Is escaping deer browse just a matter of time in Picea sitchensis? A chemical and dendroecological approach

We combined chemical and dendroecological analyses to understand the mechanisms that are involved in escaping deer browse by young Sitka spruce (Picea sitchensis) exposed to browsing by Sitka black-tailed deer (Odocoileus hemionus sitchensis) on Haida Gwaii (British Columbia, Canada). We compared chemical defences (terpenes), nutritive compounds (nitrogen, non-structural constituents, cellulose, and lignin), as well as age and radial growth of two young spruce categories growing side by side: (1) stunted spruces that were heavily browsed, shorter than the browse line, and (2) escaped spruces that were taller than the browse line but still browsed below the browse line. Escaped and stunted spruces did not differ in terpene concentrations, or in nutritive compound contents, suggesting that they had similar palatability. Escaped spruces were older that stunted spruces. Stunted and escaped trees had similar slow growth when young, suggesting no difference in initial browsing between the two spruce categories. For escaped spruce, there was a dramatic increase in radial growth at about 12-13 years old, suggesting that the apex of the trees had escaped deer browse. Because the two categories of spruces were equally accessible and did not differ in chemical defences or in nutritive compounds, and because escaped spruces were older than stunted trees and had a similar slow radial growth in their first 12-13 years, we conclude that morphological differences between stunted and escaped browsed trees are due to age and that it is only a matter of time before spruce escape deer on Haida Gwaii.     

Linking deer browsing and terpene production among genetic identities in Chamaecyparis nootkatensis and Thuja plicata (Cupressaceae)

To investigate whether differential herbivore browsing reflects genetic variation in plant defense expression, variation in needle terpenes and damage caused by black-tailed deer (Odocoileus hemionus) was analyzed on yellow-cedar (Chamaecyparis nootkatensis) and western redcedar (Thuja plicata). In a 100-genet yellow-cedar population, three genets that were heavily browsed and had extremely low levels of monoterpenes (0-0.36% dry matter), sesquiterpenes, and diterpenes were compared to unbrowsed genets (0.85-3.83% monoterpenes in dry matter). These differences were maintained in individuals protected from browsing, suggesting genetically based variation in constitutive terpene production. In western redcedar, heavily browsed trees had significantly lower total monoterpene concentrations (1.69% dry matter) than lightly browsed trees (3.32% dry matter). One heavily browsed tree expressed no monoterpenes. No differences were found for diterpenes. In both species, the genotypes with extremely low monoterpene concentrations came from the same open-pollinated families.     

Disentangling the causes of damage variation by deer browsing on young Thuja plicata

Long‐lived trees experience different levels of damage due to mammalian herbivores. To untangle the mechanisms that underlie this variation, we combined chemical with dendrochronological analyses to study variation in browsing on Western redcedars (Thuja plicata) on Haida Gwaii (British Columbia, Canada). Since the last glaciation, Haida Gwaii forests had lacked large herbivore browser until Sitka black‐tailed deer (Odocoileus hemionus sitkensis) were introduced at the beginning of the 20th century. Dendrochronology yielded information on radial growth and plant annual responses to environmental stresses including herbivory. Secondary metabolite content and plant nutritional quality provided insights into proximate causes of food choices made by herbivores. We sampled lightly‐ and heavily‐browsed young trees at four sites: three clear‐cut sites with high browsing pressure and one old‐growth forest site where browsing pressure had, until recently, been lower. Heavily‐browsed young trees had lower concentrations of secondary metabolites and were of lower nutritive value than lightly‐browsed trees at all sites. Under high browsing pressure, tree growth patterns suggested that all young trees were initially severely browsed until some trees, currently scored as lightly‐browsed, started to escape deer. At the old‐growth site, both lightly‐ and heavily‐browsed trees tended to have lower overall average secondary metabolite concentrations than those of all other sites, a trend possibly related to greater canopy closure. Lightly‐browsed trees were older than heavily‐browsed ones which resulted, during the period of lower browsing pressure, in higher growth rate and a same pattern of change in growth from one year to the next year. This suggests that, under low browsing pressure, selection of young trees related to chemical defense was weak and that growth differences due to other factors than browsing could be expressed. Under strong browsing pressure, however, all young trees had equally low growth rates until trees with better genetic potential to produce effective defenses were able to escape deer. This suggests that selection by deer could occur on a long‐lived tree.     

