body grazer populations can interact through competitive, parasitic, commensalist, or amensalist relationships, depending on the way they partition the resource as well as their relative populations and the dynamics of resource renewal. The models provide new concepts of multispecies carrying capacity (stock equilibrium) in grazed systems with implications for conservation and management. We conclude that consumer species are not substitutable; therefore, the use of rangeland management concepts such as "livestock units" may be inappropriate.

How simple grazing rules can lead to persistent boundaries in vegetation communities.

Natural landscape boundaries between vegetation communities are dynamically influenced by the selective grazing of herbivores. Here we show how this may be an emergent property of very simple animal decisions, without the need for any sophisticated choice rules etc., using a model based on biased diffusion. Animal grazing intensity is coupled with plant competition, resulting in reaction-diffusion dynamics, from which stable boundaries spontaneously emerge. In the model, animals affect their resources by both consumption and trampling. It is assumed that forage consists of two heterogeneously distributed competing resource species, one that is preferred (grass) over the other (heather) by the animals. The solutions to the resulting system of differential equations for three cases a) optimal foraging, b) random walk foraging and c) taxis-diffusion are presented. Optimal and random foraging gave unrealistic results, but taxis-diffusion accorded well with field observations. Persistent boundaries between patches of near-monoculture vegetation were predicted, with these boundaries drifting in response to overall grazing pressure (grass advancing with increased grazing and vice versa). The reaction-taxis-diffusion model provides the first mathematical explanation for such vegetation mosaic dynamics and the parameters of the model are open to experimental testing.

Limpet grazing and loss of Ascophyllum nodosum canopies on decadal time scales

The role of limpet grazing in preventing the development of algal canopies is a recurrent theme in intertidal ecology. Less is known about interactions of limpets with the long-term dynamics of established canopies. Aerial photographs indicate that intertidal canopy cover has declined over the past 44 yr in Strangford Lough, Northern Ireland. There has been a loss of the previously continuous cover of Ascophyllum nodosum (L.) Le Jolis in the mid-shore. A barnacles dominated assemblage now fills gaps in the A. nodosum canopy. The rates at which barnacle patches become established and grow have increased since 1990. Changes in canopy cover have been accompanied by increases in limpet densities since the 1980s. Measurements between 2003 and 2004 showed no increase in length of A. nodosum fronds when limpets Patella vulgata had access to the algal holdfasts. In contrast, when limpets were experimentally excluded from the holdfasts, there was net frond growth. In the Isle of Man, which is climatically similar to Strangford Lough but has fewer limpets, growth occurred regardless of limpet grazing. The breaking force for A. nodosum declined with increasing local densities of limpets. A. nodosum is a sheltered shore species, potentially vulnerable to changes in wave exposure. There is no evidence, however, that Strangford Lough has become windier over the past 3 decades. Variation in wave exposure among locations within the lough was not related to rates of barnacle patch creation or expansion, Limpet population density has increased following a series of mild winters. Climate change may have a role in causing canopy loss, not by direct effects on the limpet populations.

Individual trade-offs between nutrition and risk of interspecific transmission of disease by grazing: cows, badger latrines and bovine tuberculosis

1. We examine whether various measures of herbivore current physiological state (age, breeding and immune status) and genetic potential can be used as indicators of exposure to and risk from disease. We use dairy cattle and the risks of tuberculosis (TB) transmission posed to them by pasture contaminated with badger excreta (via the fecal-oral route) as a model system to address our aim.

Risk of disease from wildlife reservoirs: Badgers, cattle, and bovine tuberculosis

Livestock face complex foraging options associated with optimizing nutrient intake while being able to avoid areas posing risk of parasites or disease. Areas of tall nutrient-rich swards around fecal deposits may be attractive for grazing, but might incur fitness costs from parasites. We use the example of dairy cattle and the risks of tuberculosis transmission posed to them by pastures contaminated with badger excreta to examine this trade-off. A risk may be posed either by aerosolized inhalation through investigation or by ingestion via grazing contaminated swards. We quantified the levels of investigation and grazing of 150 dairy cows at badger latrines (accumulations of feces and urine) and crossing points (urination-only sites). Grazing behavior was compared between strip-grazed and rotation-grazed fields. Strip grazing had fields subdivided for grazing periods of

How fragmented was the British Holocene wildwood? Perspectives on the “Vera” grazing debate from the fossil beetle record

