Ungulate browsers promote herbaceous layer diversity in logged temperate forests

Ungulates are leading drivers of plant communities worldwide, with impacts linked to animal density, disturbance and vegetation structure, and site productivity. Many ecosystems have more than one ungulate species; however, few studies have specifically examined the combined effects of two or more species on plant communities. We examined the extent to which two ungulate browsers (moose [Alces americanus]) and white‐tailed deer [Odocoileus virginianus]) have additive (compounding) or compensatory (opposing) effects on herbaceous layer composition and diversity, 5–6 years after timber harvest in Massachusetts, USA. We established three combinations of ungulates using two types of fenced exclosures – none (full exclosure), deer (partial exclosure), and deer + moose (control) in six replicated blocks. Species composition diverged among browser treatments, and changes were generally additive. Plant assemblages characteristic of closed canopy forests were less abundant and assemblages characteristic of open/disturbed habitats were more abundant in deer + moose plots compared with ungulate excluded areas. Browsing by deer + moose resulted in greater herbaceous species richness at the plot scale (169 m²) and greater woody species richness at the subplot scale (1 m²) than ungulate exclusion and deer alone. Browsing by deer + moose resulted in strong changes to the composition, structure, and diversity of forest herbaceous layers, relative to areas free of ungulates and areas browed by white‐tailed deer alone. Our results provide evidence that moderate browsing in forest openings can promote both herbaceous and woody plant diversity. These results are consistent with the classic grazing‐species richness curve, but have rarely been documented in forests.     

Habitat preferences of ungulates in closed savanna of central Africa

Habitat preferences of fifteen species of ungulate were studied between July and September 1975 in dry savanna woodland of Mwabvi Game Reserve, Southern Malawi, Central Africa. Thirteen vegetation types were recognized using vegetation transects and plotless sampling techniques (but three of these are omitted from further analysis here either because of the difficulty of sampling in a representative manner or because of their infrequency). A combination of six methods was used to study habitat preferences since each had particular advantages or disadvantages, although quadrat sampling of fresh pellets/spoor counts was the most satisfactory and so used predominantly. The most common and widely distributed species was the Greater kudu, although abundance estimates in different habitats varied according to the method used. Pellet/spoor counts showed heavy use of open grassland by kudu which was not detected by daytime drives. Temporal patterns of activity indicated that the species was moving into more closed woodland during the day and only coming out into open grassland at night. Relative abundance estimates of a species may also be affected by changing group size. The mean group size of kudu increased in more open vegetation, although group size in warthog did not show this relationship. Some species, e.g. impala, reedbuck, and klipspringer showed narrow habitat preferences, occurring in only one or a few vegetation types, whilst others, e.g. kudu, sable, duiker, suni, and grysbok showed much broader preferences. Mixed Colophospermum mopane /Acacia woodland had the highest species diversity but rather low abundance. Open grassland and Acacia/Albizia woodland had high species diversity together with the greatest abundance of ungulates. Open Julbernardia/Diplorynchus, Brachystegia/Julber-nardia woodland, and particularly Pterocarpus/Diplorynchus/Combretum hill woodland had the lowest species diversity and abundance of ungulates. Ready access to cover and availability of suitable food plants probably account for the preferences of certain vegetation types over others, whilst the different behavioural ecology and food preferences of each species account for individual species differences. Access to water probably also determines habitat preferences in particular species, and studies during the wet season may show different distributions.     

Rewilding traditional grazing areas affects scavenger assemblages and carcass consumption patterns

The abandonment of traditional livestock farming systems in Mediterranean countries is triggering a large-scale habitat transformation, which, in general, consists of the replacement of open grazing areas by woodlands through non-managed regeneration. As a consequence, wild ungulates are occupying rapidly the empty niche left by domestic ungulates. Both types of ungulates represent the main trophic resource for large vertebrate scavengers. However, a comparison of how vertebrate scavengers consume ungulate carcasses in different habitats with different ungulate species composition is lacking. This knowledge is essential to forecast the possible consequences of the current farmland abandonment on scavenger species. Here, we compared the scavenging patterns of 24 wild and 24 domestic ungulate carcasses in a mountainous region of southern Spain monitored through camera trapping. Our results show that carcasses of domestic ungulates, which concentrate in large numbers in open pasturelands, were detected and consumed earlier than those of wild ungulate carcasses, which frequently occur in much lower densities at more heterogenous habitats such as shrublands and forest. Richness and abundance of scavengers were also higher at domestic ungulate carcasses in open habitats. Vultures, mainly griffons (Gyps fulvus), consumed most of the carcasses, although mammalian facultative scavengers, mainly wild boar (Sus scrofa) and red fox (Vulpes vulpes), also contributed importantly to the consumption of wild ungulate carcasses in areas with higher vegetation cover. Our findings evidence that the abandonment of traditional grazing may entail consequences for the scavenger community, which should be considered by ecologists and wildlife managers.     

