Latitudinal variation in herbivory: influences of climatic drivers,herbivore identity and natural enemies

Although a number of investigations have concluded that lower latitudes are associated with increases in herbivore abundance and plant damage, the generality of this pattern is still under debate. Multiple factors may explain the lack of consistency in latitude–herbivory relationships. For instance, latitudinal variation in herbivore pressure may be shaped entirely or not by climatic variables, or vary among herbivore guilds with differing life‐history traits. Additionally, the strength of top–down effects from natural enemies on herbivores might also vary geographically and influence latitude–herbivory patterns. We carried out a field study where we investigated the effects of latitude and climate on herbivory by a seed‐eating caterpillar and leaf chewers, as well as parasitism associated to the former across 30 populations of the perennial herb Ruellia nudiflora (Acanthaceae). These populations were distributed along a 5° latitudinal gradient from northern Yucatan (Mexico) to southern Belize, representing one‐third of the species' latitudinal distribution and the entirety and one‐third of the precipitation and temperature gradient of this species' distribution (respectively). We found opposing latitudinal gradients of seed herbivory and leaf herbivory, and this difference appeared to be mediated by contrasting effects of climate on each guild. Specifically, univariate regressions showed that seed herbivory increased at higher latitudes and with colder temperatures, while leaf herbivory increased toward the equator and with wetter conditions. Multiple regressions including temperature, precipitation and latitude only found significant effects of temperature for seed herbivory and latitude for leaf herbivory. Accordingly, that latitudinal variation in seed herbivory appears to be driven predominantly by variation in temperature whereas latitudinal variation in leaf herbivory was apparently driven by other unexplored correlates of latitude. Parasitism did not exhibit variation with latitude or climatic factors. Overall, these findings underscore that the factors driving latitudinal clines in herbivory might vary even among herbivore species coexisting on the same host plant.     

Decreased losses of woody plant foliage to insects in large urban areas are explained by bird predation

Despite the increasing rate of urbanization, the consequences of this process on biotic interactions remain insufficiently studied. Our aims were to identify the general pattern of urbanization impact on background insect herbivory, to explore variations in this impact related to characteristics of both urban areas and insect–plant systems, and to uncover the factors governing urbanization impacts on insect herbivory. We compared the foliar damage inflicted on the most common trees by defoliating, leafmining and gall‐forming insects in rural and urban habitats associated with 16 European cities. In two of these cities, we explored quality of birch foliage for herbivorous insects, mortality of leafmining insects due to predators and parasitoids and bird predation on artificial plasticine larvae. On average, the foliage losses to insects were 16.5% lower in urban than in rural habitats. The magnitude of the overall adverse effect of urbanization on herbivory was independent of the latitude of the locality and was similar in all 11 studied tree species, but increased with an increase in the size of the urban area: it was significant in large cities (city population 1–5 million) but not significant in medium‐sized and small towns. Quality of birch foliage for herbivorous insects was slightly higher in urban habitats than in rural habitats. At the same time, leafminer mortality due to ants and birds and the bird attack intensity on dummy larvae were higher in large cities than in rural habitats, which at least partially explained the decline in insect herbivory observed in response to urbanization. Our findings underscore the importance of top‐down forces in mediating impacts of urbanization on plant‐feeding insects: factors favouring predators may override the positive effects of temperature elevation on insects and thus reduce plant damage.     

