Impact of herbivores on nitrogen cycling: contrasting effects of small and large species

Herbivores are reported to slow down as well as enhance nutrient cycling in grasslands. These conflicting results may be explained by differences in herbivore type. In this study we focus on herbivore body size as a factor that causes differences in herbivore effects on N cycling. We used an exclosure set-up in a floodplain grassland grazed by cattle, rabbits and common voles, where we subsequently excluded cattle and rabbits. Exclusion of cattle lead to an increase in vole numbers and a 1.5-fold increase in net annual N mineralization at similar herbivore densities (corrected to metabolic weight). Timing and height of the mineralization peak in spring was the same in all treatments, but mineralization in the vole-grazed treatment showed a peak in autumn, when mineralization had already declined under cattle grazing. This mineralization peak in autumn coincides with a peak in vole density and high levels of N input through vole faeces at a fine-scale distribution, whereas under cattle grazing only a few patches receive all N and most experience net nutrient removal. The other parameters that we measured, which include potential N mineralization rates measured under standardized laboratory conditions and soil parameters, plant biomass and plant nutrient content measured in the field, were the same for all three grazing treatments and could therefore not cause the observed difference. When cows were excluded, more litter accumulated in the vegetation. The formation of this litter layer may have added to the higher mineralization rates under vole grazing, through enhanced nutrient return through litter or through modification of microclimate. We conclude that different-sized herbivores have different effects on N cycling within the same habitat. Exclusion of large herbivores resulted in increased N annual mineralization under small herbivore grazing.     

Endozoochory by free-ranging, large herbivores: Ecological correlates and perspectives for restoration

Seed dispersal via ingestion and defecation by large herbivores provides a possible aid for ecological restoration of plant communities, by connecting source communities of target species with habitat restoration sites. It is also a possible threat due to invasion of weeds, grasses or exotic species. Insight into the factors determining internal seed dispersal could therefore improve the management of grazed ecosystems.

We recorded viable seed density in cattle, sheep and pony dung samples and monitored dung pat colonisation in the field. In addition, we counted the distribution of dung pats in plots spread over all habitat units in our study site.

The three herbivore species appeared to disperse large quantities of many species (61 in total) from a variety of plant families, monocots as well as dicots. The density of viable seeds in herbivore dung and the colonisation of dung pats were positively correlated with Ellenberg nitrogen indicator values and seed supply, but not with seed mass or shape.

The results imply that many seeds are dispersed from high productive to low productive parts of the grazed area. In free-ranging systems, we therefore recommend enclosure and separate management of plant communities on nutrient-poor soils with high conservation interest. For habitat restoration sites we recommend integrated grazing only with target plant communities on nutrient-poor soils and not with plant communities on nutrient-richer soils.     

Interacting effects of grass height and herbivores on the establishment of an encroaching savanna shrub

Shrub encroachment is a widely observed problem in Southern African savannas. Although the effects of herbivory and grass height on woody species recruitment have been studied individually, little information exists about how these factors interact. In this study seeds and seedlings of the encroaching shrub Dichrostachys cinerea were planted in clipped and unclipped grass plots, with and without large herbivores present. Seed germination, seedling survival and seedling predation were monitored for 8 months. Germination started earlier in plots where herbivores were excluded. Overall, the earlier the seeds germinated, the longer the seedlings survived. Clipping positively affected the number of germinated seeds, seedling growth and survival but effects varied among herbivore exclusion treatments and sites. Invertebrates caused the majority of the seedling damage. We conclude the recruitment of D. cinerea is influenced by the interplay of grass height and herbivory. In this study, the presence of large herbivores early in the wet season, and the absence of simulated grazing later on, affected the regeneration of D. cinerea negatively. However, differences in effects among sites suggest that the mechanisms found here may work differently in other habitats.      

