Tree species diversity influences herbivore abundance and damage: meta-analysis of long-term forest experiments

Plant monocultures are commonly believed to be more susceptible to herbivore attacks than stands composed of several plant species. However, few studies have experimentally tested the effects of tree species diversity on herbivory. In this paper, we present a meta-analysis of uniformly collected data on insect herbivore abundance and damage on three tree species (silver birch, black alder and sessile oak) from seven long-term forest diversity experiments in boreal and temperate forest zones. Our aim was to compare the effects of forest diversity on herbivores belonging to different feeding guilds and inhabiting different tree species. At the same time we also examined the variation in herbivore responses due to tree age and sampling period within the season, the effects of experimental design (plot size and planting density) and the stability of herbivore responses over time. Herbivore responses varied significantly both among insect feeding guilds and among host tree species. Among insect feeding guilds, only leaf miner densities were consistently lower and less variable in mixed stands as compared to tree monocultures regardless of the host tree species. The responses of other herbivores to forest diversity depended largely on host tree species. Insect herbivory on birch was significantly lower in mixtures than in birch monocultures, whereas insect herbivory on oak and alder was higher in mixtures than in oak and alder monocultures. The effects of tree species diversity were also more pronounced in older trees, in the earlier part of the season, at larger plots and at lower planting density. Overall our results demonstrate that forest diversity does not generally and uniformly reduce insect herbivory and suggest instead that insect herbivore responses to forest diversity are highly variable and strongly dependent on the host tree species and other stand characteristics as well as on the type of the herbivore.     

Mistletoe host-resemblance: A study of herbivory,nitrogen and moisture in two Australian mistletoes and their host trees

Abstract For 150 years mistletoe host-resemblance has been an unsolved puzzle. Mimicry, camouflage, host protection and shape modification by the host tree have all been advanced as possible solutions. No extended examination of herbivory of host-parasite pairs has ever been done, however, to put these explanations to the test. The study was carried out in northeastern Australia from March to July 1994. Rates of leaf herbivory were estimated for seven individuals of Amyema biniflora Barlow (a cryptic mistletoe species), Dendrophthoe glabrescens (Blakely) Barlow (a non-cryptic mistletoe species) and their host trees (Eucalyptus tessellaris F. Muell. and Eucalyptus platyphylla F. Muell., respectively). In addition three measures of leaf palatability–nitrogen content, moisture content and toughness–were also assessed. Variability in mistletoe leaf shape was quantified by measuring the leaf widths of mistletoes on a variety of host tree species. Mistletoes sustained greater levels of herbivory compared to their host trees, but herbivory did not differ between mistletoe species. The non-cryptic mistletoe had lower levels of nitrogen compared to its host tree, but there was no difference in nitrogen levels between the cryptic mistletoe and its host. The moisture content of mistletoe leaves was greater than that of their hosts but not between mistletoe or host species. The mistletoe species had tougher leaves than their host trees. Leaf shape was different for one species of mistletoe growing on different host trees, but constant for another species of mistletoe. The results contradict, in some crucial aspect, all of the mimicry hypotheses currently on offer.     

Sapling herbivory,invertebrate herbivores and predators across a natural tree diversity gradient in Germany’s largest connected deciduous forest

Tree species-rich forests are hypothesised to be less susceptible to insect herbivores, but so far herbivory–diversity relationships have rarely been tested for tree saplings, and no such study has been published for deciduous forests in Central Europe. We expected that diverse tree communities reduce the probability of detection of host plants and increase abundance of predators, thereby reducing herbivory. We examined levels of herbivory suffered by beech (Fagus sylvatica L.) and maple saplings (Acer pseudoplatanus L. and Acer platanoides L.) across a tree species diversity gradient within Germany’s largest remaining deciduous forest area, and investigated whether simple beech or mixed stands were less prone to damage caused by herbivorous insects. Leaf area loss and the frequency of galls and mines were recorded for 1,040 saplings (>13,000 leaves) in June and August 2006. In addition, relative abundance of predators was assessed to test for potential top-down control. Leaf area loss was generally higher in the two species of maple compared to beech saplings, while only beech showed a decline in damage caused by leaf-chewing herbivores across the tree diversity gradient. No significant patterns were found for galls and mines. Relative abundance of predators on beech showed a seasonal response and increased on species-rich plots in June, suggesting higher biological control. We conclude that, in temperate deciduous forests, herbivory–tree diversity relationships are significant, but are tree species-dependent with bottom-up and top-down control as possible mechanisms. In contrast to maple, beech profits from growing in a neighbourhood of higher tree richness, which implies that species identity effects may be of greater importance than tree diversity effects per se. Hence, herbivory on beech appeared to be mediated bottom-up by resource concentration in the sampled forest stands, as well as regulated top-down through biocontrol by natural enemies. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorised users.     