Impacts of urbanization on insect herbivory and plant defences in oak trees

Systematic comparisons of species interactions in urban versus rural environments can improve our understanding of shifts in ecological processes due to urbanization. However, such studies are relatively uncommon and the mechanisms driving urbanization effects on species interactions (e.g. between plants and insect herbivores) remain elusive. Here we investigated the effects of urbanization on leaf herbivory by insect chewers and miners associated with the English oak Quercus robur by sampling trees in rural and urban areas throughout most of the latitudinal distribution of this species. In performing these comparisons, we also controlled for the size of the urban areas (18 cities) and gathered data on CO₂ emissions. In addition, we assessed whether urbanization affected leaf chemical defences (phenolic compounds) and nutritional traits (phosphorus and nitrogen), and whether such changes correlated with herbivory levels. Urbanization significantly reduced leaf chewer damage but did not affect leaf miners. In addition, we found that leaves from urban locations had lower levels of chemical defences (condensed and hydrolysable tannins) and higher levels of nutrients (nitrogen and phosphorus) compared to leaves in rural locations. The magnitude of urbanization effects on herbivory and leaf defences was not contingent upon city size. Importantly, while the effects of urbanization on chemical defences were associated with CO₂ emissions, changes in leaf chewer damage were not associated with either leaf traits or CO₂ levels. These results suggest that effects of urbanization on herbivory occur through mechanisms other than changes in the plant traits measured here. Overall, our simultaneous assessment of insect herbivory, plant traits and abiotic correlates advances our understanding of the main drivers of urbanization effects on plant–herbivore interactions.     

Different mechanisms explain feeding type-specific patterns of latitudinal variation in herbivore damage among diverse feeding types of herbivorous insects

To determine whether latitudinal variation in herbivore impact exists, we examined three major herbivorous insect feeding types (chewers, gallers, and miners) on/in leaves of Japanese beech. Herbivores were collected with litter traps deployed in forests across a latitudinal gradient of 10°. Leaf litter analyses demonstrated that chewing herbivory increased with increasing latitude of collection site. However, the densities of miners and gallers decreased with latitude. To test whether latitudinal variation in herbivore damage occurs in the absence of geographically differentiated environmental cueing (e.g., physical stresses or herbivore damage), we measured both genetically determined constitutive leaf traits and herbivore damage in a common-garden experiment. In this experiment, miner density decreased with latitude, but chewing herbivory did not vary latitudinally. Galler density was higher on trees from native provenances than on trees from unrelated provenances likely because of local adaptations. Leaf mass per unit area (LMA), tannin, and phenolics all decreased with latitude of provenance. The latitudinal variation in one constitutive leaf trait (LMA) best explained latitudinal variation in chewing herbivory. Thus, different mechanisms account for feeding type-specific patterns of latitudinal variation in herbivore damage among different herbivore feeding types.     

Latitudinal variation in plant chemical defences drives latitudinal patterns of leaf herbivory

A long‐standing paradigm in ecology holds that herbivore pressure and thus plant defences increase towards lower latitudes. However, recent work has challenged this prediction where studies have found no relationship or opposite trends where herbivory or plant defences increase at higher latitudes. Here we tested for latitudinal variation in herbivory, chemical defences (phenolic compounds), and nutritional traits (phosphorus and nitrogen) in leaves of a long‐lived tree species, the English oak Quercus robur. We further investigated the underlying climatic and soil factors associated with such variation. Across 38 populations of Q. robur distributed along an 18° latitudinal gradient, covering almost the entire latitudinal and climatic range of this species, we observed strong but divergent latitudinal gradients in leaf herbivory and leaf chemical defences and nutrients. As expected, there was a negative relationship between latitude and leaf herbivory where oak populations from lower latitudes exhibited higher levels of leaf herbivory. However, counter to predictions there was a positive relationship between leaf chemical defences and latitude where populations at higher latitudes were better defended. Similarly, leaf phosphorus and nitrogen increased with latitude. Path analysis indicated a significant (negative) effect of plant chemical defences (condensed tannins) on leaf herbivory, suggesting that the latitudinal gradient in leaf herbivory was driven by an inverse gradient in defensive investment. Leaf nutrients had no independent influence on herbivory. Further, we found significant indirect effects of precipitation and soil porosity on leaf herbivory, which were mediated by plant chemical defences. These findings suggest that abiotic factors shape latitudinal variation in plant defences and that these defences in turn underlie latitudinal variation in leaf herbivory. Overall, this study contributes to a better understanding of latitudinal variation in plant–herbivore interactions by determining the identity and modus operandi of abiotic factors concurrently shaping plant defences and herbivory.     

