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

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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.     

The influence of deer browsing on the reproductive biology of Canada yew (Taxus canadensis marsh.)

Summary Browsed Canada yew (Taxus canadensis) populations have a higher proportion of males and a lower proportion of monoecious plants than unbrowsed yew populations. The proportion of monoecious plants increases with time following protection from browsing suggesting that deer browsing causes male-biased sex expression in Canada yew. In contrast, results from comparing browsed and unbrowsed populations, exclosure studies, and browse simulation experiments indicate that strobilus ratios and phenotypic gender of browsed yews may be female-biased. In part, these results correspond to the influence of size on sex expression in Canada yew; small yews tend to be male, but if monoecious, have female-biased strobilus ratios. Large yews are monoecious, but have male-biased strobilus ratios. There is, however, no consistent relationship between size and gender in Canada yew, suggesting that in some circumstances, yews shift allocation to female function in response to browsing.     

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.     

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.     

Indirect interactions between browsers and seed predators affect the seed bank dynamics of a chaparral shrub

Interactions between herbivores and seed predators may have long-term consequences for plant populations that rely on persistent seed banks for recovery after unpredictable fires. We assessed the effects of browsing by deer and seed predation by rodents, ants and birds on the densities of seeds entering the seed bank of Ceanothus cuneatus var. rigidus, a maritime chaparral shrub in coastal California. Ceanothus produced many more seeds when protected from browsers in long-term experimental exclosures than did browsed plants, but the seed densities in the soil beneath browsed and unbrowsed Ceanothus were the same at the start of an intensive one-year study. The density of seeds in the soil initially increased in both treatments following summer seed drop: while densities returned to pre-drop levels within a few weeks under browsed plants, soil seed densities remained high for 5–8 months beneath unbrowsed plants. Rodent abundance (especially deer mice) was higher near unbrowsed plants than >30 m away, and rodents removed Ceanothus seeds from dishes in the experimental plots. At least in the short term, rodent density and rates of seed removal were inversely related to the intensity of browsing. Our data have management implications for maintaining viable Ceanothus populations by regulating the intensity of browsing and the timing, intensity and frequency of fires.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Leaf miners on Ochna ciliata (Ochnaceae) growing on Aldabra Atoll: patterns of herbivory in relation to goat browsing and exposure to the sun

Abstract.

• 1 The patterns of herbivory by leaf miners on the shrub Ochna ciliata growing on Aldabra Atoll were studied in relation to browsing by feral goats and exposure to the tropical sun versus shading.
• 2 Damage due to both tunnelling (Diptera) and ‘blotch-making’ (Lepidoptera) leaf miners was statistically greater on trees that had been browsed and on those that received more extensive exposure to the sun.
• 3 Effects of exposure and browsing were additive, as evidenced by the lack of statistical interaction between these factors.
• 4 There was a significant negative correlation between the numbers of tunnelling and blotch-making miners in trees that had been browsed, but not in unbrowsed trees.
• 5 Three geographical areas were sampled to test whether the effects of browsing on leaf mining were invariant or influenced by as yet unknown environmental factors. A statistically significant interaction between browsing and geographical region indicated that, while browsed trees had significantly more mines whatever the habitat, the extent of this effect differed across the habitats.

     

Differential response of trees and shrubs to browsing and pruning: the effects on Pistacia growth and gall-inducing aphids

The effects of long term browsing on growth and morphology of P. atlantica trees and P. palaestina shrubs, and on the colonization of these hosts by their respective gall-inducing aphid guilds, were studied in natural vegetation pastures in Israel. To simulate apical dominance release by browsing, trees and shrubs were pruned and observed one year later. P. atlantica responded strongly to browsing, producing a crown of dense and sharp dry branches and thorn-like shoots, and pruning, compensating or overcompensating for the lost biomass in the studied morphological variables. Morphological responses of P. palaestina were less evident: we found no compensation in most variables. Three species of aphids produced more galls on browsed than on control shoots of P. atlantica. Two species induced more galls on pruned shoots. The five aphid species on P. palaestina colonized browsed and unbrowsed shoots equally, and produced fewer galls on pruned shoots. The role of apical dominance in architectural responses of trees and shrubs to browsing and pruning, as related to resource availability and timing, is discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