Bird predation enhances tree seedling resistance to insect herbivores in contrasting forest habitats

According to the associational resistance hypothesis, neighbouring plants are expected to influence both the insect herbivore communities and their natural enemies. However, this has rarely been tested for the effects of canopy trees on herbivory of seedlings. One possible mechanism responsible for associational resistance is the indirect impact of natural enemies on insect herbivory, such as insectivorous birds. But it remains unclear to what extent such trophic cascades are influenced by the composition of plant associations (i.e. identity of ‘associated’ plants). Here, we compared the effect of bird exclusion on insect leaf damage for seedlings of three broadleaved tree species in three different forest habitats. Exclusion of insectivorous birds affected insect herbivory in a species-specific manner: leaf damage increased on Betula pendula seedlings whereas bird exclusion had no effect for two oaks (Quercus robur and Q. ilex). Forest habitat influenced both the extent of insect herbivory and the effect of bird exclusion. Broadleaved seedlings had lower overall leaf damage within pine plantations than within broadleaved stands, consistent with the resource concentration hypothesis. The indirect effect of bird exclusion on leaf damage was only significant in pine plantations, but not in exotic and native broadleaved woodlands. Our results support the enemies hypothesis, which predicts that the effects of insectivorous birds on insect herbivory on seedlings are greater beneath non-congeneric canopy trees. Although bird species richness and abundance were greater in broadleaved woodlands, birds were unable to regulate insect herbivory on seedlings in forests of more closely related tree species.     

不同采伐强度对阔叶红松林主要树种空间分布格局和物种空间关联性的影响

分析采伐后森林群落中物种的空间格局有助于认识该格局形成的生态学过程、种群的生物学特性及其与环境因子之间的相互关系,并对制定可持续的森林经营方案具有重要意义。以长白山地区经历不同采伐方式形成的阔叶红松林次生林为研究对象,利用空间点格局分析的研究方法,探讨了采伐对阔叶红松林主要树种空间分布格局、种间关联性以及更新的影响。研究结果显示:较低强度的择伐对阔叶红松林主要树种的空间分布格局、种间关联性的改变较小,群落可以在较短时间内恢复。中等强度的择伐减少了成年树种对幼树的抑制作用,可以促进森林的天然更新。皆伐后,森林的群落结构,物种的空间分布格局、种间关联性都发生显著变化,尽管更新状况良好,但要恢复到伐前水平仍需要较长时间。择伐不仅通过改变主要树种的密度对阔叶红松林群落结构产生影响,还通过改变物种空间关联性改变群落的结构动态。因此,在制定森林生态系统经营管理方案时,不仅要选择适合的采伐强度,还要综合考虑采伐时物种的选择以及种间关系。     

Effect of Host Tree Traits on Epiphyte Diversity in Natural and Anthropogenic Habitats in Ecuador

Vascular epiphytes represent a highly diverse element of tropical rain forests, but they depend strongly on the structure and taxonomic composition of their tree communities. For conservation planning, it is therefore critical to understand the effect of host tree characteristics on epiphyte species richness in natural and anthropogenically transformed vegetation. Our study compares the effect of human land‐use on epiphyte diversity based on 220 study plots in a lowland rain forest and an Andean cloud forest in western Ecuador. We evaluate the relevance of host tree size and taxonomic identity for epiphyte species richness in contiguous primary forests, forest fragments, isolated remnant trees (IRTs), and secondary forests. At both study sites, epiphyte diversity was highest in primary forests, and it was lowest on IRTs and in secondary forests. Epiphyte species numbers of forest fragments were significantly reduced compared with the contiguous primary forest at the lowland study site, but not in the cloud forest area. Host tree size was a core predictor among secondary forests, but it had less significance within other habitat types. Taxonomic identity of the host trees also explained up to 61 percent of the variation in epiphyte diversity, especially for IRTs. The structural and taxonomic composition of the tree community in anthropogenically transformed habitat types proved to be fundamental to epiphyte diversity. This highlights the importance of deliberate selection of tree species for reforestation in conservation programs and the possible negative effects of selective logging in primary forests. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Global patterns of insect herbivory in gap and understorey environments,and their implications for woody plant carbon storage