Influence of interspecific competition on mule deer birthing and rearing site selection

Ungulates often alter behavior and space use in response to interspecific competition. Despite observable changes in behavior caused by competitive interactions, research describing the effects of competition on survival or growth is lacking. We used spatial modeling to determine if habitat use by female mule deer (Odocoileus hemionus) was affected by other ungulate species prior to, during, and after parturition. We conducted our study in the Book Cliffs region of eastern Utah, USA, during 2019 and 2020. We used resource selection function (RSF) analysis to model space use of 4 ungulate species that potentially competed with mule deer: bison (Bos bison), cattle, elk (Cervus canadensis), and feral horses. We incorporated RSF models for competing species into a random forest analysis to determine if space use by mule deer was influenced by these other ungulate species. We used survival and growth data from neonate mule deer to directly assess potential negative effects of other ungulates. Habitat use by elk was an important variable in predicting use locations of mule deer during birthing and rearing. The relationship was positive, suggesting interference competition was not occurring. Survival of neonate mule deer increased as the probability of use by elk increased (hazard ratio = 0.185 ± 0.497 [SE]). Further, probability of use by elk in rearing habitat had no influence on growth of neonate mule deer from birth to 6 months of age, suggesting that exploitative competition was not occurring.     

Resource selection,utilization and seasons influence spatial distribution of ungulates in the western Serengeti National Park

Understanding herbivore selection and utilization of vegetation types is fundamental to conservation of multispecies communities. We tested three hypotheses for how ungulate species select their habitats and how this changes with season: first, resources are distributed as a mosaic of patches so that ungulates are also distributed patchily; this distribution reflects habitat selection, which changes with season, the different ungulates behaving differently. Second, resources become scarcer in the dry season relative to those in the wet season. If interspecific competition prevails, then all species should show a contraction of habitats chosen. Third, if predation is limiting, competition will be minimal, and hence, habitat selection by herbivores will not differ between seasons. We used frequencies of occurrence in four common vegetation types in western Serengeti National Park to determine selection coefficients and utilization patterns and Chi‐square analysis to test the hypotheses. The results showed that selection changes differently in each species, agreeing with the first hypothesis. Herbivores did not all become more selective, as predicted by the competition hypothesis, nor did selection remain the same across seasons, as predicted by the predation hypothesis. These results can be useful in constructing habitat suitability maps for ungulate species with special conservation needs.     

The feeding habits of wolves in relation to large prey availability in northern Italy

We investigated wolf feeding habits in relation to the abundance of wild and domestic ungulates to test the hypothesis that large prey are preferred and that their abundance affects the use of other food categories and diet breadth. We determined diet composition by scat analysis from December 1987 to December 1992. The research was carried out in three study areas located in northern Italy and characterised by marked differences in wild and domestic ungulate abundance. In study area A (low wild and domestic ungulate availability) fruits, livestock, other vertebrates and wild ungulates made up the bulk of the diet (71% in volume). In area B (high availability of livestock) wolf diet was mainly based on sheep and wild boars (80% in volume). In study area C (high availability of wild ungulates) wild ungulates were the main food of wolves (90% in volume). Significant differences were found among study areas in the mean percentage volume of all food categories and in particular for wild ungulates, livestock, other vertebrates and fruits (p < 0.0001 in all cases). Diet breadth decreased in areas with high availability of large wild and domestic herbivores. The use of livestock species was lower where there was high abundance, richness and diversity of the wild ungulate guild. Selection for wild ungulate species was partially affected by their abundance: however other factors as prey social behaviour, adaptability to the habitat (for introduced species), and body size could have an important role in species selection by wolves. In particular in area C wild boars were selected for, roe and red deers avoided, and fallow deers and mouflons used as available. Livestock species were used in relation to their abundance and accessibility, in particular sheep were selected for and cattle avoided; but if calves bom in the pastures were considered as the only available cattle, they were selected for and sheep were used as available. Large and in particular wild herbivores were found to be of great importance for the wolf population maintenance in northern Italy, one of the most important recovery areas of Mediterranean wolves.     

