Ungulate Herbivory Research Articles

Long-term surveys of ungulates' effects on tree and shrub species in mountainous forests -outcomes and potential limits.

Ungulate herbivory might induce different effects on the diversity and growth of trees and shrubs. The density, distribution, and the species of ungulates as well as plant communities' composition and other factors determine whether ungulate herbivory promotes or limits plants' diversity and growth. The impacts of ungulates on woody plants are commonly surveyed with exclosure-control approaches. In practice, such surveys frequently only cover short periods of time, addressing immediate management needs. Long-term surveys, documenting lasting effects of ungulate herbivory, are highly needed, but still rare. However, the general transferability of outcomes of long-term surveys might be limited due to different disturbing factors. This study addresses two basic aspects of long-term monitoring in mountainous forests, based on a unique 30-year data set: (1) Possible long-term effects of herbivores on forest vegetation (e.g., species/structural diversity of woody plants) and (2) potential differences between short-term and long-term surveys in terms of height growth patterns. In our study, diversity of woody plant species showed great variability with no significant impact of ungulate herbivory. The presence of ungulates had a significantly negative effect on the vertical structural diversity and growth of trees. Due to the slower growth on control plots, it took trees longer to reach a 160 cm height-threshold with their terminal shoots than on exclosure plots. Our long-term control-exclosure data set indicated that long-term survey data indeed might differ from growth patterns represented by short-term surveys. This can be induced by several factors, like site-specific growth patterns of trees, occurrences of natural abiotic disturbances that influence the functional life of exclosures, and others.

Complex interactions of deer herbivory, soil chemistry, and competing vegetation explain oak–hickory forest tree regeneration in central Pennsylvania, USA

The root causes of forest tree regeneration failure are difficult to resolve, although numerous studies show ungulate herbivory, soil conditions, and competition from undesirable vegetation as likely contributors. To better understand the relative importance of each issue, we conducted a 7-year manipulative experiment to assess the interactive effects of white-tailed deer ( Odocoileus virginianus) herbivory, soil acidity, and competing vegetation on tree regeneration in oak–hickory forests of central Pennsylvania, USA. Outcomes depended on initial tree seedling abundance, and all three factors had significant interactions. At low initial seedling abundance, fencing resulted in the greatest increase, but all treatments had a positive effect on seedling growth and abundance. At higher initial seedling abundance, abundance failed to recover 7 years after herbicide treatment and soil pH was an important predictor. When soil pH was >4.6 from lime application, seedling growth and abundance in unfenced controls with high initial abundance was comparable to the fenced-only treatment. Competing vegetation, assumed to be a symptom of excessive, long-term deer herbivory, does not seem to be the primary factor limiting tree regeneration in our study area. Ameliorating acid deposition warrants greater consideration as a management action because it could provide long-lasting benefits compared to short-term fence installations.

Responses of oak seedlings to increased herbivory and drought: a possible trade-off?

Anthropogenic disturbances are causing a co-occurring increase in biotic (ungulate herbivory) and abiotic (drought) stressors, threatening plant reproduction in oak-dominated ecosystems. However, we wonder whether herbivory could compensate for the adverse impact of drought by reducing evapotranspiration. Thus, we investigate the isolated and joint effects of herbivory and drought on oak seedlings of two contrasting Mediterranean species that differ in leaf habit and drought resistance. California oak seedlings from the evergreen, and more drought-resistant, Quercus agrifolia and the deciduous Q. lobata (n=387) were assigned to a fully crossed factorial design with herbivory and drought as stress factors. Seedlings were assigned in a greenhouse to 3-4 clipping levels simulating herbivory and 3-4 watering levels, depending on the species. We measured survival, growth, and leaf attributes (chlorophyll, secondary metabolites, leaf area and weight) once a month (May-Sep) and harvested above- and below-ground biomass at the end of the growing season. For both oak species, simulated herbivory enhanced seedling survival during severe drought or delayed its adverse effects, probably due to reduced transpiration resulting from herbivory-induced leaf area reduction and compensatory root growth. Seedlings from the deciduous, and less drought-resistant species, benefitted from herbivory at lower levels of water stress, suggesting different response across species. We also found complex interactions between herbivory and drought on their impact on leaf attributes. In contrast to chlorophyll content which was not affected by herbivory, anthocyanins increased with herbivory - although water stress reduced differences in anthocyanins due to herbivory. Herbivory seems to facilitate Mediterranean oak seedlings to withstand summer drought, potentially alleviating a key bottleneck in the oak recruitment process. Our study highlights the need to consider ontogenetic stages and species-specific traits in understanding complex relationships between herbivory and drought stressors for the persistence and restoration of multi-species oak savannas.