Insect herbivory is thought to favour carbon allocation to storage in juveniles of shade‐tolerant trees. This argument assumes that insect herbivory in the understorey is sufficiently intense as to select for storage; however, understoreys might be less attractive to insect herbivores than canopy gaps, because of low resource availability and – at temperate latitudes – low temperatures. Although empirical studies show that shade‐tolerant species in tropical forests do allocate more photosynthate to storage than their light‐demanding associates, the same pattern has not been consistently observed in temperate forests. Does this reflect a latitudinal trend in the relative activity of insect herbivory in gap versus understorey environments? To date there has been no global review of the effect of light environment on insect herbivory in forests. We postulated that if temperature is the primary factor limiting insect herbivory, the effect of gaps on rates of insect herbivory should be more evident in temperate than in tropical forests; due to low growing season temperatures in the oceanic temperate forests of the Southern Hemisphere, the effect of gaps on insect herbivory rates should in turn be stronger there than in the more continental temperate climates of the Northern Hemisphere. We examined global patterns of insect herbivory in gaps versus understories through meta‐analysis of 87 conspecific comparisons of leaf damage in contrasting light environments. Overall, insect herbivory in gaps was significantly higher than in the understorey; insect herbivory was 50% higher in gaps than in understoreys of tropical forests but did not differ significantly between gaps and understories in temperate forests of either hemisphere. Results are consistent with the idea that low resource availability – and not temperature – limits insect herbivore activity in forest understoreys, especially in the tropics, and suggest the selective influence of insect herbivory on late‐successional tree species may have been over‐estimated.     

Edge disturbance drives liana abundance increase and alteration of liana–host tree interactions in tropical forest fragments

Closed‐canopy forests are being rapidly fragmented across much of the tropical world. Determining the impacts of fragmentation on ecological processes enables better forest management and improves species‐conservation outcomes. Lianas are an integral part of tropical forests but can have detrimental and potentially complex interactions with their host trees. These effects can include reduced tree growth and fecundity, elevated tree mortality, alterations in tree‐species composition, degradation of forest succession, and a substantial decline in forest carbon storage. We examined the individual impacts of fragmentation and edge effects (0–100‐m transect from edge to forest interior) on the liana community and liana–host tree interactions in rainforests of the Atherton Tableland in north Queensland, Australia. We compared the liana and tree community, the traits of liana‐infested trees, and determinants of the rates of tree infestation within five forest fragments (23–58 ha in area) and five nearby intact‐forest sites. Fragmented forests experienced considerable disturbance‐induced degradation at their edges, resulting in a significant increase in liana abundance. This effect penetrated to significantly greater depths in forest fragments than in intact forests. The composition of the liana community in terms of climbing guilds was significantly different between fragmented and intact forests, likely because forest edges had more small‐sized trees favoring particular liana guilds which preferentially use these for climbing trellises. Sites that had higher liana abundances also exhibited higher infestation rates of trees, as did sites with the largest lianas. However, large lianas were associated with low‐disturbance forest sites. Our study shows that edge disturbance of forest fragments significantly altered the abundance and community composition of lianas and their ecological relationships with trees, with liana impacts on trees being elevated in fragments relative to intact forests. Consequently, effective control of lianas in forest fragments requires management practices which directly focus on minimizing forest edge disturbance.     

Soil heterogeneity in tree mixtures depends on spatial clustering of tree species

Heterogeneity in soil characteristics promotes and maintains coexistence between a diverse set of species. In forests, trees have species-specific impacts on soil abiotic characteristics and mixing of tree species is being promoted as a tool to ensure high levels of diversity and functioning. Yet, limited knowledge is available on the effect of tree species composition and spatial clustering on heterogeneity in soil characteristics. In this paper we derived heterogeneity of key characteristics of the leaf litterfall, the forest floor and the mineral topsoil (C, N and base cation concentration, C:N ratio and mass) in 53 plots of 7 different tree species compositions. We found that heterogeneity increased from the leaf litterfall, through the forest floor down to the mineral topsoil. Mixing tree species did not lead to an increased heterogeneity in the forest floor and topsoil compared to monocultures. However, we did find that mixed plots where conspecific trees stand in groups are more heterogeneous than plots where species are intimately mixed. Our results imply that heterogeneity in soil characteristics does not necessarily increase with tree diversity, but that within mixed stands the spatial organization of tree species should be considered in relation to the scale at which heterogeneity is desired.     

