Patterns Of Herbivory Research Articles

Hiding in the background: community-level patterns in invertebrate herbivory across the tundra biome

Invertebrate herbivores depend on external temperature for growth and metabolism. Continued warming in tundra ecosystems is proposed to result in increased invertebrate herbivory. However, empirica ...

Leaf traits mediate changes in invertebrate herbivory along broad environmental gradients on Mt. Kilimanjaro, Tanzania.

Temperature, primary productivity, plant functional traits, and herbivore abundances are considered key predictors of leaf herbivory but their direct and indirect contributions to community-level herbivory are not well understood along broad climatic gradients. Here, we determined elevational herbivory patterns and used a path analytical approach to disentangle the direct and indirect effects of climate, land use, net primary productivity (NPP), herbivore abundance, and plant functional traits on community-level invertebrate herbivory along the extensive elevational and land use gradients at Mt. Kilimanjaro, Tanzania. We recorded standing leaf herbivory caused by leaf chewers, leaf miners and leaf gallers on 55 study sites distributed in natural and anthropogenic habitats along a 3,060m elevation gradient. We related the total community-level herbivory to climate (temperature and precipitation), NPP, plant functional traits (specific leaf area, leafcarbon-to-nitrogen [CN]ratio and leaf nitrogen-to-phosphorus [NP] ratio) and herbivore abundances. Leaf herbivory ranged from 5% to 11% along the elevation gradient. Total leaf herbivory showed unimodal pattern in natural habitats but a strongly contrasting bimodal pattern in anthropogenic habitats. We also detected some variation in the patterns of leaf herbivory along environmental gradients across feeding guilds with leaf chewers beingresponsible for a disproportionally large part of herbivory. Path analyses indicated that the variation in leaf herbivory was mainly driven by changes in leaf CN and NP ratios which were closely linked to changes in NPP in natural habitats. Similarly, patterns of leaf herbivory in anthropogenic habitats were best explained by variation in leaf CN ratios and a negative effect of land use. Our study elucidates the strong role of leaf nutrient stoichiometry and its linkages to climate and NPP for explaining the variation in leaf herbivory along broad climatic gradients. Furthermore, the study suggests that climatic changes and nutrient inputs in the course of land use change may alter leaf herbivory and consequently energy and nutrient fluxes in terrestrial habitats.

Grazing animals drove domestication of grain crops.

In addition to large-seeded cereals, humans around the world during the mid-Holocene started to cultivate small-seeded species of herbaceous annuals for grain, including quinoa, amaranth, buckwheat, the millets and several lost crops domesticated in North America. The wild ancestors of these crops have small seeds with indigestible defences and do not germinate readily. Today, these wild plants exist in small fragmentary stands that are not appealing targets for foragers. This combination of traits has led many to argue that they must have been a food of last resort. We propose a new explanation: the domestication of small-seeded annuals involved a switch from endozoochoric dispersal (through animal ingestion) to human dispersal. Humans encountered these plants in dense stands created by grazing megafauna, making them easy to harvest. As humans began to cultivate these plants they took on the functional role of seed dispersers, and traits associated with endozoochory were lost or reduced. The earliest traits of domestication-thinning or loss of indigestible seed protections, loss of dormancy and increased seed size-can all be explained by the loss of the ruminant dispersal process and concomitant human management of wild stands. We demonstrate, by looking at rangeland ecology and herd animal herbivory patterns, that the progenitors of all of these species evolved to be dispersed by megafaunal ruminants and that heavy herbivory leads to dense homogenous clusters of endozoochoric plants. Hence, easily harvested stands on nitrogen hot spots near water sources would have existed in regions where these plants were domesticated. Future experimental and ecological studies could enhance our understanding of the relationships between specific crops and their possible ruminant dispersers.