Acorn removal and seedling age determine oak (Quercus ilex L. and Q. suber L.) restoration outcome in ungulate-dominated Mediterranean environments

Key messageWe test the potential benefits of planting 2-year vs. 1-year-old seedlings to restore Mediterranean oak-dominated systems. Planting 2-year-old Quercus suber L. seedlings is recommended for improved survival and resilience against wild boar (Sus scrofa L.) and drought in dry sandy soils. The removal of acorns in seedlings did not apparently influence leaf biochemical traits and could reduce wild boar damage, particularly in 1-year-old seedlings.ContextIn the face of anthropogenic global change, Mediterranean oak-dominated ecosystems confront increased biotic (ungulate herbivory) and abiotic (drought) stressors, compromising forest regeneration. Restoration measures are imperative to address this scenario.AimsThis study assesses the impact of different mitigation measures on the survival and biochemical traits of two oak species.MethodsWe planted Quercus ilex L. and Q. suber L. seedlings in Cabañeros and Doñana National Parks (Spain), subjecting them to three treatments: cotyledon/acorn removal, seedling age (1- vs. 2-year-old), and herbivore protection (fenced vs. non-fenced).ResultsWild boar (Sus scrofa L.) damage peaked in winter and early spring, while drought prevailed from late spring to fall. In sandy soils, wild boar uprooted 1-year-old more often than 2-years-old seedlings (40% vs. 18%). One-year-old seedlings without acorns showed higher survival rates against wild boar only in sandy soils. The removal of acorns in seedlings did not influence plant biochemical traits.ConclusionsPlanting 2-year-old seedlings in sandy soils may mitigate wild boar damage and improve drought resilience. Seedling age seems more important than acorn removal against biotic and abiotic stressors although acorn removal could reduce wild boar damage in 1-year-old seedlings. Our results underscore the importance of considering multiple stressors in oak restoration strategies.

Ungulate herbivores promote beta diversity and drive stochastic plant community assembly by selective defoliation and trampling: From a four‐year simulation experiment

Abstract Ungulate herbivores shape grassland plant communities at multiple scales, ultimately affecting ecosystem function. However, ungulates have complex effects on grasslands, including defoliation, trampling, excreta return and their interactions. Moreover, the effects of ungulate density on grasslands are regulated by these three mechanisms. Nevertheless, how these three mechanisms affect biodiversity at multiple scales and community assembly remains poorly understood. Here, we conducted a 4‐year novel field experiment to disentangle the effects of defoliation, trampling, and excreta return by ungulates on plant community assembly in a temperate grassland in Inner Mongolia, China. This experiment set two different scenarios: moderate ungulate density (Moderate, characterised by selective defoliation and moderate trampling) and high ungulate density (Intense, characterised by non‐selective defoliation and heavy trampling), including different combinations of defoliation, trampling and excreta return in each scenario. We found that defoliation and trampling increased stochasticity in community assembly and promoted alpha and beta diversity under both scenarios. Specifically, defoliation promoted the coexistence of species with multiple resource acquisition strategies (higher functional trait diversity) by reducing interspecific competition; trampling tended to facilitate random species colonisation. Conversely, excreta return favoured grasses, promoting deterministic assembly and impacting species coexistence. Notably, selective defoliation in the Moderate scenario led to a dominance of stochastic processes during community assembly, whereas non‐selective defoliation still did not change the dominance of deterministic processes. Further, communities subject to selective defoliation were insensitive to changes in soil properties caused by trampling and excreta return, maintaining a high‐level beta diversity and the stochastic of community assembly. Synthesis: Our study provides important insights into the mechanisms by which ungulate herbivores influence plant community assembly, suggesting that defoliation and trampling have the potential to drive stochastic processes, while excreta return plays the opposite role. Our study also suggests that selective foraging by ungulates acts as stronger stochastic forces during community assembly compared to non‐selective defoliation. These results imply that considering ungulate feeding preferences and foraging behaviour in grassland management will help prevent biodiversity loss and biotic homogenisation.