Influence of herbivores on a perennial plant: variation with life history stage and herbivore species

Herbivores have diverse impacts on their host plants, potentially altering survival, growth, fecundity, and other aspects of plant performance. Especially for longer-lived plant species, the effects of a single herbivore species can vary markedly throughout the life of the host plant. In addition, the effects of herbivory during any given life history stage of a host plant may also vary considerably with different types of herbivores. To investigate the effects of herbivory by black-tailed deer (Odocoileus hemionus columbianus) and snails (Helminthoglypta arrosa and Helix aspersa) on a nitrogen-fixing shrub, Lupinus chamissonis, we established three exclosure experiments in a sand dune system on the coast of northern California. These experiments documented that deer browsing significantly reduced the volume and growth rate of lupines in the seedling and juvenile life stages. Since plant volume was strongly correlated with aboveground dry biomass for lupines, such herbivore-induced reductions in volume should translate into losses of aboveground biomass. Deer browsing also significantly altered the likelihood of attack by and density of a leaf-galling cecidomyid fly (Dasineura lupinorum), suggesting that a vertebrate herbivore indirectly affected an invertebrate herbivore in this system. Although deer did not significantly affect the survival of lupine seedlings and juveniles, individuals protected from deer had consistently greater survival in the two separate experiments. Our results revealed that snails did not have a significant effect on the survival or growth of juvenile plants, despite being common on and around lupines. An exclosure experiment revealed that herbivory by deer significantly reduced the shoot lengths of mature shrubs, but led only to a minimal reduction in growth rates. In addition, we found that browsed shrubs had significantly greater inflorescence production, but also produced individual seeds with significantly reduced mass. Collectively, these data indicate that deer and snails have widely differing effects on their shared host plant; browsing by deer indirectly affects insect herbivores, and the impacts of deer change markedly with the life history stage of their host plant.     

Effects of Deer Browsing, Fabric Mats, and Tree Shelters on Quercus rubra Seedlings

We examined the effects of deer browsing, fabric mats, and tree shelters on the growth and survival of Quercus rubra (northern red oak) seedlings planted as part of a reforestation project in southcentral Minnesota. Browsing by white-tailed deer occurred on 68.6% of the seedlings growing without tree shelters, whereas only 3.6% of the trees were browsed by mice or voles and 2.5% were browsed by rabbits. Fabric mats, used to control competition from herbaceous plants for roughly half of the seedlings included in the study, had a detrimental effect overall. Seedlings grown with mats had a greater frequency of deer browsing and a greater chance of dying than seedlings grown without mats. Stem height for seedlings browsed by deer was less overall than for nonbrowsed seedlings, although this pattern varied with use of fabric mats and plot location. The use of plastic tree shelters effectively prevented deer browsing and reduced the mortality rate from 34.6% to 3.2%. Our results indicate that fabric mats should not be used in restoration projects with large deer populations. They also suggest that planting seedlings away from existing forest edges and using seedling protection measures such as tree shelters will increase seedling survival and growth in future restoration projects.     

Cumulative effects of wild ungulate and livestock herbivory on riparian willows

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

Response of young<Emphasis Type="Italic"> Tsuga heterophylla</Emphasis> to deer browsing: developing tools to assess deer impact on forest dynamics

We used dendroecology to describe and understand the consequences of deer browsing on regenerating western hemlock (Tsuga heterophylla). We compared tree shape, growth rate, height and age at four different sites in Haida Gwaii (British Columbia, Canada) that had trees representative of the range of deer impact on trees: (1) trees showing no sign of browsing, (2) escaped trees which were still browsed below the browse line and (3) stunted and heavily browsed trees. Repeated and intense browsing resulted in the small size, compact heavily ramified shape of stunted trees and in the short compact and ramified lower branches of escaped trees. These contrasted with the shape of non-browsed trees, a shape that was also found in escaped trees above the browse line. Before release, all browsed trees experienced stagnation in growth characterised by narrow rings (0.3 mm/year) and a small annual height increment (2.5 cm/year). At release, growth rate increased and stabilised: rings were wider (1.3 mm/year) and annual height increments were greater (10.5 cm/year). Non-browsed trees had a mean ring-width of 1.3 mm/year and an annual height increment of 22 cm/year. Delay in tree recruitment caused by deer varied from site to site. It had been about 15 years for the escaped trees and is estimated at 30–40 years for the stunted trees. Spatial variation in deer impact may reflect spatial variation of browsing pressure resulting from local differences in the availability of preferred forage or to differences in tree chemical defences/nutritional values.     