Macroevolutionary effects on primate trophic evolution and their implications for reconstructing primate origins

The visual-predation hypothesis proposes that certain derived features shared by crown primates reflect an insectivorous ancestry. Critics of this idea have argued that because insectivory is uncommon among extant primates it is unlikely to have been a major influence on early primate evolution. According to this perspective, the low frequency of insectivory indicates that it is an apomorphic deviation from the mostly conserved primate ecological pattern of herbivory. The present study tests two alternative hypotheses that are compatible with an insectivorous ancestor: (1) that trophic evolution was biased, such that herbivory evolved repeatedly with few shifts back to insectivory, and (2) that insectivorous lineages have diversified at a lower rate than herbivorous lineages owing to differential trophic effects on speciation and extinction probabilities. Model-based analysis conducted using trait data for 307 extant primate species indicates that rates of transition into and out of insectivory are similar, rejecting the hypothesis of biased trophic evolution. On the other hand, the hypothesis of asymmetric diversification is supported, with insectivorous lineages having a lower rate of diversification than herbivorous lineages. This correlation is mediated by activity pattern: insectivory occurs mostly in nocturnal lineages, which have a lower diversification rate than diurnal lineages. The frequency of insectivory also appears to have been shaped by repeated transitions into ecological contexts in which insectivory is absent (large body size) or rare (diurnality). These findings suggest that the current distribution of trophic strategies among extant primates is the result of macroevolutionary processes that have favored the proliferation and persistence of herbivory relative to insectivory. This conclusion implies that the low frequency of insectivory is not necessarily evidence against the visual-predation hypothesis.

Functional attributes of two Croton species in different successional stages of tropical dry forest: effects on herbivory and fluctuating asymmetry patterns

Tropical dry forests are among the most threatened ecosystems in the world. After habitat perturbation occurs, the habitat recovers naturally through ecological succession. This succession can modify functional attributes of plants, which in turn, can affect herbivorous insects’ performance. We analyzed morphological, functional, and chemical traits associated with herbivory patterns in Croton roxanae and C. suberosus, that occur in mature and secondary forests in the tropical dry forest of Chamela, Jalisco. Leaf area and leaf thickness were higher in secondary forest, while leaf density and fresh leaf mass were higher in mature forest. Dry leaf mass, specific leaf area, chlorophyll content, and water content showed variation between species in both forest conditions. The concentration of secondary metabolites showed variation between species and forest conditions. Croton roxanae showed higher herbivory in mature forest, and C. suberosus did not show differences between the two conditions. Leaves in secondary forest were slightly longer and broader than leaves in mature forest. Croton species showed higher fluctuating asymmetry in secondary forest. Herbivory was not associated with levels of fluctuating asymmetry levels in both Croton species. Our results suggest that plant attributes are influenced by forest condition, which in turn, indirectly affect the attack of herbivores.

Effects of insularity on insect leaf herbivory and chemical defences in a Mediterranean oak species

AbstractAimResearch on plant–herbivore interactions has shown that islands typically have low abundances and diversity of herbivores because of barriers to dispersal, isolation and reduced land area. Islands commonly have lower levels of herbivory relative to mainland regions, and, as a consequence, insular plants should exhibit lower levels of defences than their mainland counterparts. Despite these predictions, there are significant gaps in our understanding of insularity effects on plant–herbivore interactions. For instance, most work addressing the effects of insularity on plant–herbivore interactions have compared one or a few islands with a single mainland site. In addition, studies have measured herbivory or plant defences but not both, and the influence of abiotic factors has been neglected.LocationMediterranean Basin (from Spain to Greece).TaxonQuercus ilex L.MethodsWe conducted a large‐scale study to investigate whether insect leaf herbivory and plant chemical defences in holm oak (Quercus ilex L.) differ between insular versus mainland populations. We further investigated mechanisms by which insularity effects on herbivory may take place by assessing the influence of defences and climatic variables on herbivory.ResultsWe found that insular populations exhibited lower herbivory and higher defences (condensed tannins) than their mainland counterparts. Our analyses, however, suggest that these concomitant patterns of insect herbivory and plant defences were seemingly unrelated as island versus mainland differences in defences did not account for the observed pattern in herbivory. Furthermore, climatic factors did not explain insularity effects on either herbivory or plant defences.Main conclusionsOverall, this study provides one of the most robust assessments to date on insularity effects on herbivory and builds towards a better understanding of the ecology and evolution of plant–insect interactions in insular ecosystems.