A test of the latitudinal defense hypothesis: herbivory, tannins and total phenolics in four North American tree species

It is widely believed that insect herbivory is less intense at higher latitudes, due to winter mortality which would tend to keep insect herbivores from reaching density-limitation of their populations. One prediction of this theory is that plants should tend to be better defended at lower latitudes. Here we investigated latitudinal trends in herbivory and tannins, in four species of common North American trees. Our comparisons spanned 15° of latitude in Acer rubrum, Fagus grandifolia, and Quercus alba, and 10° latitude in Liquidambar styraciflua. Sun leaves on forest edges were sampled, at phenologically equivalent times of year. Analysis revealed significant differences between populations, including those at similar latitudes, but no significant latitudinal trend in herbivory, condensed and hydrolyzable tannins, or total phenolics measured as Folin–Denis reactives in any of the four species. Our findings contradict the theory that low latitude plants are better defended, in that lower latitude populations of the four tree species showed no greater amounts of phenolics. The possible implications for community ecology are discussed. Jonathan M. Adams, Brian Rehill, and Yangjian Zhang are equal joint lead authorship on this paper.     

Abundance and diversity of birch‐feeding leafminers along latitudinal gradients in northern Europe

Latitudinal patterns in biotic interactions, including herbivory, have been widely debated during the past years. In particular, recent meta‐analysis questioned the hypothesis that herbivory increases from the poles towards the equator. Our study was designed to verify this hypothesis by exploring latitudinal patterns in abundance and diversity of birch‐feeding insect herbivores belonging to the leafminer guild in northern Europe, from 59° to 69°N. We collected branches from five mature trees of two birch species (Betula pendula and B. pubescens) at each study site (ten sites for each of five latitudinal gradients) twice per season (in early and late summer of 2008–2011) and attributed all mines found on leaves of these branches to a certain taxon of insects. Latitudinal patterns were quantified by calculating Spearman rank correlation coefficients between both abundance and diversity of leafmining taxa and latitudes of sampling sites. In general, both abundance and diversity of leafminers significantly decreased with latitude. However, we discovered pronounced variation in patterns of latitudinal changes among study years and leafminer taxa. Variation among study years was best explained by mean temperatures in July at the northern ends of our gradients. During cold years, abundance of leafminers significantly decreased with latitude, while during warm years the abundance was either independent of latitude or even increased towards the pole. In the northern boreal forests (66° to 69°N), herbivores demonstrated larger changes in densities in response to temperature variations than in the boreo‐nemoral forests (59° to 62°N). Our data suggest that climate warming will result in a stronger increase in herbivory at higher latitudes than at lower latitudes.     

The Effects of Urban Warming on Herbivore Abundance and Street Tree Condition

Trees are essential to urban habitats because they provide services that benefit the environment and improve human health. Unfortunately, urban trees often have more herbivorous insect pests than rural trees but the mechanisms and consequences of these infestations are not well documented. Here, we examine how temperature affects the abundance of a scale insect, Melanaspis tenebricosa (Comstock) (Hemiptera: Diaspididae), on one of the most commonly planted street trees in the eastern U.S. Next, we examine how both pest abundance and temperature are associated with water stress, growth, and condition of 26 urban street trees. Although trees in the warmest urban sites grew the most, they were more water stressed and in worse condition than trees in cooler sites. Our analyses indicate that visible declines in tree condition were best explained by scale-insect infestation rather than temperature. To test the broader relevance of these results, we extend our analysis to a database of more than 2700 Raleigh, US street trees. Plotting these trees on a Landsat thermal image of Raleigh, we found that warmer sites had over 70% more trees in poor condition than those in cooler sites. Our results support previous studies linking warmer urban habitats to greater pest abundance and extend this association to show its effect on street tree condition. Our results suggest that street tree condition and ecosystem services may decline as urban expansion and global warming exacerbate the urban heat island effect. Although our non-probability sampling method limits our scope of inference, our results present a gloomy outlook for urban forests and emphasize the need for management tools. Existing urban tree inventories and thermal maps could be used to identify species that would be most suitable for urban conditions.     