Trait‐mediated indirect interactions: Moose browsing increases sawfly fecundity through plant‐induced responses

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Foliar nutrients shape fungal endophyte communities in Western white pine (Pinus monticola) with implications for white-tailed deer herbivory

Asymptomatic fungal endophytes colonize tissues of woody plants worldwide, with largely unknown ecological effects. Using culture-based methods and ITS1-5.8S-ITS2 rDNA sequence analysis, we investigated differences between foliar endophyte communities in disease-resistant hybrid and wild-type Pinus monticola (Western white pine) trees with observed variation in tree growth, vigor, and browsing damage by white-tailed deer (Odocoileus virginianus). We isolated 69 phylotypes of endophytic fungi in at least 39 genera, including 26 that have not previously been reported in P. monticola. Principal components analysis revealed that endophyte communities differed between browsed seedlings, unbrowsed seedlings, and unbrowsed adult trees. Sulfur, nitrate and calcium concentrations correlated with endophyte community differences among tree groups based on a distance-based redundancy analysis. Our results indicate that foliar nutrient variation influences endophyte community assembly and deer herbivory in P. monticola on a small landscape scale (80 hectares).     

Trade-off between plant growth and defense? A comparison of sagebrush populations

We used ecotypic variation in big sagebrush (Artemisia tridentata) to examine potential trade-offs between inherent growth rate and tolerance or resistance to herbivory. Seeds were obtained from seven geographic populations, and 1,120 seedlings were established in a common garden. In one set of plots, plants were subjected to five treatments: control, regular insecticide spray, moderate browsing, severe browsing, or moderate browsing plus insecticide. Plants in a second set of plots were all untreated, and were used to estimate ambient growth, flower production, and susceptibility to herbivorous insects. In the first growing season, population differences in relative growth rate produced approximately seven-fold variation in mean biomass. Two populations of basin big sagebrush (A. tridentata tridentata) and one population of mountain big sagebrush (A. tridentata vaseyana) grew fastest; those of Wyoming big sagebrush (A. tridentata wyomingensis) showed the slowest growth. Bi-weekly application of insecticide for two growing seasons had no effect on the growth of either browsed or unbrowsed plants. All populations showed compensatory growth (but not overcompensation) in response to browsing, but the degree of compensation was unrelated to inherent growth rate. Similarly, there was no consistent relationship between plant growth rate and flower production in the second growing season. Some insects colonized fast-growing populations more frequently than slow-growing ones, but patterns of insect colonization were species-specific. At the level of geographic populations and subspecies, we found little evidence of a built-in trade-off between inherent growth rate and the ability to tolerate or resist herbivory. Because population ranks for growth rate changed substantially between seasons, attempts to correlate growth and defense characters need to account for differences in the growth trajectories of perennial plants.     

Ungulate browsing affects subsequent insect feeding on a shared food plant,bilberry (Vaccinium myrtillus)

Browsing by ungulates may induce plant responses and affect subsequent plant food quality for other animals. Populations of many deer species have increased to unprecedented levels in Europe and North America. In Norway, population densities of red deer (Cervus elaphus) have increased over the past decades, but little is known about how increased deer browsing pressure may change the palatability of key food plants for other taxa in the boreal ecosystem. We conducted a cafeteria experiment to assess if long-term deer-browsing intensity affected the palatability of bilberry (Vaccinium myrtillus) leaves for leaf-eating larvae (mainly Lepidoptera). We found that leaf-eating insect larvae preferred bilberry leaves from the lightly browsed bilberry plants; the larvae consumed twice as much leaf biomass from the lightly browsed plants than from the unbrowsed and moderately browsed ones, and four times more than from highly browsed plants. Larvae never selected leaves from highly browsed plants as their first choice. Our study suggests that browsing-induced changes in the quality of shared food plants may be important in mediating indirect interactions between browsers of widely separated taxa. Whereas low levels of long-term red deer browsing increases the palatability of bilberry leaves for leaf-eating larvae, high browsing pressure reduces food consumption. Whether changes in palatability lead to changes in population densities of leaf-eating larvae remains to be studied, but any such adverse effects could have cascading ecological consequences for insectivorous birds and mammals.     

