From protégé to nurse plant: establishment of thorny shrubs in grazed temperate woodlands

Question: Thorny shrubs play keystone roles in grazed ecosystems by defending non‐protected plants against herbivores, but their establishment in grazed ecosystems is poorly understood. Which factors control establishment of recruits of thorny nurse shrubs in grazed temperate woodlands? Location: Ancient grazed temperate woodlands (52°32′N, 6°36′E), The Netherlands. Methods: We surveyed biotic and abiotic factors for saplings of thorny nurse shrubs in plots with and without saplings. To disentangle these factors, we performed a transplantation experiment over two growing seasons with nurse shrub saplings (Prunus spinosa and Crataegus monogyna) planted in two dominant vegetation types – tall unpalatable swards and short grazed lawns – half of them protected from herbivory via exclosures. Results: Plots with shrub saplings had taller surrounding vegetation, higher soil pH and higher soil moisture than plots without saplings. These plots predominantly contained unpalatable sward species, while plots without saplings mainly contained palatable lawn species. After transplantation, sapling survival was higher in exclosures than in the open, and higher in sward exclosures than in lawn exclosures. Sapling growth was higher in swards than in lawns, higher inside than outside exclosures, and higher for Prunus than Crataegus, while browsing on saplings was higher in lawns. Conclusion: Unpalatable swards form essential establishment niches for thorny shrubs in grazed temperate woodlands: they protect against herbivores before thorns fully develop in saplings, and sapling growth is better due to improved micro‐environmental conditions. Once established and thorny, shrub saplings grow out of the protective range of the swards and in turn facilitate tree seedlings, which are essential for long‐term persistence of grazed temperate woodlands. This study shows that nurse plants may start as protégés before becoming facilitators for other plants in a later life stage. This may be common for nurse plants in various ecosystems. We argue that improved understanding of establishment of nurse plants and their constraining factors is crucial for effective conservation and restoration in various ecosystems.     

Experimental evidence that deer browsing reduces habitat suitability for breeding Common Nightingales Luscinia megarhynchos

The ecological impacts of increasing populations of deer (Cervidae) in Europe and North America are becoming more widespread and pronounced. Within Britain, it has been suggested that declines in several woodland bird species, particularly those dependent on dense understorey vegetation, may be at least partly due to these effects. Here we present experimental evidence of the effects of deer browsing on the fine‐scale habitat selection and habitat use by a bird species in Europe. The study was conducted in a wood in eastern England where a decrease in Common Nightingale Luscinia megarhynchos numbers has coincided with a large increase in deer numbers. Eight woodland plots were cut to produce young coppice regrowth (a favoured habitat for Nightingales). Deer were excluded from half of each plot using steel fences, thus creating eight experimental pairs of exclosures (unbrowsed) and controls (browsed). Radiotelemetry and territory mapping of male Nightingales showed strong selection of exclosures. The density of territories was 15 times greater in the exclosures than in grazed controls. Selection for exclosures was significant for the minimum convex polygon, 95% kernel and 50% core home‐ranges used by seven radiotracked males. Tracked birds spent 69% of their time in the 6% of the study area protected from deer. Intensified browsing by deer influenced local settlement patterns of Nightingales, supporting the conclusion that increased deer populations are likely to have contributed to declines of Nightingales in Britain, and potentially those of other bird species dependent on dense understorey.     