Applying and refining DNA analysis to determine the identity of plant material extracted from the digestive tracts of katydids.

BackgroundFeeding habits are central to animal ecology, but it is often difficult to characterize the diet of organisms that are arboreal, nocturnal, rare, or highly mobile. Genetic analysis of gut contents is a promising approach for expanding our understanding of animal feeding habits. Here, we adapt a laboratory protocol for extracting and sequencing plant material from gut contents and apply it to Neotropical forest katydids (Orthoptera: Tettigoniidae) on Barro Colorado Island (BCI) in Panama.MethodsOur approach uses three chloroplast primer sets that were previously developed to identify vegetation on BCI. We describe the utility and success rate of each primer set. We then test whether there is a significant difference in the amplification and sequencing success of gut contents based on the size or sex of the katydid, the time of day that it was caught, and the color of the extracted gut contents.ResultsWe find that there is a significant difference in sequencing success as a function of gut color. When extracts were yellow, green, or colorless the likelihood of successfully amplifying DNA ranged from ~30–60%. When gut extracts were red, orange, or brown, amplification success was exceptionally low (0–8%). Amplification success was also higher for smaller katydids and tended to be more successful in katydids that were captured earlier in the night. Strength of the amplified product was indicative of the likelihood of sequencing success, with strong bands having a high likelihood of success. By anticipating which samples are most likely to succeed, we provide information useful for estimating the number of katydids that need to be collected and minimizing the costs of purifying, amplifying, and sequencing samples that are unlikely to succeed. This approach makes it possible to understand the herbivory patterns of these trophically important katydids and can be applied more broadly to understand the diet of other tropical herbivores.

Multi-Trophic Species Interactions Shape Seascape-Scale Coral Reef Vegetation Patterns

How species interactions shape habitat structure is a longstanding question in ecology. A curious phenomenon reflecting ecological self-organization around reef habitat structures exists on coral reefs: large-scale (hundreds to hundreds of thousands of m2) halo-like patterns surrounding patch reefs, i.e., individual coral reefs that are often separated by seagrass or macroalgal meadows. These “halos,” long known to occur in various locations worldwide, reflect a distinct band of unvegetated sediments surrounding coral patch reefs. However, the full suite of mechanisms controlling them have never been rigorously explored, perhaps due to the common assumption dating back nearly 50 years that they arise solely from reef-based herbivory patterns shaped by anti-predator behavior. Here we provide empirical evidence from a set of halos within Australia's Great Barrier Reef that risk-averse foraging and a previously unrecognized functional group contribute to halo formation, demonstrating that these halos cannot be explained by any one mechanism in isolation. Our results show that halos are a more complex ecological phenomenon than previously assumed by the majority of studies of halos. Specifically, risk-averse grazing by herbivores is likely a key mechanism behind the formation of halos, as generally assumed, but bioturbators also play a central role. This knowledge furthers our understanding of how small-scale species interactions can structure habitat at landscape scales. These large-scale habitat features are important because they affect at least one important ecosystem function, carbon storage, and potentially others (e.g., biological nutrient transfer). These results also raise the question of whether other self-organized ecological patterns may be more nuanced than is currently assumed. This study capitalizes on recent advances in high resolution satellite imagery accessibility that allow ecologists to measure landscape-scale habitat features nearly everywhere on land and in shallow seas. Our results suggest that halos may hold potential as the basis for a tool for remotely observing ecological interactions and measuring large-scale ecosystem change on coral reefs.