Responses of arthropod populations to warming depend on latitude: evidence from urban heat islands

Biological effects of climate change are expected to vary geographically, with a strong signature of latitude. For ectothermic animals, there is systematic latitudinal variation in the relationship between climate and thermal performance curves, which describe the relationship between temperature and an organism's fitness. Here, we ask whether these documented latitudinal patterns can be generalized to predict arthropod responses to warming across mid‐ and high temperate latitudes, for taxa whose thermal physiology has not been measured. To address this question, we used a novel natural experiment consisting of a series of urban warming gradients at different latitudes. Specifically, we sampled arthropods from a single common street tree species across temperature gradients in four US cities, located from 35.8 to 42.4° latitude. We captured 6746 arthropods in 34 families from 111 sites that varied in summer average temperature by 1.7–3.4 °C within each city. Arthropod responses to warming within each city were characterized as Poisson regression coefficients describing change in abundance per °C for each family. Family responses in the two midlatitude cities were heterogeneous, including significantly negative and positive effects, while those in high‐latitude cities varied no more than expected by chance within each city. We expected high‐latitude taxa to increase in abundance with warming, and they did so in one of the two high‐latitude cities; in the other, Queens (New York City), most taxa declined with warming, perhaps due to habitat loss that was correlated with warming in this city. With the exception of Queens, patterns of family responses to warming were consistent with predictions based on known latitudinal patterns in arthropod physiology relative to regional climate. Heterogeneous responses in midlatitudes may be ecologically disruptive if interacting taxa respond oppositely to warming.     

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.     

Latitudinal patterns of herbivore pressure in a temperate herb support the biotic interactions hypothesis

The longstanding biotic interactions hypothesis predicts that herbivore pressure declines with latitude, but the evidence is mixed. To address gaps in previous studies, we measured herbivory and defence in the same system, quantified defence with bioassays, and considered effects of leaf age. We quantified herbivory and defence of young and mature leaves along a continental gradient in eastern North America in the native herb Phytolacca americana L. Herbivory in the field declined with latitude and was strongly correlated with lepidopteran abundance. Laboratory bioassays revealed that leaf palatability was positively correlated with latitude of origin. Young leaves were more damaged than mature leaves at lower latitudes in the field, but less palatable in bioassays. Both defence and palatability displayed non‐linear latitudinal patterns, suggesting potential mechanisms based on biological or climatic thresholds. In sum, observational and experimental studies find patterns consistent with high herbivore pressure and stronger plant defences at lower latitudes.     

Urbanization disrupts latitude‐size rule in 17‐year cicadas

Many ectotherms show a decrease in body size with increasing latitude due to changes in climate, a pattern termed converse Bergmann's rule. Urban conditions—particularly warmer temperatures and fragmented landscapes—may impose stresses on development that could disrupt these body size patterns. To test the impact of urbanization on development and latitudinal trends in body size, we launched a citizen science project to collect periodical cicadas (Magicicada septendecim) from across their latitudinal range during the 2013 emergence of Brood II. Periodical cicadas are long‐lived insects whose distribution spans a broad latitudinal range covering both urban and rural habitats. We used a geometric morphometric approach to assess body size and developmental stress based on fluctuating asymmetry in wing shape. Body size of rural cicadas followed converse Bergmann's rule, but this pattern was disrupted in urban habitats. In the north, urban cicadas were larger than their rural counterparts, while southern populations showed little variation in body size between habitats. We detected no evidence of differences in developmental stress due to urbanization. To our knowledge, this is the first evidence that urbanization disrupts biogeographical trends in body size, and this pattern highlights how the effects of urbanization may differ over a species’ range.     