How Herbivore Browsing Strategy Affects Whole-Plant Photosynthetic Capacity

If a browse damage index indicates that a tree has been 50% browsed by herbivores, does this mean half the leaves are entirely eaten or are all the leaves half eaten? Were the affected leaves old or young? Large or small? In sunshine or shade? Understanding what effect browsing will have on the photosynthetic capacity and the plant’s survival ability clearly requires a greater understanding of browsing strategy across the canopy than can be given by a single index value. We developed stochastic models of leaf production, growth and consumption using data from kamahi (Weinmannia racemosa) trees in New Zealand which have been browsed by possums (Trichosurus vulpecula), to ascertain which of six feasible browsing strategies possums are most likely to be employing. We compared the area distribution of real fallen leaves to model output in order to select the best model, and used the model to predict the age distribution of leaves on the tree and thus infer its photosynthetic capability. The most likely browsing strategy that possums employ on kamahi trees is a preference for virgin (i.e. previously unbrowsed) leaves, consistent with the idea that browsing increases the production of chemical plant defences. More generally, our results show that herbivore browsing strategy can significantly change the whole-plant photosynthetic capability of any plant and hence its ability to survive, and therefore, herbivore damage indices should be used in conjunction with more detailed information about herbivore browsing strategy.     

Herbivores Influence the Growth,Reproduction, and Morphology of a Widespread Arctic Willow

Shrubs have expanded in Arctic ecosystems over the past century, resulting in significant changes to albedo, ecosystem function, and plant community composition. Willow and rock ptarmigan (Lagopus lagopus, L. muta) and moose (Alces alces) extensively browse Arctic shrubs, and may influence their architecture, growth, and reproduction. Furthermore, these herbivores may alter forage plants in such a way as to increase the quantity and accessibility of their own food source. We estimated the effect of winter browsing by ptarmigan and moose on an abundant, early-successional willow (Salix alaxensis) in northern Alaska by comparing browsed to unbrowsed branches. Ptarmigan browsed 82–89% of willows and removed 30–39% of buds, depending on study area and year. Moose browsed 17–44% of willows and browsed 39–55% of shoots. Browsing inhibited apical dominance and activated axillary and adventitious buds to produce new vegetative shoots. Ptarmigan- and moose-browsed willow branches produced twice the volume of shoot growth but significantly fewer catkins the following summer compared with unbrowsed willow branches. Shoots on browsed willows were larger and produced 40–60% more buds compared to unbrowsed shoots. This process of shoot production at basal parts of the branch is the mechanism by which willows develop a highly complex “broomed” architecture after several years of browsing. Broomed willows were shorter and more likely to be re-browsed by ptarmigan, but not moose. Ptarmigan likely benefit from the greater quantity and accessibility of buds on previously browsed willows and may increase the carrying capacity of their own habitat. Despite the observed tolerance of willows to browsing, their vertical growth and reproduction were strongly inhibited by moose and ptarmigan. Browsing by these herbivores therefore needs to be considered in future models of shrub expansion in the Arctic.     

The influence of deer browsing on the reproductive biology of Canada yew (Taxus canadensis marsh.)

Summary Canada yew (Taxus canadensis) populations currently browsed by white-tailed deer (Odocoileus virginianus) or browsed by deer in the past had significantly lower production of male strobili, female strobili, and seeds than unbrowsed yew populations. Exclosure studies showed that protected yews produced significantly more male and female strobili than unprotected yews, but only after several years of protection. Seed production did not respond as readily to protection from deer perhaps because of reduced pollination levels in browsed yew populations. Previously unbrowsed yews were clipped at different levels of removal of available browse (control (no removal), 25%, 50%, 75%, and 100% removal) to simulate deer browsing. Reduction in male strobilus production was linearly related to clipping intensity in three years of observation. Female strobilus production was significantly reduced only at the 100% level of removal. Intermediate levels of clipping may have even stimulated production of female strobili. Analysis of covariance, with previous year's branch production as the covariate, showed no significant effect of clipping on male strobilus production except in the 100% removal group. Female strobilus production showed no such covariance with branch production. Effects of clipping on seed production could not be reliably assessed in 1984 and 1985 due to low seed production. Seed production in 1986 was significantly reduced only in the 100% removal group. Field observations of deer browsing of Canada yew indicate that 100% levels of removal are typical of natural levels of browsing.     