Ecosystem changes associated with grazing in subhumid South American grasslands

Question: What are the changes in vegetation structure, soil attributes and mesofauna associated with grazing in mesic grasslands? Location: Southern Campos of the Río de la Plata grasslands, in south‐central Uruguay. Methods: We surveyed seven continuously grazed and ungrazed paired plots. Plant and litter cover were recorded on three 5‐m interception lines placed parallel to the fence in each plot. We extracted soil fauna from a 10 cm deep composite sample and analysed the oribatids. Soil attributes included bulk density, water content, organic carbon (in particulate and mineral associated organic matter) and nitrogen content and root biomass at different depths. Changes in floristic, Plant Functional Types and mesofauna composition were analysed by Non‐metric Multidimensional Scaling. Results: Species number was lower in ungrazed than in grazed plots. Of 105 species in grazed plots only three were exotics. Shrub and litter cover were significantly higher inside the exclosures, while the cover of Cyperaceae‐Juncaceae was lower. Grazing treatments differed significantly in plant and oribatid species composition. Grazing exclusion significantly reduced soil bulk density and increased soil water content. Carbon content in particulate organic matter was lower in the upper soil of ungrazed sites, but deeper in the profile, grazing exclosures had 8% more carbon in the mineral associated organic matter. Conclusions Our results generally agree with previous studies but deviate from the results of previous analyses in (1) the increase of shrub cover in ungrazed sites; (2) the redistribution of the soil organic carbon in the profile and (3) the low invasibility of the prairies regardless of grazing regime.     

The occurrence of the Broad‐toothed Rat Mastacomys fuscus in relation to feral Horse impacts

Feral Horse (Equus caballus) impacts in northern Kosciuszko National Park, New South Wales, Australia are directly occurring in habitat of the nationally threatened Broad‐toothed Rat (Mastacomys fuscus). This species is endemic primarily to the mountain regions of south‐eastern mainland Australia and Tasmania, with a disjunct population at Barrington Tops. The Broad‐toothed Rat's preferred habitat is being increasingly impacted by browsing and trampling associated with the expansion of feral horse populations. This study surveyed 180 sites supporting preferred habitat for this species to determine Broad‐toothed Rat presence and relative abundance in relation to the level of feral horse impacts within the reserve. There was a significant negative relationship between feral horse impacts and both Broad‐toothed Rat presence and abundance. No scats were identified at localities where feral horse impacts were severe, and at moderate horse impact sites, there was a proportion (34%) without scats found. Locations with low horse impacts had little impact on Broad‐toothed Rat occurrence. As feral horse populations increase, Broad‐toothed Rat populations may be further impacted. Such impacts will be due to the loss of vegetation cover from feral horse trampling and grazing, making animals more vulnerable to predation by predators or impacting on their ability to disperse to more suitable habitat. Habitat remnants and vegetation corridors along drainage lines require protection from feral horses to prevent localized extinctions of Broad‐toothed Rat.     

Does small-perimeter fencing inhibit mule deer or pronghorn use of water developments?

Wildlife water development can be an important habitat management strategy in western North America for many species, including both pronghorn (Antilocapra americana) and mule deer (Odocoileus hemionus). In many areas, water developments are fenced (often with small-perimeter fencing) to exclude domestic livestock and feral horses. Small-perimeter exclosures could limit wild ungulate use of fenced water sources, as exclosures present a barrier pronghorn and mule deer must negotiate to gain access to fenced drinking water. To evaluate the hypothesis that exclosures limit wild ungulate access to water sources, we compared use (photo counts) of fenced versus unfenced water sources for both pronghorn and mule deer between June and October 2002–2008 in western Utah. We used model selection to identify an adequate distribution and best approximating model. We selected a zero-inflated negative binomial distribution for both pronghorn and mule deer photo counts. Both pronghorn and mule deer photo counts were positively associated with sampling time and average daily maximum temperature in top models. A fence effect was present in top models for both pronghorn and mule deer, but mule deer response to small-perimeter fencing was much more pronounced than pronghorn response. For mule deer, we estimated that presence of a fence around water developments reduced photo counts by a factor of 0.25. We suggest eliminating fencing of water developments whenever possible or fencing a big enough area around water sources to avoid inhibiting mule deer. More generally, our results provide additional evidence that water development design and placement influence wildlife use. Failure to account for species-specific preferences will limit effectiveness of management actions and could compromise research results. © 2011 The Wildlife Society.     

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.