Assessing the latitudinal gradient in herbivory

Plant–herbivore interactions occur in all ecosystems and provide a major avenue for energy flow to higher trophic levels. A long‐standing hypothesis to explain the latitudinal gradient in species diversity proposes that the relatively stable and frost‐free climate of the tropics should lead to more intense biotic interactions in tropical compared with temperate environments, giving rise to a greater diversity of plants and herbivores. Herbivory rates have been compared across latitudes to test this biotic interactions hypothesis, with herbivory typically being measured from observable leaf damage. However, we argue that a measure of percentage leaf damage alone does not straightforwardly reflect the cost of herbivory to the plant, and on its own does not constitute an appropriate test of the biotic interactions hypothesis. For a given amount of herbivory, the impact of herbivory is dependent upon many factors, such as the construction cost of the leaf, the growth and replacement rates and leaf life span. We investigate the latitudinal gradient in herbivory by analysing a large dataset of herbivory rates for 452 tree species and separating the species into those with short and long leaf life spans. We show that annual herbivory rates tend to be greater at lower latitudes for evergreen species (which have long‐lived leaves), but no trend in herbivory rate with latitude was found for species with short leaf life spans. Phylogenetic least squares regression assuming Ornstein‐Uhlenbeck processes also showed a negative effect of latitude on herbivory rate for evergreen trees, but we caution that viewing herbivory as a species trait is problematic. An integrative approach that incorporates leaf life span, as well as the costs of investment in growth and potential costs of losing leaf tissue, is needed to further our understanding of the ecological and evolutionary dynamics of herbivory.     

Poleward increase in feeding efficiency of leafminer Stigmella lapponica (Lepidoptera: Nepticulidae) in a latitudinal gradient crossing a boreal forest zone

Damage to plant communities imposed by insect herbivores generally decreases from low to high latitudes. This decrease is routinely attributed to declines in herbivore abundance and/or diversity, whereas latitudinal changes in per capita food consumption remain virtually unknown. Here, we tested the hypothesis that the lifetime food consumption by a herbivore individual decreases from low to high latitudes due to a temperature-driven decrease in metabolic expenses. From 2016 to 2019, we explored latitudinal changes in multiple characteristics of linear (gallery) mines made by larvae of the pygmy moth, Stigmella lapponica, in leaves of downy birch, Betula pubescens. The mined leaves were larger than intact leaves at the southern end of our latitudinal gradient (at 60°N) but smaller than intact leaves at its northern end (at 69°N), suggesting that female oviposition preference changes with latitude. No latitudinal changes were observed in larval size, mine length or area, and in per capita food consumption, but the larval feeding efficiency (quantified as the ratio between larval size and mine size) increased with latitude. Consequently, S. lapponica larvae consumed less foliar biomass at higher latitudes than at lower latitudes to reach the same size. Based on space-for-time substitution, we suggest that climate warming will increase metabolic expenses of insect herbivores with uncertain consequences for plant–herbivore interactions.     

Abundance–body mass relationships among insects along a latitudinal gradient

Abstract We investigated the relationship between abundance and body size (body mass) of 162 insect herbivore species on the host plant Acacia falcata along its entire coastal latitudinal distribution (eastern Australia), spanning a gradient in mean annual temperature of 4.3°C. We extend previous research by assessing these relationships at different spatial scales (latitudes pooled, among latitudes and within latitudes) and at different taxonomic levels (insect phytophages pooled, phytophagous Coleoptera and Hemiptera, and five component suborders/superfamilies). Insect species were collected from two orders (Hemiptera and Coleoptera) and five component suborders/superfamilies. There were no consistent trends in the relationships (linear or polygonal/hump‐shaped) between abundance and body mass when latitudes were pooled, among latitudes, or when phytophagous insect species were separated into their component suborder/superfamily groups. The reason for the lack of consistent trends might be due to the insect herbivores not fully exploiting their host plant and the relative absence of competition among herbivore species for food resources. This is further assessed in relation to the lack of a consistent pattern in species richness of Coleoptera and Hemiptera herbivores from the same dataset and rates of chewing and sap‐sucking herbivory along the same latitudinal gradient. Future studies of abundance–body size relationships are discussed in relation to sampling across environmental gradients and accounting for the influence of host plant identity and insect phylogeny.     

