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.     

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.     

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 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.     

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.     

Urbanization drives unique latitudinal patterns of insect herbivory and tree condition

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

Latitudinal variation in resistance and tolerance to herbivory of a salt marsh shrub

Interactions between plants and herbivores often vary on a geographic scale. Although theory about plant defenses and tolerance is predicated on temporal or spatial variation in herbivore damage, no single study has compared the pattern of herbivory, plant defenses and tolerance to herbivory of a single species across a latitudinal gradient. In 2002–2005 we surveyed replicate salt marshes along the Atlantic coast of the United States from Florida to Maine. At each field site we scored leaves of Iva frutescens for herbivore damage. In laboratory experiments we measured constitutive resistance and induced resistance in I. frutescens from high and low latitude sites along the Atlantic Coast. In another common garden experiment we studied tolerance to herbivory of I. frutescens from various sites. Theory predicts that constitutive resistance should matter more when damage is high, and induced resistance when herbivory is high but variable. In the field, average levels of herbivore damage, and spatial and temporal variation in herbivore damage were all greater at low versus high latitudes, indicating that constitutive as well as induced resistance should be stronger at low latitudes. Consistent with this prediction, constitutive resistance to herbivory was stronger at low latitudes. Induced resistance to herbivores was also stronger at low latitudes: it was deployed faster and lasted longer. Theory also predicts that tolerance to herbivory should be greater where average herbivory damage is greater; however, tolerance to herbivory in Iva did not depend on geographic origin. Our results emphasize the value of considering multiple ways in which plants respond to herbivores when examining geographic variation in plant–herbivore interactions.     

Biogeographic consequences of nutrient enrichment for plant–herbivore interactions in coastal wetlands

A major challenge in ecology is to understand broadscale trends in the impact of environmental change. We provide the first integrative analysis of the effects of eutrophication on plants, herbivores, and their interactions in coastal wetlands across latitudes. We show that fertilisation strongly increases herbivory in salt marshes, but not in mangroves, and that this effect increases with increasing latitude in salt marshes. We further show that stronger nutrient effects on plant nitrogen concentration at higher latitudes is the mechanism likely underlying this pattern. This biogeographic variation in nutrient effects on plant–herbivore interactions has consequences for vegetation, with those at higher latitudes being more vulnerable to consumer pressure fuelled by eutrophication. Our work provides a novel, mechanistic understanding of how eutrophication affects plant–herbivore systems predictably across broad latitudinal gradients, and highlights the power of incorporating biogeography into understanding large‐scale variability in the impacts of environmental change.     

Corymbia leaf oils,latitude, hybrids and herbivory: A test using common‐garden field trials

Foliar oils, particularly monoterpenes, can influence the susceptibility of plants to herbivory. In plants, including eucalypts, monoterpenes are often associated with plant defence. A recent analysis revealed an increase in foliar oil content with increasing latitudinal endemism, and we tested this pattern using three eucalypt taxa comprising a latitudinal replacement cline. We also examined the relative concentrations of two monoterpenes (α‐pinene and 1,8‐cineole), for which meta‐analyses also showed latitudinal variation, using hybrids of these three taxa with Corymbia torelliana. These, and pure C. torelliana, were then assessed in common‐garden field plots for the abundance and distribution of herbivory by four distinct herbivore taxa. Differing feeding strategies among these herbivores allowed us to test hypotheses regarding heritability of susceptibility and relationships to α‐pinene and 1,8‐cineole. We found no support for an increase in foliar oil content with increasing latitude, nor did our analysis support predictions for consistent variation in α‐pinene and 1,8‐cineole contents with latitude. However, herbivore species showed differential responses to different taxa and monoterpene contents. For example, eriophyid mites, the most monophagous of our censused herbivores, avoided the pure species, but fed on hybrid taxa, supporting hypotheses on hybrid susceptibility. The most polyphagous herbivore (leaf blister sawfly Phylacteophaga froggatti) showed no evidence of response to plant secondary metabolites, while the distribution and abundance patterns of Paropsis atomaria showed some relationship to monoterpene yields.     