Urbanization drives unique latitudinal patterns of insect herbivory and tree condition

Urban landscapes are characterized by high proportions of impervious surface resulting in higher temperatures than adjacent natural landscapes. In some cities, like those at cooler latitudes, trees may benefit from warmer urban temperatures, but trees in many cities are beset with problems like drought stress and increased herbivory. What drives patterns of urban tree health across urbanization and latitudinal temperature gradients? In natural systems, latitude–herbivory relationships are well‐studied, and recent temperate studies have shown that herbivory generally increases with decreasing latitudes (warmer temperatures). However, the applicability of this latitude–herbivory theory in already‐warmed urban systems is unknown. In this study, we investigated how the interaction of urbanization, latitudinal warming and scale insect abundance affected urban tree health. We predicted that trees in warmer, lower latitude cities would be in poorer health at lower levels of urbanization than trees at cooler, higher latitudes due to the interaction of urbanization, latitudinal temperature and herbivory. To evaluate our predictions, we surveyed the abundance of scale insect herbivores on a single, common tree species Acer rubrum in eight US cities spanning 10° of latitude. We estimated urbanization at two extents, a local one that accounted for the direct effects on an individual tree, and a larger one that captured the surrounding urban landscape. We found that urban tree health did not vary with latitudinal temperature but was best predicted by local urbanization and herbivore abundance. We did not observe increased herbivore abundance in warmer, lower latitudes cities, but instead herbivore abundance peaked in the mid latitudes of our study. This study demonstrates that urban landscapes may deviate from classical theory developed in natural systems and reinforces the need for research reconciling ecological patterns in urban landscapes.

Human altered disturbance patterns and forest succession: impacts of competition and ungulate herbivory.

Human activities are altering patterns of ungulate herbivory and wildfire regimes globally with large potential impacts on plant community succession and ecosystem resilience. Aspen (Populus tremuloides) is a keystone species which co-exists with conifer species across temperate forests in North America. Aspen sucker regeneration which is the foundation of aspen-conifer forests succession is often a targeted food source by multiple ungulate species. Using a region-wide exclosure network across a broad gradient of aspen-conifer overstory abundance, we empirically tested the effects of ungulate herbivory and conifer competition (that increases with fire suppression), on the regeneration and recruitment of aspen forests over a 4-year period. The study results indicate that ungulate herbivory and increasing abundance of overstory conifers dramatically reduced aspen regeneration and recruitment success. The average height of aspen suckers exposed to ungulate herbivory was 72% shorter than aspen suckers in fenced plots and resulted in 24% less recruitment. There was a 9% decrease in aspen recruitmentand 12% decrease in averageaspen height with every 20% increase in overstory conifer density. Aspen suckers were most vulnerable to herbivory at 70cm height, with the probability of herbivory decreasing under 50cm or above 90cm. Steep slope angles and higher winter precipitation increased aspen regeneration and recruitment success. Reduction in aspen recruitment in response to ungulate herbivory and competition by conifers may result in loss of biodiversity, altered forest function and loss of key ecosystem services because of the important role that aspen plays in facilitating forest succession and biodiversity.

Biases in studies of spatial patterns in insect herbivory

AbstractThe properties of the human mind are responsible for a number of biases that affect the quality of scientific research. However, scientists working in the fields of ecology and environmental science rarely take these biases into account. We conducted a meta‐analysis of data extracted from 125 publications comparing woody plant damage by defoliating insects in different environments in order to understand the extent to which our knowledge on spatial patterns in herbivory is affected by various biases. We asked which research methods are most prone to biases and whether these biases lead to overestimation of the effects under study. The effect sizes (ESs) decreased with increases in the numbers of plant species involved in the study, with 61% lower ESs for herbivory estimated on all plants growing in study plots compared to herbivory on selected species. ESs also depended on the leaf sampling procedure: when all leaves from a tree or branch were sampled for measurements of herbivory or when random or systematic selection protocols were applied, ESs were 74% smaller than in cases of more subjective haphazard selection. In addition, ESs were 97% and 135% greater when the person conducting sampling and measuring was aware of the research hypothesis or sample origin, when compared with situations when the observer was blinded to these factors. The impacts of cognitive biases on the study outcomes significantly decreased with the increase in publication year; however, this pattern emerged mostly due to high‐ranked journals and was non‐significant for other journals. Using the studies of spatial patterns in herbivory as an example, we showed that our ecological and environmental knowledge is considerably biased due to an unconscious tendency of researchers to find support for their hypotheses and expectations, which generally leads to overestimation of the effects under study. Cognitive biases can be avoided by using different methods, such as applying randomization procedures in sampling and blinding of research hypotheses and sample origins. These measures should be seen as obligatory; otherwise, accumulation of the biased results in primary studies may ultimately lead to false general conclusions in subsequent research synthesis.