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.     

Habitat Preferences of Ungulates in Hunted and Nonhunted Areas in the Calakmul Forest, Campeche, Mexico1

We investigated habitat preference in the community of ungulates in the Calakmul Forest of Southern Mexico through systematically counting the tracks of six species in randomly placed transects. Tracks were associated with one of four major forest types according to Pennington and Sarukhan (1998) . Forest type availability was estimated through 393 independent points on the same transects. We surveyed 90 transects in three hunted areas and one large nonhunted area, with a total of 206 km walked. A total of 1672 tracks of the six species were found. Brocket deer (Mazama americana and Mazama pandora) preferred low‐dry forest in the nonhunted area and low‐flooded forest in the hunted areas. Collared peccary (Pecari tajacu) was a generalist species in the nonhunted area, whereas in the hunted areas, it preferred the subperennial forest. White‐lipped peccary (Tayassu pecari) was found only in subperennial forest in the nonhunted area and favored low‐flooded forest in the hunted areas. White‐tailed deer (Odocoileus virginianus) preferred low‐flooded forest in the hunted areas, while it was a generalist in the nonhunted area. Tapir (Tapirus bairdii) preferred low‐flooded forest in the hunted areas. The most evident habitat difference among hunted and nonhunted areas was a major use of low‐flooded forest in the hunted areas for the species. Conservation of ungulate species in the Calakmul region requires protection of all major habitat types in hunted and nonhunted areas.     

Climate and Species Richness Predict the Phylogenetic Structure of African Mammal Communities

We have little knowledge of how climatic variation (and by proxy, habitat variation) influences the phylogenetic structure of tropical communities. Here, we quantified the phylogenetic structure of mammal communities in Africa to investigate how community structure varies with respect to climate and species richness variation across the continent. In addition, we investigated how phylogenetic patterns vary across carnivores, primates, and ungulates. We predicted that climate would differentially affect the structure of communities from different clades due to between-clade biological variation. We examined 203 communities using two metrics, the net relatedness (NRI) and nearest taxon (NTI) indices. We used simultaneous autoregressive models to predict community phylogenetic structure from climate variables and species richness. We found that most individual communities exhibited a phylogenetic structure consistent with a null model, but both climate and species richness significantly predicted variation in community phylogenetic metrics. Using NTI, species rich communities were composed of more distantly related taxa for all mammal communities, as well as for communities of carnivorans or ungulates. Temperature seasonality predicted the phylogenetic structure of mammal, carnivoran, and ungulate communities, and annual rainfall predicted primate community structure. Additional climate variables related to temperature and rainfall also predicted the phylogenetic structure of ungulate communities. We suggest that both past interspecific competition and habitat filtering have shaped variation in tropical mammal communities. The significant effect of climatic factors on community structure has important implications for the diversity of mammal communities given current models of future climate change.     

Lack of feral livestock interference with native guanaco during the dry season in a South American desert

Analyzing coexistence of exotic and native ungulates in arid areas is important from both a theoretical and a species conservation perspective. We assess the habitat use patterns and possible interference between guanaco (Lama guanicoe) and feral livestock (donkey and cattle) in arid environments of South America. To determine habitat use and niche overlap between exotic and native ungulate species, ten sites with different habitats and six natural waterholes were selected. Plots (20 at each site, ten around each waterhole) were randomly set up and characterized by environmental variables and relative use by cattle, donkey and guanaco through faecal pellet counts. Aggregation, niche breadth and niche overlap of the three herbivores were analyzed at habitat level (mesoscale). A direct redundancy analysis was used to examine the relationships between abundance of herbivore faeces and environmental variables at microscale. Mesoscale analyses showed (i) an extensive use of the area by all three species, with guanaco having the highest niche breadth followed by donkey and cattle and (ii) a large, broad guanaco–donkey and donkey–cattle habitat overlap. However, results at a finer scale showed high spatial aggregation of feral livestock species and an independent use of territory by guanacos. This study is the first to provide information about habitat partitioning between guanacos and feral livestock in the hyper-arid Monte Desert biome and points to an apparent lack of negative effects on the native ungulate.     

What Cues Do Ungulates Use to Assess Predation Risk in Dense Temperate Forests?