Seedling Mortality and Herbivory Damage in Subtropical and Temperate Populations: Testing the Hypothesis of Higher Herbivore Pressure Toward the Tropics

Herbivory rates are generally thought to be higher in tropical than in temperate forests. Nevertheless, tests of this biogeographic prediction by comparing a single plant species across a tropical-temperate range are scarce. Here, we compare herbivore damage between subtropical and temperate populations of the evergreen tree Aextoxicon punctatum (Olivillo), distributed between 30° and 43° S along the Pacific margin of Chile. To assess the impact of herbivory on Olivillo seedlings, we set up 29 experimental plots, 1.5 × 3 m: 16 in forests of Fray Jorge National Park (subtropical latitude), and 13 in Guabún, Chiloé Island (temperate latitude). Half of each plot was fenced around with chicken wire, to exclude small mammals, and the other half was left unfenced. In each half of the plots we planted 16 seedlings of Olivillo in December 2003, with a total of 928 plants. Seedling survival, leaf production and herbivory by invertebrates were monitored over the next 16 mo. Small mammal herbivores killed ca 30 percent of seedlings in both sites. Nevertheless, invertebrate herbivory was greater in the temperate forest, thus contradicting the expected trend of increasing herbivore impact toward the tropics. Seedling growth was greater in subtropical forest suggesting better conditions for tree growth or that higher invertebrate herbivory depressed seedling growth in the temperate forest. Invertebrate herbivory increased toward temperate latitudes while small mammal herbivory was similar in both sites. We suggest that comparison of single species can be useful to test generalizations about latitudinal patterns and allow disentangling factors controlling herbivory patterns across communities.     

Latitudinal variation in resistance and tolerance to herbivory in the perennial herb Lythrum salicaria is related to intensity of herbivory and plant phenology

Both the length of the growing season and the intensity of herbivory often vary along climatic gradients, which may result in divergent selection on plant phenology, and on resistance and tolerance to herbivory. In Sweden, the length of the growing season and the number of insect herbivore species feeding on the perennial herb Lythrum salicaria decrease from south to north. Previous common‐garden experiments have shown that northern L. salicaria populations develop aboveground shoots earlier in the summer and finish growth before southern populations do. We tested the hypotheses that resistance and tolerance to damage vary with latitude in L. salicaria and are positively related to the intensity of herbivory in natural populations. We quantified resistance and tolerance of populations sampled along a latitudinal gradient by scoring damage from natural herbivores and fitness in a common‐garden experiment in the field and by documenting oviposition and feeding preference by specialist leaf beetles in a glasshouse experiment. Plant resistance decreased with latitude of origin, whereas plant tolerance increased. Oviposition and feeding preference in the glasshouse and leaf damage in the common‐garden experiment were negatively related to damage in the source populations. The latitudinal variation in resistance was thus consistent with reduced selection from herbivores towards the northern range margin of L. salicaria. Variation in tolerance may be related to differences in the timing of damage in relation to the seasonal pattern of plant growth, as northern genotypes have developed further than southern have when herbivores emerge in early summer.     

Plant damage and herbivore performance change with latitude for two old‐field plant species,but rarely as predicted

A long standing hypothesis in biogeography is that latitudinal gradients in plant defenses (LGPD) should arise because selection for plant defenses is greater in the tropics compared to temperate areas. Previous studies have focused on plant traits thought to confer resistance, yet many traits may not actually confer resistance (putative resistance) or interact to influence herbivore performance. In this study, I used a multi‐trophic approach to examine relationships between latitude, herbivore pressure, and plant resistance (measured as the growth rates of herbivores) of two old‐field plant species (Solanum carolinense and Solidago altissima) using a field survey across a 12 degree gradient in the eastern US combined with laboratory bioassays measuring the performance of generalist and specialist herbivores. I used structural equation modeling to examine the direct and indirect pathways by which latitude influences herbivore pressure and plant resistance. A latitudinal gradient in plant damage was observed in the expected direction for S. caroliense (damage decreased with latitude), but the opposite relationship was observed for S. altissima. Damage to both plant species was mediated by herbivore abundances, which was in turn influenced by predator abundances. Resistance to herbivores also varied with latitude but the form of the relationship was dependent on herbivore and plant species. There were direct, non‐linear relationships between latitude and resistance (for Spodoptera exigua and Schistocerca americana feeding on S. altissima; S. exigua and Manduca sexta feeding on S. carolinense). Herbivore growth rates were also mediated by the density of S. carolinense for Leptinotarsa juncta and S. americana feeding on S. carolinense. There was no relationship between plant resistance and herbivore pressure and no indication of feedbacks. Results from this study indicate that latitudinal variation in plant resistance is complex, possibly constrained by resource availability and tradeoffs in plant defenses.     