Invasive Species Appear to Disrupt the Top-Down Control of Herbivory on a Mexican Oceanic Island

Oceanic islands are simplified ecosystems, which are very useful for understanding the processes related to the structuring of natural communities and for assessing cascade effects after perturbations. Socorro Island in the Pacific Ocean reaches up to 1040 m in elevation, and the northeastern side that faces trade winds is more humid than the leeward southwestern side. Apparently, the lack of freshwater throughout most of the island has prevented the spread of invasive cats and sheep to the north, but these species have severely impacted land bird communities on the southern side, where they have potentially disrupted functions such as the top-down control of herbivory. We tested the relative importance of bottom-up (soil and foliage nutrient availability) and top-down (predation of insect herbivores) drivers of herbivory in three endemic tree species. By experimentally excluding birds, we assessed changes in herbivory when top-down control was disrupted, and we also evaluated herbivore predation rates using artificial caterpillars. Our findings suggest that the herbivory patterns on Socorro Island are driven by the top-down control of herbivores by land birds, because their exclusion resulted in higher herbivory (12%) despite low nitrogen content (bottom-up control) of leaves on the north side, whereas the leaves were richer in nitrogen on the south side but no change in herbivory was observed when birds were excluded, and overall, herbivory was 30% higher than in the north, suggesting release of herbivores from top down control on the south side, probably due to effects of invasive predators (cats) or sheep, which may have reduced suitable bird habitats. A better understanding of how drivers of ecological interactions operate on Socorro Island may increase the likelihood of success of future restoration programs.

The Potential use of Seagrass Herbivory Patterns as an Indicator of Herbivore Community Change after Tropical Marine Protected Area Establishment

The Potential use of Seagrass Herbivory Patterns as an Indicator of Herbivore Community Change after Tropical Marine Protected Area Establishment

Effect of Leaf Quality on Herbivory of Three Atlantic Forest Species

ABSTRACT The present study evaluated the relationship between herbivory and leaf quality of three abundant tree species (Cupania oblongifolia Mart., Siparuna guianensis Aubl. and Xylopia sericea St. Hill.) in the understory of a eucalyptus plantation and an adjacent Atlantic Rainforest fragment. Herbivory differed among species, since X. sericea was less attacked, while C. oblongifolia and S. guianensis were more. The species showed similar herbivory patterns between the investigated areas (eucalyptus plantation versus forest), although X. sericea showed higher herbivory rates in the forest fragment. Fiber values (NDF, CEL and LIG) varied significantly among the species, being lower in S. guianensis. X. sericea, the species with the highest C/N ratio, higher concentrations of total phenols, smaller leaf area and higher trichome density, showed the lowest leaf herbivory rates, corroborating the hypothesis that herbivory is lower in species with lower nutritional quality and a greater set of leaf defense features.

Mycorrhizal interactions do not influence plant-herbivore interactions in populations of Clarkia xantiana ssp. xantiana spanning from center to margin of the geographic range.

Multispecies interactions can be important to the expression of phenotypes and in determining patterns of individual fitness in nature. Many plants engage in symbiosis with arbuscular mycorrhizal fungi (AMF), but the extent to which AMF modulate other species interactions remains poorly understood. We examined multispecies interactions among plants, AMF, and insect herbivores under drought stress using a greenhouse experiment and herbivore choice assays. The experiment included six populations of Clarkia xantiana (Onagraceae), which span a complex environmental gradient in the Southern Sierra Nevada of California. Clarkia xantiana's developing fruits are commonly attacked by grasshoppers at the end of the growing season, and the frequency of attack is more common in populations from the range center than range margin. We found that AMF negatively influenced all metrics of plant growth and reproduction across all populations, presumably because plants supplied carbon to AMF but did not benefit substantially from resources potentially supplied by the AMF. The fruits of plants infected with AMF did not differ from those without AMF in their resistance to grasshoppers. There was significant variation among populations in damage from herbivores but did not reflect the center‐to‐margin pattern of herbivory observed in the field. In sum, our results do not support the view that AMF interactions modulate plant–herbivore interactions in this system.