     

An assessment of feral horse impacts on treeless drainage lines in the Australian Alps

The feral Horse (Equus caballus) is widespread across the Australian Alps. Feral horses degrade alpine and sub‐alpine ecosystems and damage habitat of a range of threatened species. Despite this, there is little published work to document the extent and severity of these impacts. This study investigated impacts of feral horses on treeless drainage lines at 186 sites across the Australian Alps. The study included sites in the Australian Capital Territory, New South Wales and Victoria. We assessed nine variables related to soil and stream stability and vegetation cover, which in turn influence ecosystem function and habitat quality. We found significant differences among horse‐occupied and horse‐free sites for all soil and stream stability variables assessed. For all variables assessed, the average score (and hence, condition) was worse in horse‐occupied areas. The sites in poorest condition were occupied by horses. Impacts from other mammalian herbivores species appeared to be minor. Management intervention is necessary if these impacts of feral horses are to be addressed.     

Deer Herbivory as an Ecological Constraint to Restoration of Degraded Riparian Corridors

Ungulate herbivory can impact riparian vegetation in several ways, such as by reducing vigor or reproductive output of mature plants, and through increased mortality of seedlings and saplings. Much work has focused on the effects of livestock grazing within riparian corridors, while few studies have addressed the influence of native ungulate herbivory on riparian vegetation. This study investigated the effect of deer herbivory on riparian regeneration along three streams with degraded riparian corridors in Mendocino County, California. We utilized existing stream restoration efforts by private landowners and natural resource agencies to compare six deer exclosures with six upstream control plots. Livestock were excluded from both exclosure and control plots. Three of the deer exclosures had been in place for 15 years, one for 6 years, and two for 4 years. The abundance and size distribution of woody riparian plant species such as Salix exigua, S. laevigata, S. lasiolepis, Alnus rhombifolia, and Fraxinus latifolia were quantified for each exclosure and control plot. The mean density of saplings in deer exclosures was 0.49 ± 0.15/m², while the mean density of saplings in control plots was 0.05 ± 0.02/m². Within exclosures, 35% of saplings were less than 1 m and 65% were greater than 1 m; within control plots, 97% of saplings were less than 1 m in height. The fact that little regeneration had occurred in control plots suggests that deer herbivory can substantially reduce the rate of recovery of woody riparian species within degraded riparian corridors. Exclusionary fencing has demonstrated promising results for riparian restoration in a region with intense deer herbivory.     

Woodlands on farms in southern New South Wales: A longer-term assessment of vegetation changes after fencing

Summary Fencing incentive programmes have been widely used throughout Australia to assist landholders to fence remnant woodland vegetation, to control grazing and improve native vegetation condition. This study investigated vegetation and soil condition in remnant woodlands fenced for 7–9 years in the Murray catchment area in southern New South Wales. Surveys were undertaken at 42 sites, where vegetation condition was assessed in paired fenced and unfenced sites. Semi‐structured interviews were also conducted with landholders to gather management information. Woodlands surveyed were Yellow Box/Blakely's Red Gum (Eucalyptus melliodora/E. blakelyi, 15 sites), Grey Box (E. microcarpa, 13 sites) and White Cypress Pine (Callitris glaucophylla, 14 sites). Fencing resulted in a range of responses which were highly variable between sites and vegetation types. In general, fenced sites had greater tree regeneration, cover of native perennial grasses, less cover of exotic annual grasses and weeds, and less soil compaction than unfenced sites. However, there was greater tree recruitment in remnants to the west of the study area, and tree recruitment was positively correlated with time since fencing. Within sites, tree recruitment tended to occur in more open areas with a good cover of native perennial grasses, as compared to sites with a dense tree canopy, or dominated by exotic annuals grasses or weeds. Forty‐eight per cent of fenced sites had no tree regeneration. There was a significant decline in native perennial grasses, and increase of several unpalatable weeds in many fenced areas, suggesting certain ecological barriers may be preventing further recovery. However, drought conditions and associated grazing are the most likely cause of this trend. A range of grazing strategies was implemented in fenced sites which require further research as a conservation management tool. Continued long‐term monitoring is essential to detect key threats to endangered woodland remnants.     