Influence of Soil Quality on the Browsing Intensity of Ungulate Herbivores on Tree Species in European Forests

Influence of Soil Quality on the Browsing Intensity of Ungulate Herbivores on Tree Species in European Forests

Ungulate herbivores promote contrasting modifications of soil properties and organic carbon stabilization in a grazed grassland versus rewilded woodland environment

Large vertebrate herbivores are ecosystem engineers, provoking cascading effects on soil properties through modifications of plant communities as well as direct modulation of organic matter incorporated into soils. However, most studies have been carried out with domestic ungulates in grasslands, and the effects which predominate in grasslands may not be generalizable to other ecosystems. It is not well understood how trophic rewilding with wild ungulates may affect soil processes, especially in non-grassland systems albeit rewilding of natural areas is a process associated with larger trends of land abandonment, shrub encroachment or land use change. In this study we investigated ungulate herbivore exclusion in two geographically and pedo-climatically equivalent but ecologically distinct contexts: a Mediterranean grassland with domesticated herbivores, and a reforested area populated by a community of wild ungulates. In each context, long-term herbivore exclusion zones were used to understand the effect of herbivores on soil properties and particularly on soil organic carbon (SOC). Samples from the topsoil (0–5 cm) were taken inside and outside the exclosures distributed between the two contexts, and ecosystem-level effects and responses to grazing were investigated. We found that soil properties’ response to grazing depended on the ecological context in which the grazing occurred. SOC contents in the grassland increased by up to 37% with grazing, while grazing reduced SOC concentrations by 23% in the rewilded area. In both cases, grazing was also associated with significant alterations of soil microbial biomass, the microbial metabolic quotient (qCO2), and aggregate stability, but the effects were opposite: On one hand, domesticated herbivores in the grassland positively influenced carbon stabilization which was strongly associated with soil microbial processes and characteristics, and additionally the proportion of stable macroaggregates was increased. In contrast, wild ungulate herbivory in the forest ecosystem reduced SOC concentrations, accompanied by decreased microbial biomass and increased microbial maintenance costs, and stable macroaggregates were reduced. The results indicate the importance of ecological context in determining the effect of herbivores on soil properties and processes and suggest that the directionality of effects on SOC may change owing to the predominance of different mechanisms of SOC stabilization. Consequently, this study provides evidence that the management of ungulates in rangelands or forests can be associated with different outcomes as related to crucial soil properties such as SOC contents and aggregation.

How Effective Is Environmental Protection for Ensuring the Vitality of Wild Orchid Species? A Case Study of a Protected Area in Italy.