Different facets of tree sapling diversity influence browsing intensity by deer dependent on spatial scale

Browsing of tree saplings by deer hampers forest regeneration in mixed forests across Europe and North America. It is well known that tree species are differentially affected by deer browsing, but little is known about how different facets of diversity, such as species richness, identity, and composition, affect browsing intensity at different spatial scales. Using forest inventory data from the Hainich National Park, a mixed deciduous forest in central Germany, we applied a hierarchical approach to model the browsing probability of patches (regional scale) as well as the species‐specific proportion of saplings browsed within patches (patch scale). We found that, at the regional scale, the probability that a patch was browsed increased with certain species composition, namely with low abundance of European beech (Fagus sylvatica L.) and high abundance of European ash (Fraxinus excelsior L.), whereas at the patch scale, the proportion of saplings browsed per species was mainly determined by the species’ identity, providing a “preference ranking” of the 11 tree species under study. Interestingly, at the regional scale, species‐rich patches were more likely to be browsed; however, at the patch scale, species‐rich patches showed a lower proportion of saplings per species browsed. Presumably, diverse patches attract deer, but satisfy nutritional needs faster, such that fewer saplings need to be browsed. Some forest stand parameters, such as more open canopies, increased the browsing intensity at either scale. By showing the effects that various facets of diversity, as well as environmental parameters, exerted on browsing intensity at the regional as well as patch scale, our study advances the understanding of mammalian herbivore–plant interactions across scales. Our results also indicate which regeneration patches and species are (least) prone to browsing and show the importance of different facets of diversity for the prediction and management of browsing intensity and regeneration dynamics.     

Feeding patterns by roe deer and rabbits on pine, willow and birch in relation to spatial arrangement

The location of a plant and its association with the surrounding vegetation may be a strong determinant of herbivore foraging decisions. The attractiveness of a food plant may be reduced if associated with a less preferred one (repellent–plant hypothesis) or, conversely, it may escape herbivory via association with a more preferred species (attractant–decoy hypothesis). In this study we tested the hypothesis that selection of the same food plants by two herbivore species with different body mass, i.e. roe deer and rabbits, is affected by the spatial disposition of preferred plants in relation to less preferred ones. We planted willows, birches and pines, representing food species of higher, intermediate and lower preference, respectively, in different spatial arrangements, to manipulate patch quality and accessibility to herbivores. Contrasting patches were constructed by planting willows or pines in the midst of birches and single-species arrays of all three species, in a blocked design, replicated six times in an area occupied by both roe deer and rabbits. Preference patterns were studied by recording browsing on current years' shoots and older plant parts. Across all trees in the experiment, we confirmed that roe deer clearly preferred willow over birch, pine was not browsed at all. There was greater herbivory by roe deer on birches occurring peripherally in an array, when the associated central species was willow or pine, as compared to birches in a single-species birch patch. Therefore food choices by roe deer appear to be determined by both patch and individual level selection. The rabbits only showed preference at a species level and did not respond to spatial arrangement of trees. The results from this study indicate that the location and association of trees can strongly affect foraging patterns of roe deer, but not in a manner wholly consistent with the repellent–plant or the attractant–decoy hypotheses.     