Rapid, landscape scale responses in riparian tundra vegetation to exclusion of small and large mammalian herbivores

Productive tundra plant communities composed of a variety of fast growing herbaceous and woody plants are likely to attract mammalian herbivores. Such vegetation is likely to respond to different-sized herbivores more rapidly than currently acknowledged from the tundra. Accentuated by currently changing populations of arctic mammals there is a need to understand impacts of different-sized herbivores on the dynamics of productive tundra plant communities. Here we assess the differential effects of ungulate (reindeer) and small rodent herbivores (voles and lemmings) on high productive tundra vegetation. A spatially extensive exclosure experiment was run for three years on river sediment plains along two river catchments in low-arctic Norway. The river catchments were similar in species pools but differed in species abundance composition of both plants and vertebrate herbivores. Biomass of forbs, deciduous shrubs and silica-poor grasses increased by 40–50% in response to release from herbivory, whereas biomass of silica-rich grasses decreased by 50–75%. Hence both additive and compensatory effects of small rodents and reindeer exclusion caused these significant changes in abundance composition of the plant communities. Changes were also rapid, evident after only one growing season, and are among the fastest and strongest ever documented in Arctic vegetation. The rate of changes indicates a tight link between the dynamics of productive tundra vegetation and both small and large herbivores. Responses were however not spatially consistent, being highly different between the catchments. We conclude that despite similar species pools, variation in plant species abundance and herbivore species dynamics give different prerequisites for change.     

Seasonal impact of waterfowl on communities of macrophytes in a shallow lake

From 2005 to 2007, we established bird-proof enclosures in a small, shallow and semi-permanent lake, lacking fish, at Brown Moss, Shropshire, UK, to investigate the effects of aquatic birds on seasonal growth of submerged and emergent macrophytes. The highest density of birds on the lake was in winter (110 individuals ha⁻¹) and the lowest in summer 2005 (6 ha⁻¹). Plant growth varied with season but there were significantly different (F = 8.03, p < 0.05, df = 1) standing crops of macrophytes between bird-proof enclosures (proportion of volume occupied, 0.47 ± 0.04) and control treatments (0.36 ± 0.11). Different densities of birds occurred in different areas and this was reflected in their effects. Ducks, mainly mallard (Anas platyrhynchos, Linnaeus), and teal (Anas crecca, Linnaeus), damaged plants by direct consumption, uprooting and trampling, whereas larger birds, such as mute swan (Cygnus olor, Gmelin), were able to remove Typha latifolia (Linnaeus). In summer, grazing pressure was reduced as the population of birds declined. Waterfowl caused seasonal impacts on the re-development of the water plant community. However, waterfowl herbivory had low potential to shift a macrophyte-dominated state into a phytoplankton-dominated state because aquatic plants could recover, during the growing season, when bird populations declined.     

The impact of grazing and human disturbance on the dynamics of woodland in S. W. Ireland

Abstract. Pollen data from three mor humus profiles taken from within native woodland in the Killarney National Park are presented. The pollen records illustrate the woodland history over about 250 years. The dynamics of woodland on the mainland are compared with those of a small ungrazed lake island. Human disturbance of the woodland in the form of felling and burning is recorded at both locations. Following the disturbance, open vegetation predominated in which Arbutus unedo was widespread. This species declined as canopy woodland re-developed. The pollen records illustrate the dynamics of the developing woodland during which the role of Ilex aquifolium and Taxus baccata was strongly influenced by grazing. The past importance of Taxus baccata in western Irish woodland is considered.     

Effects of two types of herbivores on the population dynamics of a perennial herb

The joint effects of multiple herbivores on their shared host plant have received increasing interest recently. The influence of herbivores on population dynamics of their host plants, especially the relative roles of different types of damage, is, however, still poorly understood. Here, we present a modelling approach, including both deterministic and stochastic matrix modelling, to be used in estimating fitness effects of multiple herbivores on perennial plants. We examined the effects and relative roles of two specialist herbivores, a pre-dispersal seed predator, Euphranta connexa, and a leaf-feeding moth, Abrostola asclepiadis, on the population dynamics and long-term fitness of their shared host plant, a long-lived perennial herb Vincetoxicum hirundinaria (Asclepiadaceae). We collected demographic data during 3 years and combined these data with the effects of natural levels of herbivory measured from the same individuals. We found that both seed predation and leaf herbivory reduced population growth of V. hirundinaria, but only very high damage levels changed the growth trend of the vigorously growing study populations from positive to negative. Demographic modelling indicated that seed predation had a greater impact on plant population growth than leaf herbivory. The effect of leaf herbivory was weaker and diminished with increasing level of seed predation. Evaluation of individual fitness components, however, suggested that leaf herbivory contributed more strongly to host plant fitness than seed predation. Our results emphasize that understanding the effects of a particular herbivore on plant population dynamics requires also knowledge on other herbivores present in the system, because the effect of a particular type of herbivory on plant population dynamics is likely to vary according to the intensity of other types of herbivory. Furthermore, evaluating herbivore impact from using individual fitness components does not necessarily reflect the long-term effects on total plant fitness.     