Orchids are among the plants most threatened by anthropic impact and environmental changes. Therefore, all known orchid species are protected in several countries by regional, national and international legislation. Several studies have cast doubts on the effectiveness of legislation to ensure the protection of wild orchids. We assessed the vitality of four orchid populations in a coastal area in Northern Italy, by monitoring the vegetative and reproductive traits of the orchid populations growing both in the protected sites comprising the Natura 2000 network, and in non-protected sites. We also monitored the level of environmental threat to orchid vitality. The early-flowering deceptive species (Ophrys sphegodes and Anacamptis morio) exhibited high vegetative vitality and experienced similar levels of environmental threat in the protected and non-protected areas. However, their reproductive success was strongly jeopardized, probably by a failed pollination that impeded the fruit set. The late-flowering, partially or totally rewarding species (Anacamptis pyramidalis and Anacamptis coriophora) were more strongly impacted by spring mowing and ungulate herbivory and alien species. Only for A. coriophora were the herbivory and alien species invasions lower at the protected vs. non-protected sites, which ensured a higher population vitality at the protected sites. We conclude that the environmental protection in our study area is ineffective for preserving orchids without targeted actions against specific environmental threats.

Seedling responses to moderate and severe herbivory: a field-clipping experiment with a keystone Mediterranean palm.

Plant-ungulate interactions are critical in shaping the structure of Mediterranean plant communities. Nevertheless, there is a dearth of knowledge on how plant intrinsic and extrinsic factors mediate the sign and strength of plant-ungulate interactions. This is most relevant when addressing natural or assisted restoration of plant communities in human-disturbed areas. We conducted field-clipping experiments simulating how different intensities of ungulate herbivory may affect the natural regeneration and establishment of the Mediterranean dwarf palm (Chamaerops humilis), a keystone species in Mediterranean ecosystems. We quantified seedling survival and size in two human-disturbed sites (SW Spain) where wild and domestic ungulates exert high herbivory pressure on vegetation. Severe clipping and seedling aging reduced rates of seedling survival. In contrast, moderate clipping did not affect seedling survival, suggesting a certain degree of C. humilis tolerance to herbivory. Severe clipping reduced seedling height strongly but not seedling diameter, and these effects seem to have decreased seedling survival. Nurse shrubs increased seedling size, which likely improved seedling survival. We also found seedling compensatory growth which varied between study sites. Field-clipping experiments can help disentangle effects of plant extrinsic and intrinsic factors on the sign and strength of plant-ungulate interactions and their ecological consequences on the dynamics of human-disturbed ecosystems. We call attention to the importance of appropriately managing scenarios of severe herbivory and summer droughts, particularly frequent in Mediterranean ecosystems, as synergic effects of such key drivers can negatively affect the structure and dynamics of plant communities and endanger their conservation.

Does Ungulate Herbivory Translate into Diversity of Woody Plants? A Long-Term Study in a Montane Forest Ecosystem in Austria

Different species-specific traits of woody plant species, feeding preferences of herbivores together with resulting effects on plant competition are expected to translate into different plant community structures and expressions of biodiversity. We studied the diversity of woody plant species (trees and shrubs) and structural diversity of forest trees, using a 30-year and an 18-year dataset of ungulate exclosure-control plot pairs in a mixed alpine forest community in Austria. We surveyed the tallest individuals per tree species and plot and analyzed the collective of top-height individuals per plot pair. Incidence data for exclosure and control plots were aggregated. Comparing species diversity and diversity of height classes on the plots throughout time, we calculated diversity profiles based on Hill numbers. Diversity of top height individuals and structural diversity, expressed by height classes, were two diversity aspects that differed between exclosures and control plots. Other diversity estimates of woody plant species showed huge variation without significant differences between plots. Height growth was significantly suppressed by ungulate herbivory. Effects of ungulate herbivores in forest ecosystems are highly complex and context-dependent and thus not reducible to simple top-down forces. Long-term surveys provide data that reflect “ultimate” effects of herbivory interacting with other drivers of community dynamics.