Feeding behaviour of Red Deer (Cervus elaphus) offered Sitka Spruce saplings (Picea sitchensis) grown under different light and nutrient regimes

1. In addition to exhibiting preferences for particular plant species, vertebrate herbivores select particular individuals of these species whilst leaving others undamaged. This pattern of diet selection may reflect differences in the chemical composition (and hence nutritional quality) between individual plants, and/or variability in the physical constraints on intake rate, such as plant structure.
2. An experiment was conducted to test the effects of environmental manipulations on the morphology and chemical composition of Sitka Spruce saplings, and to evaluate the consequences for herbivory by Red Deer. Fertilizing the trees increased tree height, branch span and leader length, twig width and needle width, and decreased the concentrations of total phenolics, condensed tannins, fibre and lignin but monoterpene content was not altered. Shading also reduced phenolic and tannin concentrations.
3. When the fertilized and shaded trees were offered to deer in feeding trials, the probability of a tree being visited by a deer and the biomass removed were influenced by tree morphological variables, as were the bite rate and intake rate of the deer. More biomass was removed from larger trees.
4. Once the effects of tree morphology had been taken into account, there was no effect of the fertilizer and shade treatments on deer browsing behaviour that could be attributed to changes in chemical composition of the trees. The relationship between intake rate and bite size at each tree varied between individual deer, but the functional response relationship between intake rate and bite size was not influenced by the treatments applied to the trees.
5. Tree morphology may have a larger influence on deer feeding behaviour than tree chemical composition.     

Chemical and morphological variation of Mediterranean woody evergreen species: Do plants respond to ungulate browsing?

Abstract. Plant defense theories suggest that chemical or structural defences should be maximized when and where browsing is most likely to occur. We tested this hypothesis on four evergreen woody species growing in a Mediterranean area with a high density of ungulates. In this system, levels of browsing are more intense in the winter (due to the lack of annual plants) and young foliage is often preferred. Therefore we predicted that the chemical defences of these species, namely their phenolic content, would vary with leaf age, season and damage intensity. In addition, we tested whether ungulates preferentially selected species containing lower phenolic levels, and also whether browsing induced either chemical or morphological changes in damaged plants. Phenolic levels varied greatly between plant species; ungulates browsed preferentially on the species with the lowest phenolic levels. No difference in phenolic content was found between browsed and unbrowsed trees. Morphological changes in heavily browsed trees included an increase in shoot and leaf density and a net decrease in leaf size. We suggest that for Mediterranean plants, which have evolved under high browsing pressure from large mammals, the production of small leaves and dense shoots in response to browsing might decrease ungulate foraging efficiency and hence reduce the rate of further damage as effectively as high levels of chemical defence.     

Secondary metabolites and nutrients of woody plants in relation to browsing intensity in African savannas

Carbon-based secondary metabolites (CBSMs) are assumed to function as defences that contribute to herbivore-avoidance strategies of woody plants. Severe browsing has been reported to reduce concentrations of CBSMs and increase N concentrations in individual plants, causing heavily browsed plants to be characterised by N-rich/C-poor tissues. We hypothesised that concentrations of condensed tannins (CT) and total polyphenols (TP) should decrease, or N increase, in relation to increasing intensity of browsing, rendering severely browsed plants potentially more palatable (increased N:CT) and less N-limited (increased N:P) than lightly browsed ones. We sampled naturally browsed trees (taller than 2 m) of four abundant species in southern Kruger National Park, South Africa. Species-specific relationships between N:CT, CT, TP and P concentrations and increasing browsing intensity were detected, but N and N:P were consistently invariable. We developed a conceptual post-hoc model to explain diverse species-specific CBSM responses on the basis of relative allocation of C to total C-based defence traits (e.g. spines/thorns, tough/evergreen leaves, phenolic compounds). The model suggests that species with low allocation of C to C-based defence traits become C-limited (potentially more palatable) at higher browsing intensity than species with high allocation of C to C-based defences. The model also suggests that when N availability is high, plants become C-limited at higher browsing intensity than when N availability is low.     