The induction of spinescence in European holly leaves by browsing ungulates

Evidence is presented which suggests that the spinescence of leaves of European holly, Ilex aquifolium, deters feeding by ungulates and is induced by browsing. Spinescence decreased as leaf size increased; hence, spinescence may be achieved by reducing adult leaf size. Holly shrubs with very spiny leaves were browsed less often than less spiny shrubs. In the absence of browsing ungulates during a one year period, the spinescence of leaves of holly shrubs significantly decreased. Browsed shrubs exhibited reduced annual shoot growth, increased branching, and produced smaller leaves with high spinescence. The regrowth on browsed branches of holly trees was characterized by increased leaf spinescence relative to unbrowsed branches. Hence, the induced response was localized, thereby reducing the ability of browsing ungulates to exert selective pressures on holly trees.     

Effects of competition,herbivory and substrate disturbance on growth and size structure in pin cherry (Prunus pensylvanica L.) seedlings

We examined separate and interactive effects of intraspecific competition, vertebrate browsing and substrate disturbance on the growth and size structure of pin cherry (Prunus pensylvanica L.) in the first two seasons of growth after clearcutting, in a hardwoods forest in New Hampshire, United States. Over the 15-month study period, 97.5% of 1801 individuals survived, and mean plant height increased from 4-fold at high density to 5-fold at low density. Relative height growth was significantly lower at higher plant densities in two of the three growth periods examined. Vertebrate browsers (moose and deer) significantly preferred taller plants. Browsed plants had higher relative height growth following browsing than unbrowsed plants (compensatory growth) at low and intermediate densities. The degree of compensation declined with density and compensation was not significant at the highest density level. At low and intermediate densities, plants browsed early in life regained height dominance through compensatory growth; they failed to regain dominance at high density. Because compensatory growth tended to offset the effects of size-selective browsing, there was no difference in the degree of size inequality between browsed and unbrowsed plots. However, size inequality increased with plant density. Substrate disturbance caused by logging had no significant effects on either relative height growth or size inequality. The slope of the relationship between relative height growth and initial height increased significantly with density and time, and was higher in unbrowsed than in browsed plots, suggesting that competition among plants was size-asymmetric. Despite the preference of browsers for large plants, there was a clear net growth advantage for plants of large initial size, when the effects of competition, browsing and compensatory growth were combined. The interactive effects of density and browsing demonstrate the importance of a multifactorial approach to the analysis of individual plant performance and population structure.     

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.     

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.     

Winter browsing on Alaska feltleaf willow twigs improves leaf nutritional value for snowshoe hares in summer

In boreal forests, browsing by mammals on winter-dormant twigs increases leaf nitrogen, leaf greenness, and leaf size. This suggests browsing reduces competition among meristems for mineral nutrients, and in particular, competition for nitrogen. Winter browsing also reduces the shoot carbohydrate reserves used by leaves to produce condensed tannin. These effects of winter browsing are predicted to improve the nutritional value of leaves for mammals because they increase the mass of digestible nitrogen in leaves. This hypothesis was tested using Alaska feltleaf willow and the snowshoe hare as the experimental system. Six in vivo indicators of leaf nutritional quality were used to compare leaves from winter-browsed plants with leaves from unbrowsed plants. The indicators used were dry matter intake, nitrogen intake, condensed tannin intake, dry matter digestibility, apparent digestibility of nitrogen and nitrogen retention. The results obtained were in agreement with the above hypothesis. In early summer, at the time snowshoe hares and other northern herbivores reproduce, hares fed leaves from browsed plants consumed more nitrogen, digested more of the nitrogen they consumed, and retained more of the nitrogen they digested than did hares fed leaves from unbrowsed plants. The high nitrogen content and low tannin content of leaves from browsed plants may explain this browsing caused increase in leaf nutritional value. How these positive effects of winter browsing on snowshoe hare nutrition at the time of reproduction might affect hare population dynamics are briefly discussed.     