Divergent roles of herbivory in eutrophying forests

Ungulate populations are increasing across Europe with important implications for forest plant communities. Concurrently, atmospheric nitrogen (N) deposition continues to eutrophicate forests, threatening many rare, often more nutrient-efficient, plant species. These pressures may critically interact to shape biodiversity as in grassland and tundra systems, yet any potential interactions in forests remain poorly understood. Here, we combined vegetation resurveys from 52 sites across 13 European countries to test how changes in ungulate herbivory and eutrophication drive long-term changes in forest understorey communities. Increases in herbivory were associated with elevated temporal species turnover, however, identities of winner and loser species depended on N levels. Under low levels of N-deposition, herbivory favored threatened and small-ranged species while reducing the proportion of non-native and nutrient-demanding species. Yet all these trends were reversed under high levels of N-deposition. Herbivores also reduced shrub cover, likely exacerbating N effects by increasing light levels in the understorey. Eutrophication levels may therefore determine whether herbivory acts as a catalyst for the “N time bomb” or as a conservation tool in temperate forests.

Ungulate herbivores as drivers of aspen recruitment and understory composition throughout arid montane landscapes

AbstractHerbivory by wild and domestic ungulates can influence tree recruitment and understory forest communities throughout the world. Herbivore‐driven declines in tree recruitment have been observed for quaking aspen (Populus tremuloides), a foundation species whose health and management is recognized as a critical priority throughout much of its range. Livestock fencing is commonly used to promote aspen regeneration, but its effectiveness is rarely assessed, especially across large spatial scales. Using a livestock‐reduction experiment, we evaluated the effects of ungulate herbivory on aspen in the Great Basin and southern Cascades, an expansive and environmentally heterogeneous region where aspen faces the interacting threats of climate change, conifer encroachment, and herbivory. We found that livestock fencing only reduced the intensity of herbivore browsing on aspen when wild ungulate abundance was low and did not increase stem densities of aspen recruits. Contrary to expectations, wild ungulate abundance was a strong driver of browsing intensity on juvenile aspen within fenced, but not unfenced, aspen stands, and when the abundance of these herbivores was high, browsing intensity in fenced stands exceeded that in unfenced stands. The density of aspen recruits decreased with browsing intensity on juvenile aspen and with the density of both adult aspen and conifers, suggesting that both herbivory and intra‐ and interspecific competition are important drivers of recruitment. Fire history was also an important driver of recruitment, with stands that burned 10–20 years ago having the greatest density of aspen recruits. Finally, in the stand understory, we found that livestock fencing decreased forb cover, increased shrub species richness, and increased the cover of exotic annual grasses, a group dominated by Bromus tectorum. This latter finding suggests that livestock fencing may not be appropriate in areas where controlling the spread of this invader is a priority. In sum, our findings indicate that aspen recruitment is limited by browsing by both wild and domestic ungulates, is mediated by competition with neighboring trees and fire history, and will require management actions beyond livestock fencing, as this approach does not control browsing by wild ungulates.

Trophic Interactions Drive Tree Establishment in the Serengeti Ecosystem

Photo 1. Tree seedlings in savanna ecosystems, such as this Vachellia tortillis seedling in Tanzania's Serengeti National Park, face competition from grasses and are regularly damaged by fire and browsers. These filters to survival result in demographic bottlenecks in savanna tree populations. We found that grass competition accounted for the majority of early seedling mortality. Photo credit: Jason Donaldson. Photo 2. Regular fires, fueled by a buildup in grass biomass, kill high numbers of tree seedlings with mortality rates highest in areas where grass competition is historically high. Photo credit: Jason Donaldson. Photo 3. Impala (Aepyceros melampus) and other browsing ungulate herbivores remove vegetative growth and kill seedlings, exacerbating the negative effects of fire and grass competition on seedling establishment. There was no evidence that browsers in the Serengeti were responding to lion kill risk in a way that changed their spatial impacts on tree establishment. Photo credit: Jason Donaldson. Photo 4. Expansive herds of wildebeest (Connochaetes taurinus), zebra (Equus quagga), and other ungulate grazers in the Serengeti remove grass biomass and can reduce the detrimental effects of grass competition and fire, resulting in increased tree seedling establishment. Despite this, lion kill risk does not meaningfully impact grazer effects on tree seedling survival. Photo credit: Jason Donaldson. Photo 5. Tree seedling establishment will substantially increase in systems containing high densities of grazing livestock, where wild browsing ungulates have been removed, and tree seedlings are released from grass competition and fire. It is likely that this scenario is occurring over broad areas of savanna globally. Photo credit: Jason Donaldson. These photographs illustrate the article “Fire, grazers and browsers interact with grass competition to determine tree establishment in an African savanna” by Jason E. Donaldson, Ricardo Holdo, Jeremia Sarakikya, T. Michael Anderson published in Ecology. https://doi.org/10.1002/ecy.3715