Large herbivores influence the composition and diversity of shrub-steppe communities in the Rocky Mountains, USA

It is widely believed that wild and domestic herbivores have modified the structure and composition of arid and semi-arid plant communities of western North America, but these beliefs have rarely been tested in long-term, well-replicated studies. We examined the effects of removing large herbivores from semi-arid shrublands for 40–50 years using 17 fenced exclosures in western Colorado, USA. Shrub cover was greater (F=5.87, P=0.0020) and cover (F=3.01, P=0.0601) and frequency (F=3.89, P=0.0211) of forbs was less inside the exclosures (protected) relative to grazed plots. However, we found no significant effects (minimum P=0.18) of protection from grazing on cover or frequency of grasses, biotic crusts, or bare soil. Although mean species richness and diversity were similar between treatments, protected areas had much higher dominance by fewer species, primarily sagebrush. Exclusion of herbivores changed the relationship between species richness and evenness. Consistent with theoretical expectations, species evenness was positively correlated with richness in protected plots (r ²=0.54). However, contrary to theory, evenness and richness were inversely related in grazed plots (r ² _adjacent=0.72, r ² _distant=0.84). We suggest that these differences resulted because grazing acts as a stressor promoting facilitative relationships between plant species that might compete for resources in the absence of grazing. We conclude that exclusion of grazing in the sites we studied caused minor changes in cover and diversity of herbaceous plants, but caused a clear increase in the cover of shrubs. Importantly, the exclusion of ungulates changed the relationship between evenness and richness.     

Effects of spines and thorns on Australian arid zone herbivores of different body masses

Summary We investigated the effects of thorns and spines on the feeding of 5 herbivore species in arid Australia. The herbivores were the rabbit (Oryctolagus cuniculus), euro kangaroo (Macropus robustus), red kangaroo (Macropus rufus), sheep (Ovis aries), and cattle (Bos taurus). Five woody plants without spines or thorns and 6 woody plants with thorns were included in the study. The spines and thorns were not found to affect the herbivores' rates of feeding (items ingested/min), but they did reduce the herbivores' rates of biomass ingestion (g-dry/item). The reduction in biomass ingested occurred in two ways: at a given diameter, twigs with spines and thorns had less mass than undefended plants, and the herbivores consumed twigs with smaller diameters on plants with spines and thorns. The relative importance of the two ways that twigs with spines and thorns provided less biomass varied with herbivore body mass. Reduced twig mass was more important for small herbivores, while large herbivores selected smaller diameters. The effectiveness of spines and thorns as anti-herbivore defenses did not vary with the evolutionary history of the herbivores (i.e. native vs. introduced). Spines and thorns mainly affected the herbivores' selection of maximum twig sizes (reducing diameter and mass), but the minimum twig sizes selected were also reduced.     

Plant community response to loss of large herbivores: comparing consequences in a South African and a North American grassland

Loss of biodiversity poses one of the greatest threats to natural ecosystems throughout the world. However, a comprehensive understanding of the impacts of species losses from upper trophic levels is still emerging. Here we compare the impacts of large mammalian herbivore species loss on grassland plant community structure and composition in a South African and North American grassland. Herbaceous plant communities were surveyed at sites without large mammalian herbivores present and at sites with a single species of herbivore present in both locations, and additionally at one site in South Africa with multiple herbivore species. At both the North American and South African locations, plant communities on sites with a single herbivore species were more diverse and species rich than on sites with no herbivores. At the multi-herbivore site in South Africa, plant diversity and richness were comparable to that of the single herbivore site early in the growing season and to the no herbivore site late in the growing season. Analyses of plant community composition, however, indicated strong differences between the multi-herbivore site and the single and no herbivore sites, which were more similar to each other. In moderate to high-productivity ecosystems with one or a few species of large herbivores, loss of herbivores can cause a significant decrease in plant diversity and richness, and can have pronounced impacts on grassland plant community composition. In ecosystems with higher herbivore richness, species loss may also significantly alter plant community structure and composition, although standard metrics of community structure may obscure these differences.     