Anti-predator responses by ungulates can be based on habitat features or on the near-imminent threat of predators. In dense forest, cues that ungulates use to assess predation risk likely differ from half-open landscapes, as scent relative to sight is predicted to be more important. We studied, in the Białowieża Primeval Forest (Poland), whether perceived predation risk in red deer (Cervus elaphus) and wild boar (Sus scrofa) is related to habitat visibility or olfactory cues of a predator. We used camera traps in two different set-ups to record undisturbed ungulate behavior and fresh wolf (Canis lupus) scats as olfactory cue. Habitat visibility at fixed locations in deciduous old growth forest affected neither vigilance levels nor visitation rate and cumulative visitation time of both ungulate species. However, red deer showed a more than two-fold increase of vigilance level from 22% of the time present on control plots to 46% on experimental plots containing one wolf scat. Higher vigilance came at the expense of time spent foraging, which decreased from 32% to 12% while exposed to the wolf scat. These behavioral changes were most pronounced during the first week of the experiment but continuous monitoring of the plots suggested that they might last for several weeks. Wild boar did not show behavioral responses indicating higher perceived predation risk. Visitation rate and cumulative visitation time were not affected by the presence of a wolf scat in both ungulate species. The current study showed that perceived predation risk in red deer and wild boar is not related to habitat visibility in a dense forest ecosystem. However, olfactory cues of wolves affected foraging behavior of their preferred prey species red deer. We showed that odor of wolves in an ecologically equivalent dose is sufficient to create fine-scale risk factors for red deer.     

Effects of herbivore species richness on the niche dynamics and distribution of blue sheep in the Trans-Himalaya

Aim To understand the community structure of mountain ungulates by exploring their niche dynamics in response to sympatric species richness. Location Ladakh and Spiti Regions of the Western Indian Trans‐Himalaya. Methods We used the blue sheep Pseudois nayaur, a relatively widely distributed mountain ungulate, as a model species to address the issue. We selected three discrete valleys in three protected areas with similar environmental features but varying wild ungulate species richness, and studied blue sheep’s diet and habitat utilization in them. Habitat variables such as slope angle, distance to cliff and elevation at blue sheep locations were recorded to determine the habitat width of the species. Faecal pellets were collected and microhistological faecal analysis was carried out to determine the diet width of blue sheep in the three areas with different ungulate species richness. Blue sheep’s niche width in terms of habitat and diet was determined using the Shannon’s Index. Results The habitat width of blue sheep had a negative relationship with the number of sympatric species. However, contrary to our expectation, there was a hump‐shaped relationship between blue sheep’s diet width and the sympatric species richness, with the diet width being narrower in areas of allopatry as well as in areas with high herbivore species richness, and the greatest in areas with moderate species richness. Main conclusions We suspect that the narrow diet width in allopatry is out of choice, whereas it is out of necessity in areas with high herbivore species richness because of resource partitioning that enables coexistence. We suggest that interactions with sympatric species lead to niche adjustment of mountain ungulates, implying that competition may play a role in structuring Trans‐Himalayan mountain ungulate assemblages. Given these results, we underscore the importance of including biotic interactions in species distribution models, which have often been neglected.     

Factors Related to the Recovery of Subalpine Woodland on Mauna Kea,Hawaii1

We measured mature tree and sapling density, tree associations, crown size, age structure, recovery from ungulate browsing, and grass cover at four study sites in two types of subalpine woodland on Mauna Kea volcano, island of Hawaii. Beginning in 1981, introduced ungulates were reduced in number to allow regeneration of Sophora chrysophylla (mamane) in habitat supporting the endangered Hawaiian finch, Loxioides bailleui (palila). We found Sophora regeneration at all four study sites, but regeneration was higher in mixed species woodland with codominant Myoporum sandwicense (naio) than in areas where Sophora dominated. Regeneration of Myoporum was uniformly very low in comparison. Invasive grass cover, which suppresses Sophora germination, was highest in mid-elevation woodland where Sophora dominated. The distribution of mature and sapling Sophora were both related to study site, reflecting previous ungulate browsing and uneven recovery due to grasses. Densities of Sophora snags were not different among any of the sites, suggesting a more even distribution in the past. Selective browsing before ungulate reduction may have favored Myoporum over Sophora, leading to high densities of mature Myoporum in codominant woodland. After ungulate reduction, however, we found no pattern of competitive inhibition by Myoporum on regeneration of Sophora. Reduction of Myoporum is not likely to enhance habitat for Loxioides as much as supplemental plantings of Sophora, glass control, and continued ungulate eradication. Mid-elevation Sophora woodland areas, where Loxioides forage and nest in high densities, would benefit the most from these management actions.     