Arthropod assemblages in a focal tree species (Eucalyptus microcarpa) depends on the species mix in restoration plantings

The composition of arthropods in trees has long been a topic of interest, with many studies exploring ways in which arthropod communities differ among tree species or entire forests. Few studies, however, have examined arthropods from trees in restoration plantings, and little is known about how different tree plantings might lead to different biodiversity outcomes. The aim of our study was to determine if a focal tree species hosted a different arthropod fauna depending on its context. We examined arthropod assemblages from the foliage of Eucalyptus microcarpa (Grey Box) trees present in two kinds of plantings: (i) simple plantings with only Eucalyptus trees, and (ii) mixed plantings with both Eucalyptus and Acacia trees. We examined the composition of the assemblages, and looked for associations between ant and psyllid (Hemiptera) diversity across each kind of planting. We found more species of psyllids, beetles and ants in E. microcarpa trees when they were grown in simple plantings compared with the mixed species plantings. We also found that psyllid richness and abundance was positively correlated with ant richness and abundance across all plantings. Our study shows that a more diverse context (i.e. a mixed planting) is not necessarily associated with a more diverse fauna within a focal tree species, and that simple Eucalyptus-only plantings may encourage higher numbers of insect herbivores. The apparent association between psyllids and ants suggests a complex interaction that may drive patterns in tree arthropod assemblages.     

Differential Responses of Herbivores and Herbivory to Management in Temperate European Beech

Forest management not only affects biodiversity but also might alter ecosystem processes mediated by the organisms, i.e. herbivory the removal of plant biomass by plant-eating insects and other arthropod groups. Aiming at revealing general relationships between forest management and herbivory we investigated aboveground arthropod herbivory in 105 plots dominated by European beech in three different regions in Germany in the sun-exposed canopy of mature beech trees and on beech saplings in the understorey. We separately assessed damage by different guilds of herbivores, i.e. chewing, sucking and scraping herbivores, gall-forming insects and mites, and leaf-mining insects. We asked whether herbivory differs among different forest management regimes (unmanaged, uneven-aged managed, even-aged managed) and among age-classes within even-aged forests. We further tested for consistency of relationships between regions, strata and herbivore guilds. On average, almost 80% of beech leaves showed herbivory damage, and about 6% of leaf area was consumed. Chewing damage was most common, whereas leaf sucking and scraping damage were very rare. Damage was generally greater in the canopy than in the understorey, in particular for chewing and scraping damage, and the occurrence of mines. There was little difference in herbivory among differently managed forests and the effects of management on damage differed among regions, strata and damage types. Covariates such as wood volume, tree density and plant diversity weakly influenced herbivory, and effects differed between herbivory types. We conclude that despite of the relatively low number of species attacking beech; arthropod herbivory on beech is generally high. We further conclude that responses of herbivory to forest management are multifaceted and environmental factors such as forest structure variables affecting in particular microclimatic conditions are more likely to explain the variability in herbivory among beech forest plots.     

The negative effect of lianas on tree growth varies with tree species and season

Lianas reduce tree growth, reproduction, and survival in tropical forests. Liana competition can be particularly intense in isolated forest fragments, where liana densities are high, and thus, host tree infestation is common. Furthermore, lianas appear to grow particularly well during seasonal drought, when they may compete particularly intensely with trees. Few studies, however, have experimentally quantified the seasonal effects of liana competition on multiple tree species in tropical forests. We used a liana removal experiment in a forest fragment in southeastern Brazil to test whether the effects of lianas on tree growth vary with season and tree species identity. We conducted monthly diameter measurements using dendrometer bands on 88 individuals of five tree species for 24 months. We found that lianas had a stronger negative effect on some tree species during the wet season compared to the dry season. Furthermore, lianas significantly reduced the diameter growth of two tree species but had no effect on the other three tree species. The strong negative effect of lianas on some trees, particularly during the wet season, indicates that the effect of lianas on trees varies both seasonally and with tree species identity. Abstract in Portuguese is available with online material.     

Observations on the Ecology of Weaver Ants (Oecophylla smaragdina Fabricius) in a Thai Mangrove Ecosystem and Their Effect on Herbivory of Rhizophora mucronata Lam.