Long-term impacts of goat browsing on bush-clump dynamics in a semi-arid subtropical savanna

The effects of 16 years of continuous browsing by goats in a South African savanna at stocking rates intended for bush control were compared with plots unbrowsed for the same period of time. Differences in bush-clump density, structure and species composition were recorded. Bush-clump density did not differ between browsed and unbrowsed plots. Within individual bush-clumps, browsing was shown to impact more on structure than species composition, with smaller, shorter bush-clumps, containing fewer species but much greater stem-densities. Although species presence/absence was little affected by browsing, many species showed differences in abundance, growth and location within browsed and unbrowsed bush-clumps. Species reduced in abundance in browsed plots included Cussonia spicata, Ehretia rigida, Grewia occidentalis, Jasminum angulare and Senecio linifolius. Several species showed reduced growth in browsed plots, particularly those located at bush-clump edges. The relatively unpreferred Aloe ferox was a notable exception. Although browsing had little effect on the composition of the main clump founding species, emergents or late arrivals, there were twice as many single plants in browsed plots and emergence of several species was restricted to the middle of bush-clumps. Comparison of our findings with aerial photographic evidence and other literature suggest that browsing alone is unlikely to significantly reduce scrub cover, although it can clearly control further expansion. Combinations of fire and browsing, rather than one factor alone, are considered likely to act fastest and most effectively to significantly reduce or remove scrub cover altogether.     

Interactions among elk, aspen, galling sawflies and insectivorous birds

Using two years of observational and experimental data, we examined the hypothesis that browsing by elk on aspen indirectly affects the distribution of a leaf-galling sawfly, which in turn affects insect diversity and foraging patterns of insectivorous birds. We found that: i) in an analyses of 33 arthropod species, the presence of sawflies significantly increased arthropod richness and abundance by 2 X and 2.5 X, respectively. ii) browsing by elk reduced sawfly gall abundance such that 90% of the galls were found on unbrowsed aspen ramets. iii) insectivorous birds attacked 60–74% of the galls on unbrowsed shoots compared to 11% on browsed shoots. When leaf-galler abundance was experimentally held constant on browsed and unbrowsed shoots, predation by insectivorous birds did not differ significantly. This result suggests that browsing affects the patterns of avian predation by altering the distribution of a galling insect. These data argue that bottom-up, top-down, and lateral factors can act in concert to affect the distribution of a galler, structure arthropod communities and affect predation by insectivorous birds.     

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.     

Long‐term response of temperate canopy trees to removal of browsing from an invasive arboreal herbivore in New Zealand

Defoliation of forest tree canopies by herbivores and other agents, leading to tree mortality and reduced productivity, threatens the ecological stability of forests globally. This study shows that long‐term control of a mammalian arboreal folivore (brushtail possums; Trichosurus vulpecula Phalangeridae) reduces crown dieback and increases foliage cover in browsing‐damaged canopy trees. We monitored indices of possum density, possum browsing, tree foliage cover and crown dieback for 20 years following initiation of possum control in 1994 that repeatedly reduced possum densities to near zero every 5–6 years and kept the population below 35% of pre‐control levels over the entire period. Observable possum browsing was recorded on 20–49% of individuals of three palatable tree species at the time of first control. Those percentages fell to zero after control and never exceeded 2–10% for individual species over the next 19 years. We recorded significant increases in foliage cover attributable to recovery from defoliation by possums for all three species during the first 10 years. Large increases in foliage cover occurred on individuals that were heavily browsed in 1994 (mean increases: 36–89%), but mean population increases were modest (3–19%) because only 10–19% of trees were initially heavily browsed. Twenty‐year mortality rates were similar for plants with, or without, initial possum browsing, indicating no residual impact of pre‐control browsing on tree mortality. Times for full recovery of crown foliage cover varied from 10 years for the youngest trees and faster growing species to more than 20 years for mature individuals of the slowest growing species.