Evidence for top predator control of a grazing ecosystem

The importance of top predators in controlling ecological processes in large, intact ecosystems is unclear. In grasslands that support abundant ungulates, top–down control by predators may be particularly important, because of the tight biogeochemical linkages of ungulate prey with plants and soil microbes. Here, I examined the effects of the recent reintroduction of the gray wolf Canis lupus on ecosystem processes in Yellowstone National Park, where herds of grazing ungulates previously have been shown to stimulate several processes, including soil net nitrogen (N) mineralization. Rates of ungulate grazing intensity and soil net N mineralization were compared before and after wolf reintroduction in grasslands ranging five‐fold in aboveground production. Grazing intensity and grassland net N mineralization declined after wolf reintroduction, a likely partial function of fewer ungulates; wolf predation has been one of several factors implicated in causing the decline in Yellowstone ungulates. In addition, the spatial pattern of grazing and net N mineralization changed after reintroduction. A shift in the spatial patterns of grazer‐associated processes is consistent with a growing body of work indicating that wolves have changed habitat use patterns of ungulates in Yellowstone National Park. These findings suggest widespread wolf effects on ungulate prey, plants, and microbial activity that have spatially reorganized grassland energy and nutrient dynamics in Yellowstone Park.     

Rodents change acorn dispersal behaviour in response to ungulate presence

Small rodents are prominent seed predators, but they also favour plant recruitment as seed dispersers. The direct interactions of ungulates on plants are more one‐sided and negative, as they mainly reduce plant recruitment through predation on seeds and seedlings. The effects of small rodents and ungulates on plant recruitment have been considered and studied as independent episodes within plant regeneration cycles. However, ungulate–rodent interactions and their potential effects on plant regeneration have not been considered so far. A number of studies have recently documented ungulate effects on the abundance, diversity and spatial distribution of small rodents. Here, we hypothesize that ungulates may also affect rodent seed dispersal behaviour. We monitored acorn dispersal by small rodents (Mus spretus and Apodemus sylvaticus) in oak woodlands with and without exclosures for large ungulates, mainly red deer, Cervus elaphus, and wild boar, Sus scrofa. The study was carried out in a typical Mediterranean Holm oak, Quercus ilex, forest throughout the acorn fall season in 2003 and 2004. We found that, in both years, the proportion of acorns cached and not recovered in the short‐term was, on average, lower in the presence (1.4%) than in the absence (19.9%) of ungulates. Acorn dispersal distances were not affected by ungulate presence in either year. However, ungulates had an effect on the spatial distribution of dispersed seeds; rodents apparently avoided shrubs as caching sites in both years. This result was interpreted as a behavioural response to reduce the risk of cache pilferage by conspecifics, which are closely associated with shrubs in presence, but not in absence, of ungulates. Potential effects of different densities of rodents or predators were discarded, as none of them differed between the areas with and without ungulates. The present study found significant interactions between heterospecific seed and seedling consumers that had been considered as independent episodes within tree regeneration cycles. As a result of such interactions, ungulates may have negative indirect effects on oak recruitment by reducing (1) acorn caching frequency, and (2) the proportion of acorns cached under shrubs, key nurse‐plants for the establishment of Holm oak seedlings in Mediterranean areas.     

Ungulates,rodents, shrubs: interactions in a diverse Mediterranean ecosystem

Ungulate abundance has increased dramatically worldwide, having strong impacts on ecosystem functioning. High ungulate densities can reduce the abundance, diversity and/or body condition of small mammals, which has been attributed to reductions in cover shelter and food availability by ungulates. The densities of wild ungulates have increased recently in high-diversity Mediterranean oak ecosystems, where acorn-dispersing small rodents are keystone species. We analysed experimentally ungulate effects on seed-dispersing rodents in two types of oak woodland: a forest with dense shrub layer and in dehesas lacking shrubs. Ungulates had no significant effects on vegetation structure or rodent body mass, but they reduced dramatically rodent abundance in the lacking-shrub dehesas. In the forest, ungulates modified the spatial distribution and space use of rodents, which were more concentrated under shrubs in the presence than in the absence of ungulates. Our results point to the importance of shrubs in mediating ungulate–rodent interactions in Mediterranean areas, suggesting that shrubs serve as shelter for rodents against ungulate physical disturbances such as soil compaction, trampling or rooting. Holm oak seedling density was reduced by ungulates in dehesa plots, but not in forests. Acorn consumption by ungulates may reduce oak recruitment to a great extent. Additionally, we suggest that ungulates may have a negative effect on oak regeneration processes by reducing the abundance of acorn-dispersing rodents. Given that shrubs seem to mediate ungulate effects on acorn dispersers, controlled shrub encroachment could be an effective alternative to ungulate population control or ungulate exclusion for the sustainability of the high-diversity Mediterranean oak ecosystems.     