Perspectives on grassland conservation efforts: should we rewild to the past or conserve for the future?

Grasslands are among the most imperilled biomes of the world. Identifying the most appropriate framework for restoring grasslands is dependent on the objectives of restoration, which is inherently determined by human priorities. Debates over the appropriate conservation model for grasslands have often focused on which species of herbivores should be the focus of restoration efforts. Here we discuss three perspectives of herbivore-based conservation in North American grasslands. First, the Pleistocene rewilding perspective is based upon the idea that early humans contributed to the demise of megafauna that were important to the evolution and development of many of North America's grasslands; therefore, their aim of restoration is rewilding of landscapes to pre-human times. Second, the bison rewilding perspective considers American bison a keystone herbivore that is culturally and ecologically important to North American grasslands. A third perspective focuses on restoring the pattern and processes of herbivory on grasslands and is less concerned about which herbivore is introduced to the landscape. We evaluate each of these three conservation perspectives in terms of a framework that includes a human domain, an herbivore domain and a biophysical domain. While all conservation perspectives partly address the three domains, they all fall short in key areas. Specifically, they fail to recognize that past, current and future humans are intimately linked to grassland patterns and processes and will continue to play a role in structuring grasslands. Furthermore, these perspectives seem to only superficially consider the role of fragmentation and climate change in influencing grassland patterns and processes. As such, we argue that future grassland conservation efforts must depend on the development of a model that better integrates societal, economic and policy objectives and recognizes climate change, fragmentation and humans as an integral part of these ecosystems.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

Patrones de herbivoría en Vassobia breviflora (Solanaceae): variación en el daño foliar y selección natural mediada por herbívoros

Herbivore mediated-selection shapes the evolution of defensive plant traits. Knowledge about the role of herbivores as mediators of selection is scarce and even more if herbivore functional groups are considered. The objectives of this work were (1) to describe the variation in foliar traits between populations and both between and intra-plants within a population, (2) to explore the relationship between the variation in the herbivory level and foliar traits, (3) to determine the relationship between leaf traits and damage patterns and (4) estimate the selection regimes by different herbivore functional groups. We conducted this study in four populations of Vassobia breviflora in northwestern Argentina (Yungas). The foliar traits considered were size, leaf area (af), shape (leaf length / width ratio; laf) and proportion of leaf area removed (pafr) (N = 1582 leaves, 57 plants). The herbivores consumed 15.6 % of the leaf area and 76.8 % of the variation in the pafr occurred at the sub-individual level. The damage pattern was dominated by cutter herbivores (70 %), followed by a dotted herbivory pattern (22 %), mixed 5 % and 1 % miner. Nonlinear selection was detected for laf (γii = -0.180; EE = 0.76; P < 0.05), and correlational selection between the cutter damage and af (γij = -1.297; SE = 0.62; P < 0.05) and between the dotted damage and af (γij = -1.130; SE = 0.76; P < 0.05). Natural selection favored plants with small leaves and high foliar removal and large leaves with less damage and selection against larger leaves with greater damage was detected. In addition, deduced from the relationship between the damage type and the relative fitness, the selection would favor the dotted damage over the cutter one. The plants would resolve the conflict with the herbivores according to the damage type and natural selection would regulate the foliar display as a strategy to deal with the herbivore diversity.

Temporal patterns of ungulate herbivory and phenology of aspen regeneration and defense.