Ants of the genus Oecophylla are predators of other insects and are able to protect a variety of terrestrial plants against pest insects; however, observations on the ecology of these ants in mangrove forests are lacking. General observations on the ecology of Oecophylla smaragdina were carried out in a Thai mangrove forest to determine if these ants can protect their host plants in less favorable mangrove habitats. Leaf herbivory and the density of O. smaragdina ants were measured on Rhizophora mucronata trees at two sites. The results showed a negative correlation between ant density and herbivory. At both sites, the mean percent damaged leaf area was more than four times higher on trees without ants compared to “ant‐trees.” A significant negative correlation was found between tree mean percent leaf damage and the density of ants on the tree. Furthermore, on trees with ants, there was less herbivory on leaves close to ant nests compared to other leaves on the tree. Most damage was caused by chrysomelid beetles (62%) and sesarmid crabs (25%) and both types of herbivory were significantly reduced on ant‐trees.     

Fine spatial pattern of an epiphytic lichen species is affected by habitat conditions in two forest types in the Iberian Mediterranean region

Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random.     

Effects of fragmentation of secondary broadleaf deciduous forests on populations of the near-threatened butterfly, Sasakia charonda (Lepidoptera, Nymphalidae), in central Japan

We examined the effects of fragmentation of secondary broadleaf deciduous forests (secondary forests) on populations of the near-threatened butterfly, Sasakia charonda, in central Japan. Regression analyses revealed that the number of overwintering larvae per host tree significantly increased when the area of secondary forest patches and the Isolation Index of the forest patch increased and the distance from secondary forest patches containing the focal host trees to the nearest secondary forest patch decreased. There was a significantly positive correlation between the number of overwintering larvae and the number of host trees in the neighborhood. The host trees were primarily distributed at the edges of secondary forests. From the results of the backward elimination method of multiple linear regression analysis, independent variables other than patch area were eliminated, and the standardized partial regression coefficient of the patch area was significant. This result suggested that a contiguous distribution of large secondary forest patches with many host trees is very important to conserving this butterfly species.     

Selective logging in tropical forests decreases the robustness of liana–tree interaction networks to the loss of host tree species

Selective logging is one of the major drivers of tropical forest degradation, causing important shifts in species composition. Whether such changes modify interactions between species and the networks in which they are embedded remain fundamental questions to assess the ‘health’ and ecosystem functionality of logged forests. We focus on interactions between lianas and their tree hosts within primary and selectively logged forests in the biodiversity hotspot of Malaysian Borneo. We found that lianas were more abundant, had higher species richness, and different species compositions in logged than in primary forests. Logged forests showed heavier liana loads disparately affecting slow-growing tree species, which could exacerbate the loss of timber value and carbon storage already associated with logging. Moreover, simulation scenarios of host tree local species loss indicated that logging might decrease the robustness of liana–tree interaction networks if heavily infested trees (i.e. the most connected ones) were more likely to disappear. This effect is partially mitigated in the short term by the colonization of host trees by a greater diversity of liana species within logged forests, yet this might not compensate for the loss of preferred tree hosts in the long term. As a consequence, species interaction networks may show a lagged response to disturbance, which may trigger sudden collapses in species richness and ecosystem function in response to additional disturbances, representing a new type of ‘extinction debt’.     

Tree diversity drives associational resistance to herbivory at both forest edge and interior

Tree diversity is increasingly acknowledged as an important driver of insect herbivory. However, there is still a debate about the direction of associational effects that can range from associational resistance (i.e., less damage in mixed stands than in monocultures) to the opposite, associational susceptibility. Discrepancies among published studies may be due to the overlooked effect of spatially dependent processes such as tree location within forests. We addressed this issue by measuring crown defoliation and leaf damage made by different guilds of insect herbivores on oaks growing among conspecific versus heterospecific neighbors at forest edges versus interior, in two closed sites in SW France forests. Overall, oaks were significantly less defoliated among heterospecific neighbors (i.e., associational resistance), at both forest edge and interior. At the leaf level, guild diversity and leaf miner herbivory significantly increased with tree diversity regardless of oak location within stands. Other guilds showed no clear response to tree diversity or oak location. We showed that herbivore response to tree diversity varied among insect feeding guilds but not between forest edges and interior, with inconsistent patterns between sites. Importantly, we show that oaks were more defoliated in pure oak plots than in mixed plots at both edge and forest interior and that, on average, defoliation decreased with increasing tree diversity from one to seven species. We conclude that edge conditions could be interacting with tree diversity to regulate insect defoliation, but future investigations are needed to integrate them into the management of temperate forests, notably by better understanding the role of the landscape context.     