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.     

A paradox of latitudinal leaf defense strategies in deciduous and evergreen broadleaved trees

The classical “low latitude–high defense” hypothesis is seldom supported by empirical evidence. In this context, we tested latitudinal patterns in the leaf defense traits of deciduous broadleaved (DB) and evergreen broadleaved (EGB) tree species, which are expected to affect herbivore diversity. We examined the co-occurrence of leaf defense traits (tannin and phenol content, leaf mechanical strength, leaf dry matter content, leaf mass per area, and leaf thickness) in 741 broadleaved tree species and their correlations with species geographical range in East Asian island flora. We discovered contrasting latitudinal defense strategy gradients in DB and EGB tree species. DB species employed chemical defenses (increasing tannin and phenol content) at higher latitudes and physical defenses (softer and thinner leaves) at lower latitudes, whereas EGB tree species exhibited opposite latitudinal defense patterns. The “low latitude high defense” hypothesis included a paradoxical aspect in chemical and physical defense traits across broadleaved tree species. To reconcile paradoxical defense strategies along the latitudinal gradient, we conclude that interactive correlations among leaf traits are controlled by leaf longevity, which differs between DB and EGB tree species.     

Contrasting diversity patterns of breeding Anatidae in the Northern and Southern Hemispheres

For sustaining ecosystem functions and services, environmental conservation strategies increasingly target to maintain the multiple facets of biodiversity, such as functional diversity (FD) and phylogenetic diversity (PD), not just taxonomic diversity (TD). However, spatial mismatches among these components of biodiversity can impose challenges for conservation decisions. Hence, understanding the drivers of biodiversity is critical. Here, we investigated the global distribution patterns of TD, FD, and PD of breeding Anatidae. Using null models, we clarified the relative importance of mechanisms that influence Anatidae community. We also developed random forest models to evaluate the effects of environmental variables on the Anatidae TD, FD, and PD. Our results showed that geographical variation in Anatidae diversity is hemispheric rather than latitudinal. In the species‐rich Northern Hemisphere (NH), the three diversity indices decreased with latitude within the tropical zone of the NH, but increased in the temperate zone reaching a peak at 44.5–70.0°N, where functional and phylogenetic clustering was a predominant feature. In the Southern Hemisphere (SH), Anatidae diversity increased poleward and a tendency to overdispersion was common. In NH, productivity seasonality and temperature in the coldest quarter were the most important variables. Productivity seasonality was also the most influential predictor of SH Anatidae diversity, along with peak productivity. These findings suggested that seasonality and productivity, both consistent with the energy‐diversity hypothesis, interact with the varying histories to shape the contrasting hemispheric patterns of Anatidae diversity. Phylogenetic diversity (PD) and FD underdispersion, widespread across the species‐rich, seasonally productive mid‐to‐high latitudes of the NH, reflects a rapid evolutionary radiation and resorting associated with Pleistocene cycles of glaciation. The SH continents (and southern Asia) are characterized by a widespread tendency toward PD and FD overdispersion, with their generally species‐poor communities comprising proportionately more older lineages in thermally more stable but less predictably productive environments.     