Long-term plant community responses to resource objective wildfires in montane coniferous forests of Grand Canyon National Park, USA

Interest in use of naturally ignited wildland fires managed to meet multiple resource objectives (resource objective wildfire) is increasing among U.S. public forest managers; however, only a limited number of studies have examined this approach for conserving or restoring understory plant diversity, productivity, and community structure. We analyzed understory community changes two andtwelve yearsafter resource objective fire, using permanent sample plotsin three montane contrasting forest types in Grand Canyon National Park,AZ.Our findingsindicatedthat species compositionin the pine-oak forest rebounded to be similar to that observed before the fire, but plant cover did not recover to pre-fire levels by the twelfth year post-fire. Plant cover showed mixed results post-fire in mixed-conifer and spruce-fir forests, and species composition was still characterized by ruderal species twelve years later. Patterns observed in this study likely reflect interacting factors of burn severity, periodic drought, large ungulate herbivory, and inherent site variability. Other than cheatgrass (Bromus tectorum), we found no non-native speciesoccurring with high frequency. Continued monitoring with increased frequency and intensity could lead to better understanding of long-term changes in these forests after resource objective fire, and enhance our understanding of important drivers of variation including interactions ofclimate, burn severity and herbivory by large ungulates.

Herbivory, intraspecific trait variability and back to herbivory

Ungulate herbivory is a key driver of leaf trait syndromes and defense strategies at the ecological and macroevolutionary levels. Herbivory should also cause short‐term evolution within plant populations, but few studies have experimentally tested this prediction. We set out to experimentally assess the plastic and heritable effects of contemporary history of ungulate herbivory on 1) leaf trait syndromes at the intraspecific level, and on the defense traits, 2) leaf size and 3) size of spines.We measured leaf traits (specific leaf area, leaf dry matter content, leaf nitrogen concentration and leaf mechanical resistance) in populations of six grass and four woody legume species that had grown under contrasting pressures of domestic ungulate herbivory for at least 40 years. We then performed a common garden experiment, measuring the same traits. Next, we performed cutting experiments on the plants grown in the common garden to measure leaf trait plasticity. In the woody species, we also measured the length of spines and the leaf size (average area per leaf), in the field and in the common garden experiment.We found that in grasses field herbivory over decades produced a displacement toward more acquisitive leaf trait syndromes. Most of these changes appeared largely due to plasticity. In woody legume species, in contrast, herbivory produced a displacement towards more conservative leaves, longer spines and smaller leaves, with the trait differences between histories of herbivory being mostly hereditary. These results point out that vertebrate herbivory can lead to evolutionary processes at contemporary time scales even in long‐lived perennial plant species, but the direction of such change depends on the life form of the species. These divergent effects could depend on the predominant anti‐herbivore strategies in the different life forms (tolerance versus resistance).

Cover Image

COVER PHOTO: An African bush elephant (Loxodonta africana) and two Grévy's zebras (Equus grevyi) forage at Mpala Research Centre and Conservancy in Laikipia, Kenya during July 2019. These and other herbivores suppress plant biomass, and herbivore extinctions may trigger a cascade of broader changes throughout the ecological communities they once inhabited. In their Data Paper published in this issue, Alston, et al. (Article e3649; doi:10.1002/ecy.3649) provide twelve years of data on vegetation, rainfall, and mammal densities in the Ungulate Herbivory Under Rainfall Uncertainty (UHURU) Experiment, a series of long-term herbivore exclosures designed to mimic the effects of size-selective herbivore extinctions on plant and small mammal communities inhabiting African savannas. Photo credit: Courtney Reed.