The impact of grazing on plant communities,plant populations and soil conditions on salt marshes

Grazing an abandoned salt marsh causes retrogressive succession, since mid salt-marsh communities change into lower salt-marsh communities. Grazing and mowing are compared in detail. Both management practices enhance species diversity in an abandoned salt marsh. This can be attributed to the removal of litter. The finding that lower salt-marsh species appear more with grazing than with mowing or abandoning is not related to a higher soil salinity as compared to mowing or abandoning, but probably to locally baring of the soil by grazing animals. Only species of pioneer or unstable environments seem to have a persistent seed bank, for other species seed dispersal seems to be a limiting factor for their establishment.Nomenclature follows Heukels & van Ooststroom (1977) for species; Westhoff & den Held (1969) for syntaxa.Mrs R. Rusthoven analyzed the soil samples, Mr E. Leeuwinga drawed the figures, and Mrs J. O'Brien corrected the English text.     

Effects of quantitative and qualitative variation in phenolic compounds on feeding in three species of marine invertebrate herbivores

The deterrent effects of brown algal phenolic compounds and the terrestrial polyphenolic tannic acid on feeding by three species of invertebrate herbivores from central California, including the gastropods Tegula funebralis (Adams) and Tegula brunnea (Phillipi) and the echinoid Strongylocentrotus purpuratus (Stimpson) were examined. Algal phenolics used were the monomeric phenolic phloroglucinol and polyphloroglucinols from Fucus vesiculosus (Linnaeus), Halidrys siliquosa (Linnaeus) Lyngbye and Eisenia arborea Areschoug. All of the polyphenolics deterred feeding by all three herbivores at concentrations of 5 mg · ml⁻¹ in agar disks. Concentrations of 2 mg · ml⁻¹ also generally deterred feeding by the gastropods (these levels were not tested against S. purpuratus). Relative amounts of deterrence by different compounds were similar, especially for the gastropods. Phloroglucinol deterred feeding by the echinoids, but not by T. funebralis. Responses of the echinoids were otherwise similar to the gastropods, but more variable. I also demonstrated deterrence of S. purpuratus by tannic acid using the “tanned” kelp technique of Steinberg (1985). Reactivity of the different phenolic compounds in the Folin-Denis procedure, a common colorimetric assay used to estimate levels of phenolics in plant tissue, was similar. This suggests that measuring phenolic levels in brown algae by this technique will not be greatly confounded by the occurrence of different kinds of phenolic molecules in different brown algae. This result, in combination with the similarity of the deterrent effects of the compounds used in this study, increases the validity of previous studies in the northeastern Pacific Ocean which correlate algal phenolic levels and diets or feeding preferences of invertebrate herbivores. For plants and herbivores in this region, this assay is a reasonable measure of a biologically meaningful phenomenon — levels of phenolic deterrents in the algae.     

Aggregative responses of brent geese on salt marsh and their impact on plant community dynamics

The aggregative responses and habitat preferences of a generalist herbivore, the dark-bellied brent goose Branta bernicla bernicla, feeding on salt marshes are examined in relation to vegetation community characteristics and the abundances of individual plant species. In the autumn, feeding was strongly concentrated on the low marsh, which had the highest biomass of the preferred food plant, Salicornia europaea. There was a strong aggregative response of the geese to the abundance of S. europaea. A decline in the availability of S. europaea led to an increase in the pattern of aggregation in relation to the two other major food plants on the low marsh, Aster tripolium and Puccinellia maritima. The availability of these food plants, however, reached critically low levels in mid-winter and the geese abandoned the low marsh for the high marsh. Within the high marsh, the plant communities selected tended to be dominated by the inedible species Limonium vulgare. The food plants selected were P. maritima in the winter and P. maritima and Triglochinmaritimum in the spring. On the high marsh, aggregative responses were shown to both P. maritima and T. maritimum, but in both cases, aggregation increased up to a critical level of biomass, and then declined. The prevention of grazing with exclosures for 3 years led to an increase in the abundance of P. maritima on both high and low marshes. This change was slight on high marsh but pronounced on low marsh, where S. europaea showed a decrease in abundance in the exclosures over this time. The implications of the aggregative responses for the population dynamics of P. maritima and S. europaea are discussed. Received: 11 September 1997 / Accepted: 12 December 1997     