Ungulates increase forest plant species richness to the benefit of non‐forest specialists

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant‐plant interactions and through soil disturbance. In forest ecosystems, researchers’ attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation‐wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light‐demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non‐forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape‐level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.     

Plant responses to fertilization and exclusion of grazers on an arctic tundra heath

This study investigated the impacts of fertilization and grazing by Norwegian lemmings (Lemmus lemmus), grey‐sided voles (Clethrionomys rufocanus), and reindeer (Rangifer tarandus) on a diverse tundra plant community dominated by deciduous shrubs. Four out of eight study areas, having a size of 2500 m² each, were fertilized with a N‐P‐K fertilizer and four areas served as unfertilized controls. Two types of exclosures were used within each study area, one to exclude solely reindeer, and one to exclude both rodents and reindeer. Open, grazed plots served as controls. During 5 years following the fertilization event the changes in vegetation inside and outside the exclosures were monitored using a point frequency method. The densities of rodents on the fertilized and unfertilized areas were investigated by live trapping and by counting nests of overwintering individuals. Reindeer do not graze on the study area during the growing season but migrate through this area in autumn and spring. Fertilization increased the abundance of vascular plants while grazing by reindeer and rodents decreased the abundance of vascular plants significantly on both fertilized and unfertilized areas. Rodents preferred clearly the fertilized areas during winter, decreasing the abundance of Vaccinium myrtillus and Vaccinium vitis‐idaea, while very little grazing occurred during summer. Graminoids showed the strongest positive response to fertilization and dominated the plant community on ungrazed plots, while winter grazing by both reindeer and rodents significantly decreased the abundance of graminoids. Deciduous shrubs (Betula nana, Vaccinium myrtillus) increased slightly but significantly due to fertilization and evergreen dwarf shrubs showed no response to fertilization. However, the use of functional growth forms for predicting the responses of nutrient enrichment and grazing must be questioned, as responses to fertilization as well as preferences by herbivores were shown to be species‐specific rather than uniform within functional groups based on plant growth forms.     

Cumulative effects of wild ungulate and livestock herbivory on riparian willows

We examined the effects of wild ungulates (deer and elk) and domestic sheep browsing on the growth, structure, and reproductive effort of two common willow species, Salix boothii and S. geyeriana, in a montane northeast Oregon riparian zone. With the use of exclosures, large herbivore effects on willows were studied in an area browsed by native mammals only and an adjacent area in which domestic sheep also lightly grazed during summer months. Growth variables were repeatedly measured on individual plants over a 5-year period to understand physiognomic and flowering responses of native willows to different levels of browsing pressure. At the beginning of the study, all willows were intensely browsed but were significantly taller in the area browsed only by native mammals than in the area also grazed by sheep (69 versus 51 cm, respectively). Willows inside exclosures responded with pronounced increases in height, crown area, and basal stem diameters while the stature of browsed plants outside exclosures stayed constant or declined. In the area browsed by both sheep and wild herbivores, the size of browsed plants remained at pre-treatment levels (<60 cm in height) for the duration of the study. There was no significant difference in growth rates of enclosed willows, indicating that current herbivory was the primary cause of growth retardation in the study area. Foliar area was strongly correlated with basal stem numbers for enclosed plants but much less so for browsed plants. Willows inside exclosures had more than twice as much foliar area per stem. Stem diameters were a positive function of crown area: stem-number ratios, suggesting lower photosynthetic potential was correlated with diminished radial growth among browsed plants. No flowering was observed until 2 years after exclusion when plants inside all exclosures and browsed willows in the wild ungulate area responded with a large pulse in flowering. Browsed plants in the sheep + wild ungulate area did not flower. The number of catkins produced per plant was significantly associated with willow height and plants <70 cm in height did not flower, thus suggesting a size threshold for reproduction in these species. Our results suggest that even relatively light levels of domestic livestock grazing, when coupled with intense wild ungulate browsing, can strongly affect plant structure and limit reproduction of riparian willows.     