Seed dispersal by ungulates as an ecological filter: a trait‐based meta‐analysis

Plant communities are often dispersal‐limited and zoochory can be an efficient mechanism for plants to colonize new patches of potentially suitable habitat. We predicted that seed dispersal by ungulates acts as an ecological filter – which differentially affects individuals according to their characteristics and shapes species assemblages – and that the filter varies according to the dispersal mechanism (endozoochory, fur‐epizoochory and hoof‐epizoochory). We conducted two‐step individual participant data meta‐analyses of 52 studies on plant dispersal by ungulates in fragmented landscapes, comparing eight plant traits and two habitat indicators between dispersed and non‐dispersed plants. We found that ungulates dispersed at least 44% of the available plant species. Moreover, some plant traits and habitat indicators increased the likelihood for plant of being dispersed. Persistent or nitrophilous plant species from open habitats or bearing dry or elongated diaspores were more likely to be dispersed by ungulates, whatever the dispersal mechanism. In addition, endozoochory was more likely for diaspores bearing elongated appendages whereas epizoochory was more likely for diaspores released relatively high in vegetation. Hoof‐epizoochory was more likely for light diaspores without hooked appendages. Fur‐epizoochory was more likely for diaspores with appendages, particularly elongated or hooked ones. We thus observed a gradient of filtering effect among the three dispersal mechanisms. Endozoochory had an effect of rather weak intensity (impacting six plant characteristics with variations between ungulate‐dispersed and non‐dispersed plant species mostly below 25%), whereas hoof‐epizoochory had a stronger effect (eight characteristics included five ones with above 75% variation), and fur‐epizoochory an even stronger one (nine characteristics included six ones with above 75% variation). Our results demonstrate that seed dispersal by ungulates is an ecological filter whose intensity varies according to the dispersal mechanism considered. Ungulates can thus play a key role in plant community dynamics and have implications for plant spatial distribution patterns at multiple scales. Synthesis Plant communities are often dispersal‐limited and zoochory can be an efficient mechanism for plants to colonize new patches of potentially suitable habitat. Our analysis is the first synthesis of ungulate seed dispersal that compares characteristics from both non‐dispersed and dispersed diaspores, distinguishing the three zoochory mechanisms ungulates are involved in: endozoochory, hoof‐epizoochory and fur‐epizoochory. We confirmed that seed dispersal by ungulates is an ecological filter whose intensity increases from endozoochory, then hoof‐epizoochory to finally fur‐epizoochory. By filtering seed traits through dispersal, ungulates can thus play a key role in plant community dynamics and have implications for plant spatial distribution patterns at multiple scales.     

Migratory Disturbance Thresholds with Mule Deer and Energy Development

Fine-scale movement data has transformed our knowledge of ungulate migration ecology and now provides accurate, spatially explicit maps of migratory routes that can inform planning and management at local, state, and federal levels. Among the most challenging land use planning issues has been developing energy resources on public lands that overlap with important ungulate habitat, including the migratory routes of mule deer (Odocoileus hemionus). We generally know that less development is better for minimizing negative effects and maintaining habitat function, but we lack information on the amount of disturbance that animals can tolerate before reducing use of or abandoning migratory habitat. We used global positioning system data from 56 deer across 15 years to evaluate how surface disturbance from natural gas well pads and access roads in western Wyoming, USA, affected habitat selection of mule deer during migration and whether any disturbance threshold(s) existed beyond which use of migratory habitat declined. We used resource and step selection functions to examine disturbance thresholds at 3 different spatial scales. Overall, migratory use by mule deer declined as surface disturbance increased. Based on the weight of evidence from our 3 independent but complementary metrics, declines in migratory use related to surface disturbance were non-linear, where migratory use sharply declined when surface disturbance from energy development exceeded 3%. Disturbance thresholds may vary across regions, species, or migratory habitats (e.g., stopover sites). Such information can help with management and land use decisions related to mineral leasing and energy development that overlap with the migratory routes of ungulates. © 2020 The Wildlife Society.