Context‐dependent effects of shifting large herbivore assemblages on plant structure and diversity

Abstract Despite wide recognition of the importance of anthropogenically driven changes in large herbivore communities—including both declines in wildlife and increases in livestock—there remain large gaps in our knowledge about the impacts of these changes on plant communities, particularly when combined with concurrent changes in climate. Considering these prominent forms of global change in tandem enables us to better understand controls on savanna vegetation structure and diversity under real‐world conditions. We conducted a field experiment using complete and semi‐permeable herbivore exclosures to explore the difference in plant communities among sites with wild herbivores only, with cattle in addition to wild herbivores, and with no large herbivores. To understand variation in effects across climatic contexts, the experiment was replicated at three locations along a topoclimatic gradient in California. Critically, this is the first such experiment to compare cattle and wildlife impacts along an environmental gradient within a single controlled experiment. Vegetation structure responded strongly to herbivore treatment regardless of climate. Relative to the isolated effects of wildlife, exclusion of all large herbivores generally increased structural components related to cover and above‐ground biomass while the addition of cattle led to reductions in vegetation cover, litter, shading and standing biomass. Furthermore, wildlife had a consistent neutral or positive effect on plant diversity, while the effect of livestock addition was context dependent. Cattle had a neutral to strongly negative effect at low aridity, but a positive effect at high aridity. These results suggest that (a) herbivore effects can override climate effects on vegetation structure, (b) cattle addition can drive different effects on diversity and (c) herbivore effects on diversity are modulated by climate. Synthesis. Our results illustrate very distinctive shifts in plant communities between two realistic forms of change in ungulate herbivore assemblages—livestock addition and large herbivore losses—particularly for plant diversity responses, and that these responses vary across climatic contexts. This finding has important implications for the management and protection of plant biodiversity given that over a quarter of the Earth's land area is managed for livestock and climate regimes are changing globally.

Ecological consequences of large herbivore exclusion in an African savanna: 12 years of data from the UHURU experiment.

Diverse communities of large mammalian herbivores (LMH), once widespread, are now rare. LMH exert strong direct and indirect effects on community structure and ecosystem functions, and measuring these effects is important for testing ecological theory and for understanding past, current, and future environmental change. This in turn requires long-term experimental manipulations, owing to the slow and often nonlinear responses of populations and assemblages to LMH removal. Moreover, the effects of particular species or body-size classes within diverse LMH guilds are difficult to pinpoint, and the magnitude and even direction of these effects often depends on environmental context. Since 2008, we have maintained the Ungulate Herbivory Under Rainfall Uncertainty (UHURU) experiment, a series of size-selective LMH exclosures replicated across a rainfall/productivity gradient in a semiarid Kenyan savanna. The goals of the UHURU experiment are to measure the effects of removing successively smaller size classes of LMH (mimicking the process of size-biased extirpation) and to establish how these effects are shaped by spatial and temporal variation in rainfall. The UHURU experiment comprises three LMH-exclusion treatments and an unfenced control, applied to nine randomized blocks of contiguous 1-ha plots (n=36). The fenced treatments are MEGA (exclusion of megaherbivores, elephant and giraffe), MESO (exclusion of herbivores ≥40 kg), and TOTAL (exclusion of herbivores ≥5kg). Each block is replicated three times at three sites across the 20-km rainfall gradient, which has fluctuated over the course of the experiment. The first 5 years of data were published previously (Ecological Archives E095-064) and have been used in numerous studies. Since that publication, we have (1) continued to collect data following the original protocols, (2) improved the taxonomic resolution and accuracy of plant and small-mammal identifications, and (3) begun collecting several new data sets. Here, we present updated and extended raw data from the first 12 years of the UHURU experiment (2008-2019). Data include daily rainfall data throughout the experiment; annual surveys of understory plant communities; annual censuses of woody-plant communities; annual measurements of individually tagged woody plants; monthly monitoring of flowering and fruiting phenology; every-other-month small-mammal mark-recapture data; and quarterly large-mammal dung surveys. There are no copyright restrictions; notification of when and how data are used is appreciated and users of UHURU data should cite this data paper when using the data.