Feral horse impacts on threatened plants and animals in sub‐alpine and montane environments in Victoria,Australia

Feral herbivores are a major driver of biodiversity loss globally and can alter the structure, composition and functioning of ecosystems. The direct impacts of feral herbivores on plant communities are well studied, but the direct and indirect effect they have on wildlife is not well understood. In Victoria (south‐eastern Australia), a large feral Horse (Equus caballus) population coincides with highly sensitive and nationally endangered Alpine Sphagnum Bogs and Associated Fens communities, and several threatened animal species. We assessed the impact of feral horses on this ecological community and the Alpine Water Skink (Eulamprus kosciuskoi) and the Broad‐toothed Rat (Mastacomys fuscus) at 20 sites with varying levels of horse disturbance. We used scat counts to determine an index of feral horse abundance and quantified impacts associated with their presence in the landscape. Active searches were used for Alpine Water Skink and scat and runway surveys for Broad‐toothed Rat. We also measured the vegetation structure and the abundance of different vegetation types (life forms). Our results suggest that feral horses are associated with vegetation types and characteristics that negatively influence the presence or abundance of Alpine Water Skink and Broad‐toothed Rat. Sites with high horse activity had more low‐growing forbs, and the abundance of Alpine Water Skink was negatively related to this vegetation type. Grasses, sedges, rushes and shrubs were also less dense and lower in height in high horse activity sites, and Broad‐toothed Rat was less likely to be present in areas with these habitat attributes. We recommend that feral horses are controlled to protect these threatened vertebrate species and their Sphagnum bog habitat.     

Quantifying ecological filters: the relative impact of herbivory, neighbours, and sediment on an oligohaline marsh

The objective of this experiment was to quantify the relative importance of herbivory, interactions with neighbours and sediment addition as filters in controlling the species present in an oligohaline marsh in southeastern Louisiana, USA. We planted 16 species of wetland plants in 3×3 m plots either inside or outside herbivore exclosures, with or without presence of established vegetation, and with or without added sediment. These species, representing 12 families and diverse morphologies, included herbaceous and woody plants. At the end of the growing season above‐ and below ground biomass were measured. In cleared plots (no neighbours) inside herbivore exclosures, 12 of the 16 species grew well, indicating that they could tolerate the physical conditions in the habitat. Competition significantly reduced the biomass of four of these species (A. calamus, C. occidentalis, P. hemitomon, P. cordata), suggesting that it is the strongest species‐specific filter operating in the marsh. These results confirm the general consensus that competition from existing plants is the strongest filter operating in high biomass habitats, and further imply that competition has the potential to remove 33% of the species from the community. Facilitation occurred for one species, R. corniculata. Herbivory had the largest general effect of reducing biomass, as determined in the full ANOVA model, but on a species‐specific basis, significantly reduced the biomass of only two species (T. distichum, T. domingensis). These results support the emerging view that herbivory is an important filter in coastal wetlands. Although several other studies have argued for the importance of sedimentation in controlling plant species composition in wetlands, we found no effect of sediment addition. Competition and herbivory appear to be two critical biological filters that control plant composition of coastal wetlands, and they must be considered in future studies and restoration efforts along the Gulf of Mexico.     

Livestock exclusion increases the spatial heterogeneity of vegetation in Colorado shortgrass steppe