Conservation implications of grazing practices on the plant and dipteran communities of a turlough in Co. Mayo,Ireland

Turloughs, which are classified as priority habitats under the European Habitats Directive, are seasonally flooded depressions found almost exclusively in Ireland. In 2001, three adjacent fields with different stocking densities were selected and plant/dipteran communities within the same vegetation zone of each field (site) were investigated using quadrats and sweep netting, respectively. There was a significant positive relationship between Diptera morphospecies richness/Diptera abundance and mean vegetation height (P < 0.001). However, no significant relationship between Diptera morphospecies richness and plant species richness was found. Median Diptera morphospecies richness per sweep was lower at the site with the highest stocking density (17) than at the other two sites (22 and 31, respectively). Total species richness of Sciomyzidae was greater at the least grazed site (7) than at the more heavily grazed sites (2 and 1, respectively). The results suggest that an evaluation of turlough management practices based on plant communities alone is not sufficient and that at least some areas within the turlough basin remain ungrazed on a rotational basis to ensure maximum diversity of Diptera.     

Disproportionate effects of non‐colonial small herbivores on structure and diversity of grassland dominated by large herbivores

The response of semiarid grasslands to small, non‐colonial herbivores has received little attention, focusing primarily on the effects of granivore assemblages on annual plant communities. We studied the long‐term effects of both small and large herbivores on vegetation structure and species diversity of shortgrass steppe, a perennial semiarid grassland considered marginal habitat for small mammalian herbivores. We hypothesized that 1) large generalist herbivores would affect more abundant species and proportions of litter‐bare ground‐vegetation cover through non‐selective herbivory, 2) small herbivores would affect less common species through selective but limited consumption, and 3) herbivore effects on plant richness would increase with increasing aboveground net primary production (ANPP). Plant community composition was assessed over a 14‐year period in pastures grazed at moderate intensities by cattle and in exclosures for large (cattle) and large‐plus‐small herbivores (additional exclusion of rabbits and rodents). Exclusion of large herbivores affected litter and bare ground and basal cover of abundant, common and uncommon species. Additional exclusion of small herbivores did not affect uncommon components of the plant community, but had indirect effects on abundant species, decreased the cover of the dominant grass Bouteloua gracilis and total vegetation, and increased litter and species diversity. There was no relationship between ANPP and the intensity of effects of either herbivore body size on richness. Exclusion of herbivores of both body sizes had complementary and additive effects which promoted changes in vegetation composition and physiognomy that were linked to increased abundance of tall and decreased abundance of short species. Our findings show that small mammalian herbivores had disproportionately large effects on plant communities relative to their small consumption of biomass. Even in small‐seeded perennial grasslands with a long history of intensive grazing by large herbivores, non‐colonial small mammalian herbivores should be recognized as an important driver of grassland structure and diversity.     

Cattle grazing, raptor abundance and small mammal communities in Mediterranean grasslands

Abundance and diversity of small mammals are usually affected strongly by grazing either due to decreased food availability or quality, decreased suitability of soil for building burrow systems due to trampling and/or due to increased predation risk in the structurally simpler grazed areas. We estimated the effects of grazing-induced changes in vegetation and soil and of increased predation on small mammals in a Mediterranean grassland landscape. We measured vegetation structure, soil compaction and small mammal abundance and species composition in 22 plots of 8 Sherman live traps each, arranged according to an unbalanced two-way ANOVA design with two grazing levels (grazed areas and cattle exclosures) and two predator abundance levels (increased densities of Eurasian kestrels Falco tinnunculus by means of nest boxes and control). Plots were sampled during 2 consecutive years in early summer and early fall. Exclosure from cattle increased significantly vegetation height and volume and decreased soil compaction. Grazing-induced changes in vegetation height and volume and in soil compaction produced strong effects on small mammal abundance and species richness. Increased kestrel densities did not have significant additive or interactive effects, with the effects of grazing-induced vegetation and soil gradients on abundance or richness of small mammals. Our results suggest that the effects of grazing on small mammal communities in Mediterranean montane grasslands were mainly due to reduced food availability and by negative effects of trampling on the suitability of soils for building burrow systems. Decreased food quality and increased predation in grazed areas seemed to play a minor role, if any. Reductions in stock densities would then favor generalist predator populations in Mediterranean grasslands through the expected positive effects of such reductions on the availability of food and burrows for small mammals.     