Ungulate herbivory can create strong top-down effects on forest recruitment, especially after fire. Defense strategies of tree species against ungulate herbivory include escape through vertical growth and resistance through the production of defense compounds. Using afour-way fence design and camera traps we characterized the differential impacts of ungulate herbivores (deer, elk, cattle) on aspen forest recruitment and plant defense responses and how they vary depending on the timing of herbivory. Aspen height growth was greatest between June and August and ungulate use of aspen was highest in July and August. Threeyears after fire, height of aspendiffered among fence treatments with full ungulate exclusion > deer-only plots > native ungulate plots > fenceless plots: 108 ± 4cm, 94 ± 4cm, 89 ± 4cm, and 65 ± 4cm, respectively. Fenceless plots had the highest rates of removal of apical meristems by the end of 2014 and 2015 (61% and 53%, respectively). Native ungulate plots, and deer-only plots both had similar removal of apical meristems in 2014 (37% and 39%, respectively). The highest phenolic glycoside concentrations were associated with an 80% reduction in meristem removal and four-fold greater aspen height by the end of summer. Low nitrogen was associated with an 86% reduction in apical meristem removal and threefold greater aspen height. In conclusion, our study suggests that high ungulate abundance can have detrimental impacts on forest recruitment and that high aspen defense chemistry and lower leaf N deters ungulate herbivory, especially in the late summer.

Does local isolation allow an invasive thistle to escape enemy pressure?

Enemy release often is invoked to explain the success of invasive plants: an invader benefits from reduced attack as it escapes specialized enemies through the invasion process. Enemy release typically is thought of as occurring at large geographic scales, but local-scale interactions may also be important for invader establishment and success. Furthermore, most tests of local enemy release have been conducted over a single year even though release may be a transient phenomenon, especially at small scales. In this study, we used a multi-year field experiment to investigate whether locally isolated populations of the noxious non-native weed Cirsium arvense benefit from reduced levels of aboveground damage, and whether any initial advantage is lost over subsequent growing seasons. Populations of C. arvense were grown in plots at set distances from established source populations for 4years. In the first year of the experiment, folivory significantly declined with host isolation, but damage from specialist stem gallers and seed predators did not. However, in subsequent years of observation, folivores began colonizing isolated C. arvense populations while stem gallers exhibited very slow colonization of more isolated plots; seed predation showed no pattern with distance in any year. Local enemy escape did not result in increased plant performance, which instead negatively correlated with degree of isolation. Nonetheless, our results stress the importance of multi-year observations in tests of enemy release, since the herbivory patterns initially observed often changed within subsequent years depending on the dispersal ability and biology of the causal organism involved.

Spatial factors of white-tailed deer herbivory assessment in the central Appalachian Mountains.

Because moderate to over-abundant white-tailed deer (Odocoileus virginianus) herbivory impacts biodiversity and can alter community function, ecological benchmarks of herbivory impact are needed to assess deer impacts. We evaluated spatial patterns of deer herbivory and their relation to herbivory assessment by evaluating woody vegetation along 20 transects at each of 30 sites spread across a wide range of deer herd densities and vegetative condition throughout the biodiverse Appalachian Mountains of Virginia, USA. Surprisingly, herbivory patterns and the availability of woody forage generally were unchanged among physiographic regions and land use diversity classes. However, some relationships between browsing pattern and vegetation varied with scale. The total quantity of vegetation browsed on a given site and at the transect scale were related positively to the availability of forage, as the proportion of stems browsed decreased as stem density increased. However, this was only true when all stems were considered equally. When stem densities by species were weighted for deer preference, the proportion of stems browsed had no relationship or increased with stem density. Compared to the value from all transects sampled, on average, the mean of ≥ 3 transects within a site was within 0.1 of the browsing ratio and stem densities were within 0.5 stems m-2. Our results suggest that one transect per square kilometer with a minimum of three transects may be sufficient for most browsing intensity survey requirements to assess herbivory impacts in the Appalachian region of Virginia. Still, inclusion of spatial factors to help partition variation of deer herbivory potentially may allow for improved precision and accuracy in the design of field herbivory impact assessment methods and improve their application across various landscape contexts.