Sustained high levels of foliar herbivory of the mangrove Rhizophora stylosa by a moth larva Doratifera stenosa (Limacodidae) in north-eastern Australia

The moth larva, Doratifera stenosa (Lepidoptera: Limacodidae), was observed feeding voraciously in great numbers on mature leaves of Rhizophora stylosa in mangroves at Port Curtis in Central Queensland, NE Australia. This behaviour was considered unusual since mangroves, and the Rhizophora species in particular, reportedly harbour few herbivores and have relatively low levels of herbivory, less than 10%. During a two year period (1996–1998), larvae were observed consuming around 30–40% of leaves in the canopy each year, and the mangroves appeared able to sustain these high levels of herbivory. The impact on trees was assessed in conjunction with a study of the herbivore, its behaviour and life history, in an attempt to explain the occurrence. Larvae were 1–2 cm in length, bright green and gregarious, with numerous small, stinging hairs along their upper bodies. Feeding was in small cohort groups of 5–70 individuals that broke up immediately prior to each moult after which they regrouped in much larger numbers of mixed cohorts to form single-file processions across branches, stems and prop roots. In this way, they moved to neighbouring trees with less affected foliage. One of the outstanding characteristics of this herbivore was its ability to desist from killing host trees although it appeared quite capable of doing so had it remained on individual trees. By moving from tree to tree, the herbivore was able to heavily crop Rhizophora foliage in an apparently sustainable manner. These findings demonstrate the role and importance of foliar herbivory in severely affected forests and how such instances best not be ignored or treated as curiosities in future assessments of herbivory and forest turnover in mangrove ecosystems.     

Altitudinal patterns in host suitability for forest insects

Conspecific trees growing at high and low-elevations encounter different growing conditions and may vary in their suitability as hosts for herbivorous insects. Mountain tree populations may be more resistant to herbivory if low temperatures constrain growth more than they constrain photosynthesis, resulting in increased secondary metabolism (temperature hypothesis). Alternatively, mountain trees may be fertilized by atmospheric nitrogen deposition and become more palatable to insects (atmospheric deposition hypothesis). We evaluated these two hypotheses by comparing high- and low-elevation trees with insect bioassays and analyses of foliar nitrogen and condensed tannin. Contrary to the temperature hypothesis, high-elevation foliage had higher leaf nitrogen (six of six tree species) and allowed higher growth rates of Lymantria dispar larvae (five of six tree species). The nitrogen deposition hypothesis was broadly supported by measurements from two mountains showing that high-elevation trees tended to have higher leaf nitrogen, lower leaf tannins, and support higher insect growth performance than conspecific trees from lower elevations. The deposition hypothesis was further supported by fertilization studies showing that simulated atmospheric nitrogen deposition changed the foliar chemistry of valley trees to resemble that of high-elevation trees. Predictions that the altitudinal gradient in foliar chemistry and host suitability should be steepest on mountains receiving more deposition were largely not supported, but interpretations are complicated by lack of replication among mountains. In the northeastern United States, increased host suitability of high-elevation trees seems sufficient to influence the population dynamics and community composition of herbivores. Atmospheric nitrogen deposition offers a promising hypothesis to explain and predict some important spatial patterns in herbivory. Received: 21 September 1997 / Accepted: 12 June 1998     

Native plant diversity increases herbivory to non-natives

There is often an inverse relationship between the diversity of a plant community and the invasibility of that community by non-native plants. Native herbivores that colonize novel plants may contribute to diversity–invasibility relationships by limiting the relative success of non-native plants. Here, we show that, in large collections of non-native oak trees at sites across the USA, non-native oaks introduced to regions with greater oak species richness accumulated greater leaf damage than in regions with low oak richness. Underlying this trend was the ability of herbivores to exploit non-native plants that were close relatives to their native host. In diverse oak communities, non-native trees were on average more closely related to native trees and received greater leaf damage than those in depauperate oak communities. Because insect herbivores colonize non-native plants that are similar to their native hosts, in communities with greater native plant diversity, non-natives experience greater herbivory.