Asymmetric changes of growth and reproductive investment herald altitudinal and latitudinal range shifts of two woody species

Ongoing changes in global climate are altering ecological conditions for many species. The consequences of such changes are typically most evident at the edge of the geographical distribution of a species, where range expansions or contractions may occur. Current demographical status at geographical range limits can help us to predict population trends and their implications for the future distribution of the species. Thus, understanding the comparability of demographical patterns occurring along both altitudinal and latitudinal gradients would be highly informative. In this study, we analyse the differences in the demography of two woody species through altitudinal gradients at their southernmost distribution limit and the consistency of demographical patterns at the treeline across a latitudinal gradient covering the complete distribution range. We focus on Pinus sylvestris and Juniperus communis, assessing their demographical structure (density, age and mortality rate), growth, reproduction investment and damage from herbivory on 53 populations covering the upper, central and lower altitudes as well as the treeline at central latitude and northernmost and southernmost latitudinal distribution limits. For both species, populations at the lowermost altitude presented older age structure, higher mortality, decreased growth and lower reproduction when compared to the upper limit, indicating higher fitness at the treeline. This trend at the treeline was generally maintained through the latitudinal gradient, but with a decreased growth at the northern edge for both species and lower reproduction for P. sylvestris. However, altitudinal and latitudinal transects are not directly comparable as factors other than climate, including herbivore pressure or human management, must be taken into account if we are to understand how to infer latitudinal processes from altitudinal data.     

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.     

Latitudinal decrease in folivory within Nothofagus pumilio forests: dual effect of climate on insect density and leaf traits?

Aim The strength of consumer–plant interactions may decrease with latitude. Our objectives were to assess the spatial variation in folivory on Nothofagus pumilio and understand the influence of climate on folivory patterns as mediated by changes in folivore density and leaf traits. Location Nothofagus pumilio forests, between 38 and 55°S (Argentina). Methods We studied the correlation of leaf damage with latitude on data from 47 sampling sites, and evaluated spatial patterns of autocorrelation on latitudinally detrended data with a principal coordinates of neighbour matrices method. Path analysis was used to test the association of temperature and precipitation with leaf damage, mediated by folivore density and leaf traits. We evaluated the adequacy of this ecological model by examining the spatial pattern of autocorrelation in the residuals, and combined spatial and environmental predictors of leaf damage into partial regression. Results Leaf damage decreased with latitude, which was the only significant spatial predictor. The latitudinal decrease in temperature and precipitation was correlated with a decrease in the density of folivores and leaf size, and diminished leaf damage. Our ecological model adequately explained the spatial autocorrelation in the data: 44% of the variation in leaf damage was explained by the latitudinally structured component of the environment, whereas local environmental effects accounted for another 22%. Main conclusions We conclude that N. pumilio forests show consistent latitudinal patterns of variation in folivory, folivore density and leaf traits. Our study suggests that the latitudinal variation in folivory rates is partly driven by the influence of climate on both plants and herbivores. This warns us about the potential susceptibility of folivory rates to climate warming. We emphasize the value of large‐scale analyses as complementary to local experimental approaches to understanding the regulation of herbivory.     

Leaf herbivory and monodominance in a Cerrado–Amazonia transitional forest,Mato Grosso,Brazil

Monodominant tropical forests occur on several continents, including the Brazilian Amazon. In this study, we tested the hypothesis that seedling escape from leaf herbivory contributes to the maintenance of the monodominant Brosimum rubescens forest. The study was undertaken both in a monodominant forest of B. rubescens and in an adjacent seasonal forest in the transitional zone between the Cerrado and the Amazonian forest biomes. Percentage of leaf area damaged and herbivory rates were evaluated on young and mature leaves of seedlings of Brosimum rubescens, Protium pilosissimum and Tetragastris altissima in the understory and in the gap between the monodominant and seasonal forests. Little evidence of any significant relationship between leaf herbivory and seedling density indicates that the monodominant species does not follow the hypothesized pattern of an intensive herbivore attack in areas of higher seedling density. The escape of Brosimum rubescens from herbivore pressure under conditions of high seedling density may be part of a set of conditions that determine the maintenance of this monodominant forest.     

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.