The reconstruction and structure of the European Holocene “wildwood” has been the focus of considerable academic debate. The ability of palaeoecological data and particularly pollen analysis to accurately reflect the density of wildwood canopy has also been widely discussed. Fossil insects, as a proxy for vegetation and landscape structure, provide a potential approach to address this argument. Here, we present a review and re-analysis of 36 early and mid-Holocene (9500-2000 cal BC) sub-fossil beetle assemblages from Britain, examining percentage values of tree, open ground and dung beetles as well as tree host data to gain an insight into vegetation structure, the role of grazing animals in driving such structure and establish independently the importance of different types of trees and associated shading in the early Holocene “wildwood”. Open indicator beetle species are persistently present over the entire review period, although they fluctuate in importance. During the early Holocene (9500-6000 cal BC), these indicators are initially high, at levels which are not dissimilar to modern data from pasture woodland. However, during the latter stages of this and the next period, 6000-4000 cal BC, open ground and pasture indicators decline and are generally low compared with previously. Alongside this pattern, we see woodland indicators generally increase in importance, although there are significant local fluctuations. Levels of dung beetles are mostly low over these periods, with some exceptions to this pattern, especially towards the end of the Mesolithic and in floodplain areas. Host data associated with the fossil beetles indicate that trees associated with lighter canopy conditions such as oak, pine, hazel and birch are indeed important components of the tree canopy during the earlier Holocene (c. 9500-6000 cal BC), in accordance with much of the current pollen literature. Beetles associated with more shade-tolerant trees (such as lime and elm) become more frequent in the middle Holocene (6000-4000 cal BC) suggesting that at this stage the woodland canopy was less open than previously, although open ground and pasture areas appear to have persisted in some locations. The onset of agriculture (4000-2000 cal BC) coincides with significant fluctuations in woodland composition and taxa. This is presumably as a result of human impact, although here there are significant regional variations. There are also increases in the amounts of open ground represented and especially in the levels of dung beetles present in faunas, suggesting there is a direct relationship between the activities of grazing animals and the development of more open areas. One of the most striking aspects of this review is the variable nature of the landscape suggested by the palaeoecological data, particularly but not exclusively with the onset of agriculture: some earlier sites indicate high variability between levels of tree-associated species on the one hand and the open ground beetle fauna on the other, indicating that in some locations, open areas were of local significance and can be regarded as important features of the Holocene landscape. The role of grazing animals in creating these areas of openness was apparently minimal until the onset of the Neolithic.

Mammals and agri-environment schemes: hare haven or pest paradise?

1. Agri-environment schemes (AESs) are designed to create landscape-scale improvements in biodiversity. While the specific aims of AESs do not always include the enhancement of species of conservation concern, associated conservation strategies, such as the UK Biodiversity Action Plan, often rest on the assumption that AESs enhance environmental conditions and thereby improve the conservation status of target species. However, there is little evidence for the general efficacy of AESs in this respect. 2. To evaluate the effects of the Environmentally Sensitive Area (ESA) scheme, a widespread AES in Northern Ireland, a spotlight survey of the relative abundance of three mammal species, Irish hare Lepus timidus hibernicus, European rabbit Oryctolagus cuniculus and red fox Vulpes vulpes, was conducted. Of these, the Irish hare is a priority species for conservation action and the focus of a species action plan, while rabbit and fox are commonly considered agricultural pests. The effects of ESA designation and habitat on each species were assessed at 150 ESA and 50 non-ESA sites, matched for landscape characteristics. 3. The ESA scheme had no demonstrable effect on the abundance of Irish hares, and this agri-environment scheme did not target the landscape and habitat variables associated with hares. 4. In contrast, the abundance of rabbits and foxes was significantly greater within ESAs than the wider countryside. Agricultural factors such as reduced livestock stocking density, reduced overgrazing and field boundary enhancements may create more favourable conditions for both species. Aside from the implications for farm economics, the proliferation of rabbit populations within conservation areas may raise issues concerning the grazing of important plant communities, while increases in fox populations may adversely affect ground-nesting birds and other animal species of conservation concern. 5. Synthesis and applications: The abundance of rabbits and foxes corroborates recent work that suggests AESs may benefit common species but can not be relied upon to encourage rarer species. The Irish hare species action plan relies on agri-environment schemes to enhance the species’ status and realize the target of increasing the hare population by 2010 by promoting suitable habitat. However, the ESA scheme is unlikely to help in achieving these objectives. Targeted and evidence-based agri-environment prescriptions are clearly required in order to ensure the realization of species-specific conservation targets.