Abstract. Spatial heterogeneity, an important characteristic in semi‐arid grassland vegetation, may be altered through grazing by large herbivores. We used Moran's I, a measure of autocorrelation, to test the effect of livestock grazing on the fine scale spatial heterogeneity of dominant plant species in the shortgrass steppe of northeastern Colorado. Autocorrelation in ungrazed plots was significantly higher than in grazed plots for the cover of the dominant species Bouteloua gracilis, litter cover and density of other bunchgrasses. No species had higher autocorrelation in grazed compared to ungrazed sites. B. gracilis cover was significantly auto‐correlated in seven of eight 60‐yr ungrazed exclosures, four of six 8‐yr exclosures, and only three of eight grazed sites. Autocorrelograms showed that B. gracilis cover in ungrazed sites was frequently and positively spatially correlated at lag distances less than 5 m. B. gracilis cover was rarely autocorrelated at any sampled lag distance in grazed sites. The greater spatial heterogeneity in ungrazed sites appeared linked to patches characterized by uniformly low cover of B. gracilis and high cover of C₃ grasses. This interpretation was supported by simple simulations that modified data from grazed sites by reducing the cover of B. gracilis in patches of ca. 8 m diameter and produced patterns quite similar to those observed in ungrazed sites. In the one exclosure where we intensively sampled soil texture, autocorrelation coefficients for sand content and B. gracilis cover were similar at lag distances up to 12 m. We suggest that the negative effect of sand content on B. gracilis generates spatial heterogeneity, but only in the absence of grazing. An additional source of heterogeneity in ungrazed sites may be the negative interaction between livestock exclusion and B. gracilis recovery following patchy disturbance.     

Effects of reindeer on boreal forest floor vegetation: Does grazing cause vegetation state transitions?

Intensive reindeer grazing has been hypothesized to drive vegetation shifts in the arctic tundra from a low-productive lichen dominated state to a more productive moss dominated state. Although the more productive state can potentially host more herbivores, it may still be less suitable as winter grazing grounds for reindeer, if lichens, the most preferred winter forage, are less abundant. Therefore, such a shift towards mosses may have severe consequences for reindeer husbandry if ground-growing lichens have difficulties to recover. We tested if reindeer cause this type of vegetation state shifts in boreal forest floor vegetation, by comparing plant species composition and major soil processes inside and outside of more than 40-year-old exclosures. Lichen biomass was more than twice as high inside exclosures than in grazed controls and almost 5 times higher than in heavily grazed patches. Contrary to our predictions, net N mineralization and plant production were higher in the exclosures than in the grazed controls. The lack of response of phytometer plants in a common garden bioassay indicated that changed soil moisture may drive effects of reindeer on plant productivity in these dry Pine forest ecosystems.     

Effect of host populations on the intensity of ticks and the prevalence of tick-borne pathogens: how to interpret the results of deer exclosure experiments

Deer are important blood hosts for feeding Ixodes ricinus ticks but they do not support transmission of many tick-borne pathogens, so acting as dead-end transmission hosts. Mathematical models show their role as tick amplifiers, but also suggest that they dilute pathogen transmission, thus reducing infection prevalence. Empirical evidence for this is conflicting: experimental plots with deer removal (i.e. deer exclosures) show that the effect depends on the size of the exclosure. Here we present simulations of dynamic models that take into account different tick stages, and several host species (e.g. rodents) that may move to and from deer exclosures; models were calibrated with respect to Ixodes ricinus ticks and tick-borne encephalitis (TBE) in Trentino (northern Italy). Results show that in small exclosures, the density of rodent-feeding ticks may be higher inside than outside, whereas in large exclosures, a reduction of such tick density may be reached. Similarly, TBE prevalence in rodents decreases in large exclosures and may be slightly higher in small exclosures than outside them. The density of infected questing nymphs inside small exclosures can be much higher, in our numerical example almost twice as large as that outside, leading to potential TBE infection risk hotspots.     

Relative impacts of cattle grazing and feral animals on an Australian arid zone reptile and small mammal assemblage

The effect of different levels of cattle grazing on an arid Australian small terrestrial mammal and lizard assemblage was assessed in a long‐tem series of cross‐fence comparisons. Cross‐fenced sites were closely matched for edaphic and vegetation characteristics and experienced near identical weather patterns, to ensure that cattle grazing pressure was the principal determinant of any differences in fauna assemblages. In addition, the effects of removal of cattle, cats, foxes and rabbits from three of these long‐term monitoring sites were assessed to determine the relative impacts of cattle grazing and feral animals. Small mammal captures, with the exception of Mus musculus, revealed a significant negative response to cattle grazing pressure but this response was of a considerably lower magnitude than the dramatic increase in rodent captures and species richness within the feral animal‐proof Arid Recovery Reserve. Higher kangaroo numbers in ungrazed controls, compared with treatments grazed by cattle, possibly negated the benefits to small mammals of removing cattle grazing. No reptile species responded significantly to the grazing treatments although reptile richness and captures of geckos and skinks were the lowest and agamid captures were the highest at heavily grazed sites. Nephrurus levis was the only reptile species to increase significantly, while captures of some smaller geckoes declined, within the feral‐proof treatment. Feral predation exerted a more significant effect on most small mammal species than the levels of cattle grazing assessed in this study, yet reptile responses to grazing or feral animals were less apparent and were likely primarily driven by changes in vegetation cover or secondary trophic impacts.     