A tentative typology of European scrub and forest communities based on vegetation texture and structure

Utilitarian and fundamental, floristic and structural classifications of forest and scrub communities are compared. Their advantages and disadvantages and their interrelations are discussed. The necessity of a structural classification as complementary to a floristic classification is emphasized. It seems difficult, if not impossible, to combine the two. The existing structural classifications are reviewed. They are not very detailed and often mixed with a habitat classification. A new, detailed and purely structural system for woods and scrub in Europe is proposed in the form of an identification key. The main criteria used are — in order of diminishing importance —: (1) architecture of the dominant layer, (2) photoperiodicity (evergreen vs. deciduous) of that layer, (3) leaf size and leaf form of that layer, (4) presence/absence of thorns/spines, (5) presence of a second tree layer, shrub layer, dwarf shrub layer, herb layer and/or moss layer, (6) deciduousness of these layers, (7) main growth forms, (8) leaf size and leaf consistency, (9) leaf inclination and (10) height of these layers.In all 27 scrub types and 45 wood types are described. They have been named after a combination of two more or less characteristic and dominant genera. The system is open and hierarchical. Broader units have also been described.Comm. No. 388 of the Biological Station Wijster, Department of Silviculture and Forest Ecology, Agricultural University, Wageningen     

Indirect effects of cattle grazing on shrub spatial pattern in a mediterranean scrub community

Identifying the mechanisms and interactions that influence the spatial structure of vegetation is important for both scientific and practical purposes. Grazing is one of the most fundamental interactions in ecology but so far its effect on vegetation spatial pattern received little attention. In this study we propose a conceptual model that can be used to predict the effect of grazing on shrub spatial pattern in water-limited ecosystems where shrubs grow within a matrix of annual vegetation. According to the model, grazing may increase or decrease clumping in shrub distribution, depending on (1) the relative palatability of shrubs vs. annual plants to the herbivores, and (2) the manner (negative or positive) by which adult shrubs and annual plants affect the establishment of shrub seedlings. We tested our model in a Mediterranean scrub ecosystem by analyzing the development of shrub spatial pattern over a period of 40 years in plots characterized by contrasting intensities of cattle grazing. As predicted by the model, all plots showed a clumped pattern of shrub distribution in the absence of cattle grazing while intense cattle grazing reduced the clumpiness of the vegetation and generated a more random pattern of shrub distribution. Interestingly, plots representing the two grazing regimes did not differ significantly in their shrub cover, suggesting that shrub spatial pattern may be more sensitive to grazing than overall shrub cover.     

Effects of herbivores, nutrient enrichment, and their interactions on macroalgal proliferation and coral growth

We conducted a 20-week manipulative field experiment on shallow forereefs of the Florida Keys to assess the separate and interactive effects of herbivory and nutrient enrichment on the development of macroalgal communities and the fitness of the corals Porites porites and Siderastrea siderea. Excluding large herbivorous fishes produced macrophyte blooms both with and without nutrient enrichment. In contrast, there were no direct effects of nutrient enrichment. There were, however, small, but significant, interactive effects of herbivory and enrichment on macroalgal cover. Following nutrient enrichment, total macroalgae and the common seaweeds Dictyota spp. were suppressed in the presence, but not in the absence, of large herbivorous fishes—suggesting that fishes were selectively feeding on nutrient-enriched macrophytes. Access by large herbivores prevented algal overgrowth of corals, but these large fishes also directly grazed both corals. Excluding fishes did not alter survivorship of either coral species, but did decrease parrotfish grazing scars on both corals and increased the net growth of P. porites. Nutrient additions had no direct effects on the survivorship of corals, but there was a trend (P = 0.097) for nutrients to stimulate the growth of P. porites. The preponderance of experiments available to date indicates that loss of key herbivores is a major factor driving macroalgal blooms on coral reefs; anthropogenic nutrient pollution generally plays a more minor role.