The long‐term impacts of deer herbivory in determining temperate forest stand and canopy structural complexity

Abstract Ungulates place immense consumptive pressure on forest vegetation globally, leaving legacies of reduced biodiversity and simplified vegetative structure. However, what remains unresolved is whether browse‐induced changes occurring early in succession ultimately manifest themselves in the developed forest canopy. Understanding the development and persistence of these legacies is critical as canopy structure is an important determinant of forest ecosystem functions such as carbon sequestration and wildlife habitat. We measured how white‐tailed deer Odocoileus virginianus browse during stand initiation affected canopy structure, tree species richness, diversity, stem density, and basal area on Pennsylvania's Allegheny Plateau using a portable canopy LiDAR system. We capitalized on an historic deer enclosure experiment where forests were subjected to four deer densities (4, 8, 15, and 25 deer/km2) for 10 years following stand initiation. Deer browsing impacts on the forest canopy are apparent nearly four decades since stand initiation. The highest deer density treatment experienced a significant reduction in tree species diversity, density, and basal area with stands becoming dominated by black cherry Prunus serotina. Reductions in overstorey diversity and tree density resulted in a more open canopy with low leaf area and high horizontal leaf variability. Canopies were tallest at the lowest and highest deer densities. Synthesis and applications. Using a portable canopy LiDAR system and a former deer enclosure experiment, we show that high deer browsing pressure during stand initiation can have a decades‐long impact on stand and canopy structure. High deer densities led to stands with lower species diversity and tree density, which resulted in canopies that were taller and less dense. As remote sensing of the canopy becomes more prevalent, considering the legacy of ungulate herbivory on canopy structure may inform both land management and our understanding of ecological function, such as forest carbon sequestration, maintenance of diverse understory communities, and creation of wildlife habitat

Functional trait data for vascular plant species from northeastern North America.

Wisconsin's plant communities are responding to shifting disturbance regimes, habitat fragmentation, aerial nitrogen deposition, exotic species invasions, ungulate herbivory, and successional processes. To better understand how plant functional traits mediate species' responses to changing environmental conditions, we collected a large set of functional trait data for vascular plant species occupying Wisconsin forests and grasslands. We used standard protocols to make 76,213 measurements of 34 quantitative traits. These data provide rich information on genome size, physical leaf traits (length, width, circularity, thickness, dry matter content, specific leaf area, etc.), chemical leaf traits (carbon, nitrogen, phosphorus, potassium, calcium, magnesium, ash), life history traits (vegetative and flower heights, seed mass), and traits affecting plant palatability (leaf fiber, fat, and lignin). These trait values derive from replicate measurements on 12+ individuals of each species from multiple sites and 45+ individuals for a selected subset of species. Measurements typically reflect values for individuals although some chemical traits involved composite samples from several individuals at the same site. We also qualitatively characterized each species by plant family, woodiness, functional group, and Raunkiaer lifeform. These data allow us to characterize trait dimensionality, differentiation, and covariation among temperate plant species (e.g., leaf and stem economic syndromes). We can also characterize species' responses to environmental gradients and drivers of ecological change. With survey and resurvey data available from >400 sites in Wisconsin, we can analyze variation in community trait distributions and diversity over time and space. These data therefore allow us to assess how trait divergence vs. convergence affects community assembly and how traits may be related to half-century shifts in the distribution and abundance of these species. The data set can be used for non-commercial purposes. The data set is licensed as follows: CC-By Attribution 4.0 International. We request users cite both the OSF data set and this Ecology data paper publication.