Seed dispersal in red deer (Cervus elaphus L.) dung and its potential importance for vegetation dynamics in subalpine grasslands

Free-ranging large herbivores can influence vegetation dynamics through seed dispersal within and among habitats. We investigated the content of germinable seeds in the dung (endozoochory) of red deer (Cervus elaphus L.), the most ubiquitous wild ungulate throughout the European Alps, and compared the results with the species composition of the vegetation type in which the dung was dropped. The study was conducted in the subalpine zone of the Swiss National Park and included the three most important vegetation types for red deer: (i) intensively grazed short-grass vegetation, (ii) less intensively grazed tall-grass vegetation, and (iii) adjacent conifer forest understory vegetation. Seeds of 47 species, mostly from small-seeded herbaceous species, were recorded in dung samples with three species accounting for 65% of germinated seeds. Our results confirmed the hypotheses that (H1) small-seeded species were more likely to occur in red deer dung than larger-seeded species, though seed size was unrelated to seed density, (H2) red deer dung contained mostly seeds from short-grass vegetation, with seed species composition in dung collected from any vegetation type being most similar to species composition of relevés from short-grass vegetation, and (H3) seeds were less likely to be dispersed between vegetation types than within vegetation types, with dung dropped in short-grass vegetation having a different species composition and containing over twice as many seeds as dung dropped in the other two vegetation types. These results collectively support the hypothesis that red deer endozoochory contributes to maintaining short-grass vegetation, the favoured grazing sites of hinds in the Swiss National Park, by increasing propagule pressure of seeds from herbaceous forage species adapted to endozoochory relative to other species and especially those from later stages of secondary succession.     

Impacts of feral horses in the Australian Alps and evidence‐based solutions

New evidence of impacts by feral horses in Australia's alpine parks systems confirms they endanger threatened species and extensively damage critically endangered bog communities that could take millennia to recover. These impacts are not confounded by effects of deer and accumulate over time, even when only a small number of feral horses (~100) are present. With protected areas representing only a small proportion of the area of the Australian states of New South Wales (9.3%) and Victoria (17%), allowing feral horses to degrade reserves is not a reasonable management compromise, is contrary to the purpose of the protected area system and conflicts with international obligations. Modelling and decades of management experience indicate that trapping alone does not control feral horse numbers. Trapping and fertility control can work in small populations, but not when there are several thousand horses in remote areas. Aerial culling is needed to cost‐effectively and humanely control feral horse populations. The relatively small amount of suffering feral horses experience during a cull is outweighed by (i) avoiding suffering and death of horses from starvation and thirst, (ii) avoiding the suffering of native animals displaced by horses and (iii) avoiding the ethical concerns of driving threatened species towards extinction. Objections to aerial culling on welfare and cultural grounds are contradicted by evidence. Improving knowledge in the general community about what is at stake is long overdue because without this knowledge, small groups with vested interests and unfounded claims have been able to dominate debate and dictate management actions. As a result of ineffective management, horse populations are now expanding and causing well‐documented damage to Australia's alpine parks, placing at risk almost $10M spent on restoration after livestock grazing ended. The costs of horse control and restoration escalate the longer large horse populations remain in the alpine parks. It is crucial that feral horse numbers are rapidly reduced to levels where ecosystems begin to recover. Aerial culling is needed as part of the toolbox to achieve that reduction.