Browsing in a heterogeneons savanna

Large herbivores generally depend on and interact with a food resource that is heterogeneous at different spatial scales. Plants allocate resources to rapid growth or to defence mechanisms depending on the availability of resources relative to loss of resources from herbivory. Herbivores select food and feeding habitats in order to maximize intake rate of nutrients and digestible energy, while avoiding chemical and structural deterrents. To optimize foraging, herbivores select habitats and food items in a hierarchical way, and different attracting and deterring factors may govern selection at different scales. We studied the impact of twig biting by a guild of indigenous browsers in three vegetation types in a semi-arid savanna in Botswana. The heaviest browsing pressure was in the vegetation type richest in preferred plant species, although that type was also richest in defended species. There were large differences in relative utilization between plant species, and ranking of species was roughly similar in the different vegetation types. Browsing pressure varied between species from almost 0-30%. Overall, spinescent trees were less browsed than non-spinescent ones, and evergreen species were less browsed than deciduous ones. In two of the three vegetation types there was a negative correlation between browsing pressure on a species and its frequency. There was a high incidence of rebrowsing, and once a tree had been browsed, the probability that it would be browsed again increased. The results largely agree with predictions based on the resource availability hypothesis, the scarcity accessibility hypothesis and recent theories on the significance of plant defences and on plant's response to browsing and the subsequent response by herbivores on the plant's responses.     

Increased thorn length in Acacia depranolobium —an induced response to browsing

Summary I report here longer thorns induced by large mammal herbivory on the tree Acacia depranolobium. I compared trees that had been browsed by domestic goats to trees protected from goat browsing. Thorns on browsed branches within the reach of goats (<125 cm above the ground) were significantly longer than thorns from higher branches on the same browsed trees, and significantly longer than branches at similar heights on unbrowsed trees. It appears that increased thorn length was an induced response to large mammal herbivory in Acacia depranolobium, both among and within individual trees.     

Can browsing by deer in winter induce defence responses in bilberry (<Emphasis Type="Italic">Vaccinium myrtillus</Emphasis>)?

Wild ungulates are key determinants in shaping boreal plant communities, and may also affect ecosystem function through inducing the plant defence systems of key plant species. We examined whether winter browsing by deer could increase the resistance of bilberry (Vaccinium myrtillus). We used three indicators of induced bilberry defence: reduced growth (a), reduced reproduction (b) and decreased insect herbivory (c) in focal plants. In a field experiment, using a randomised block design, we exposed half of plants twice in winter to exogenously applied methyl jasmonate (MeJA) and crossed this factor with randomly selecting browsed and unbrowsed plants. We predicted that MeJA-plants would have significant lower growth, reproduction and insect herbivory than Control plants. We also expected that Browsed plants would experience similar negative effects and that there would be an interaction between MeJa and Browsed indicating a possible additive effect. Growth, flowering and insect herbivory were significantly lower in MeJA than in Control, as expected. We did not find the same reduction for Browsed and no significant interaction between factors. The combined treatment, unexpectedly, flowered more and showed higher levels of insect herbivory than MeJA. Our study showed that defence responses of bilberry may be induced by exogenously-applied MeJA in winter. Our study could not confirm whether winter browsing by deer can induce the same defence responses.     

Environmental context determines within- and potential between-generation consequences of herbivory

Plant tolerance to herbivory may depend on local environmental conditions. Models predict both increased and decreased tolerance with increasing resources. Transgenerational effects of herbivory may result in cross-generation tolerance. We evaluated within- and potential between-generation consequences of deer browsing in light-gap and understory habitats in the forest-edge herb, Campanulastrum americanum. Plants were assigned to deer-browsed, simulated-herbivory, and control (undamaged) treatments in the two light environments. In light gaps, plants were eaten earlier, more frequently, and had less vegetative recovery relative to uneaten plants than in the understory. As a result, browsed light-gap plants had a greater reduction in flowers and fruit than understory plants. This reduced tolerance was in part because deer browsing damaged plants in light gaps more than those in the understory. However, in the simulated herbivory treatment, where damage levels were similar between light habitats, plants growing in high-resource light gaps also had reduced tolerance of herbivory relative to those in the forest understory. C. americanum’s reproductive phenology was delayed by reduced light and the loss of the apical meristem. As a result, deer-browsed plants in the light gap flowered slightly later than uneaten plants in the understory. C. americanum has a polymorphic life history and maternal flowering time influences the frequency of annual and biennial offspring. The later flowering of deer-browsed plants in light gaps will likely result in a reduced frequency of high-fitness annual offspring and an increase in lower fitness biennial offspring. Therefore, additional between-generation costs of herbivory are expected relative to those predicted by fruit number alone.