Do cities simulate climate change? A comparison of herbivore response to urban and global warming

Cities experience elevated temperature, CO₂, and nitrogen deposition decades ahead of the global average, such that biological response to urbanization may predict response to future climate change. This hypothesis remains untested due to a lack of complementary urban and long‐term observations. Here, we examine the response of an herbivore, the scale insect Melanaspis tenebricosa, to temperature in the context of an urban heat island, a series of historical temperature fluctuations, and recent climate warming. We survey M. tenebricosa on 55 urban street trees in Raleigh, NC, 342 herbarium specimens collected in the rural southeastern United States from 1895 to 2011, and at 20 rural forest sites represented by both modern (2013) and historical samples. We relate scale insect abundance to August temperatures and find that M. tenebricosa is most common in the hottest parts of the city, on historical specimens collected during warm time periods, and in present‐day rural forests compared to the same sites when they were cooler. Scale insects reached their highest densities in the city, but abundance peaked at similar temperatures in urban and historical datasets and tracked temperature on a decadal scale. Although urban habitats are highly modified, species response to a key abiotic factor, temperature, was consistent across urban and rural‐forest ecosystems. Cities may be an appropriate but underused system for developing and testing hypotheses about biological effects of climate change. Future work should test the applicability of this model to other groups of organisms.     

Adaptive latitudinal variation in Common Blackbird Turdus merula nest characteristics

Nest construction is taxonomically widespread, yet our understanding of adaptive intraspecific variation in nest design remains poor. Nest characteristics are expected to vary adaptively in response to predictable variation in spring temperatures over large spatial scales, yet such variation in nest design remains largely overlooked, particularly amongst open‐cup‐nesting birds. Here, we systematically examined the effects of latitudinal variation in spring temperatures and precipitation on the morphology, volume, composition, and insulatory properties of open‐cup‐nesting Common Blackbirds’ Turdus merula nests to test the hypothesis that birds living in cooler environments at more northerly latitudes would build better insulated nests than conspecifics living in warmer environments at more southerly latitudes. As spring temperatures increased with decreasing latitude, the external diameter of nests decreased. However, as nest wall thickness also decreased, there was no variation in the diameter of the internal nest cups. Only the mass of dry grasses within nests decreased with warmer temperatures at lower latitudes. The insulatory properties of nests declined with warmer temperatures at lower latitudes and nests containing greater amounts of dry grasses had higher insulatory properties. The insulatory properties of nests decreased with warmer temperatures at lower latitudes, via changes in morphology (wall thickness) and composition (dry grasses). Meanwhile, spring precipitation did not vary with latitude, and none of the nest characteristics varied with spring precipitation. This suggests that Common Blackbirds nesting at higher latitudes were building nests with thicker walls in order to counteract the cooler temperatures. We have provided evidence that the nest construction behavior of open‐cup‐nesting birds systematically varies in response to large‐scale spatial variation in spring temperatures.     

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.     

Weak latitudinal gradients in insect herbivory for dominant rangeland grasses of North America

Patterns of insect herbivory may follow predictable geographical gradients, with greater herbivory at low latitudes. However, biogeographic studies of insect herbivory often do not account for multiple abiotic factors (e.g., precipitation and soil nutrients) that could underlie gradients. We tested for latitudinal clines in insect herbivory as well as climatic, edaphic, and trait‐based drivers of herbivory. We quantified herbivory on five dominant grass species over 23 sites across the Great Plains, USA. We examined the importance of climate, edaphic factors, and traits as correlates of herbivory. Herbivory increased at low latitudes when all grass species were analyzed together and for two grass species individually, while two other grasses trended in this direction. Higher precipitation was related to more herbivory for two species but less herbivory for a different species, while higher specific root length was related to more herbivory for one species and less herbivory for a different species. Taken together, results highlight that climate and trait‐based correlates of herbivory can be highly contextual and species‐specific. Patterns of insect herbivory on dominant grasses support the hypothesis that herbivory increases toward lower latitudes, though weakly, and indicates that climate change may have species‐specific effects on plant–herbivore interactions.     

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.