Does spatial variation in insect herbivory match variations in plant quality? A meta-analysis.

Variation in herbivore pressure has often been predicted from patterns in plant traits considered as antiherbivore defences. Here, we tested whether spatial variation in field insect herbivory is associated with the variation in plant quality by conducting a meta-analysis of 223 correlation coefficients between herbivory levels and the expression of selected plant traits. We found no overall correlation between herbivory and either concentrations of plant secondary metabolites or values of physical leaf traits. This result was due to both the large number of low correlations and the opposing directions of high correlations in individual studies. Field herbivory demonstrated a significant association only with nitrogen: herbivore pressure increased with an increase in nitrogen concentration in plant tissues. Thus, our meta-analysis does not support either theoretical prediction, i.e., that plants possess high antiherbivore defences in localities with high herbivore pressure or that herbivory is low in localities where plant defences are high. We conclude that information about putative plant defences is insufficient to predict plant losses to insects in field conditions and that the only bottom-up factor shaping spatial variation in insect herbivory is plant nutritive value. Our findings stress the need to improve a theory linking plant putative defences and herbivory.

Meta-analysis of elevational changes in the intensity of trophic interactions: Similarities and dissimilarities with latitudinal patterns.

The premise that the intensity of biotic interactions decreases with increasing latitudes and elevations is broadly accepted; however, whether these geographical patterns can be explained within a common theoretical framework remains unclear. Our goal was to identify the general pattern of elevational changes in trophic interactions and to explore the sources of variation among the outcomes of individual studies. Meta-analysis of 226 effect sizes calculated from 134 publications demonstrated a significant but interaction-specific decrease in the intensity of herbivory, carnivory and parasitism with increasing elevation. Nevertheless, this decrease was not significant at high latitudes and for interactions involving endothermic organisms, for herbivore outbreaks or for herbivores living within plant tissues. Herbivory similarly declined with increases in latitude and elevation, whereas carnivory showed a fivefold stronger decrease with elevation than with latitude and parasitism increased with latitude but decreased with elevation. Thus, although these gradients share a general pattern and several sources of variation in trophic interaction intensity, we discovered important dissimilarities, indicating that elevational and latitudinal changes in these interactions are partly driven by different factors. We conclude that the scope of the latitudinal biotic interaction hypothesis cannot be extended to incorporate elevational gradients.

Latitudinal gradient in the intensity of biotic interactions in terrestrial ecosystems: Sources of variation and differences from the diversity gradient revealed by meta-analysis.

The Latitudinal Biotic Interaction Hypothesis (LBIH) states that the intensity of biotic interactions increases from high to low latitudes. This hypothesis, which may partly explain latitudinal gradients in biodiversity, remains hotly debated, largely due to variable outcomes of published studies. We used meta-analysis to identify the scope of the LBIH in terrestrial ecosystems. For this purpose, we explored the sources of variation in the strength of latitudinal changes in herbivory, carnivory and parasitism (119 publications) and compared these gradients with gradients in the diversity of the respective groups of animals (102 publications). Overall, both herbivory and carnivory decreased towards the poles, while parasitism increased. The latitudinal gradient in herbivory and carnivory was threefold stronger above 50-60° than at lower latitudes and was significant due to interactions involving ectothermic consumers, studies using standardised prey (i.e. prey lacking local anti-predator adaptations) and studies aimed at testing LBIH. The poleward decrease in biodiversity did not differ between ectothermic and endothermic animals or among climate zones and was fourfold stronger than decrease in herbivory and carnivory. The discovered differences between the gradients in biotic interactions and biodiversity suggest that these two global macroecological patterns are likely shaped by different factors.

Changes in plant collection practices from the 16th to 21st centuries: implications for the use of herbarium specimens in global change research.

BACKGROUND AND AIMS: Herbaria were recently advertised as reliable sources of information regarding historical changes in plant traits and biotic interactions. To justify the use of herbaria in global change research, we asked whether the characteristics of herbarium specimens have changed during the past centuries and whether these changes were due to shifts in plant collection practices. METHODS: We measured nine characteristics from 515 herbarium specimens of common European trees and large shrubs collected from 1558 to 2016. We asked botanists to rank these specimens by their scientific quality, and asked artists to rank these specimens by their beauty. KEY RESULTS: Eight of 11 assessed characteristics of herbarium specimens changed significantly during the study period. The average number of leaves in plant specimens increased 3-fold, whereas the quality of specimen preparation decreased. Leaf size negatively correlated with leaf number in specimens in both among-species and within-species analyses. The proportion of herbarium sheets containing plant reproductive structures peaked in the 1850s. The scientific value of herbarium specimens increased until the 1700s, but then did not change, whereas their aesthetic value showed no systematic trends. CONCLUSIONS: Our findings strongly support the hypothesis that many characteristics of herbarium specimens have changed systematically and substantially from the 16th to 21st centuries due to changes in plant collection and preservation practices. These changes may both create patterns which could be erroneously attributed to environmental changes and obscure historical trends in plant traits. The utmost care ought to be taken to guard against the possibility of misinterpretation of data obtained from herbarium specimens. We recommend that directional changes in characters of herbarium specimens which occurred during the past 150‒200 years, primarily in specimen size and in the presence of reproductive structures, are accounted for when searching for the effects of past environmental changes on plant traits.

Seasonal variations in bird selection pressure on prey colouration.

The direction and strength of selection for prey colouration by predators vary in space and time and depend on the composition of the predator community. We tested the hypothesis that bird selection pressure on prey colouration changes through the season due to changes in the proportion of naïve juvenile individuals in the bird community, because naïve and educated birds differ in their responses to prey colours. Bird predation on caterpillar-shaped plasticine models in two boreal forest sites increased sevenfold from early summer to mid-summer, and the time of this increase coincides with the fledging of juvenile birds. In early summer, cryptic (black and green) models were attacked at fivefold higher rates compared with conspicuous (red and yellow) models. By contrast, starting from fledging time, cryptic and conspicuous models were attacked at similar rates, hinting at a lower selectivity by naïve juvenile birds compared with educated adult birds. Cryptic models exposed in a group together with conspicuous models were attacked by birds at a threefold lower rate than cryptic models exposed singly, thus supporting the aposematic commensalism hypothesis. However, this effect was not observed in mid- and late summer, presumably due to the lack of avoidance of conspicuous prey by the juvenile birds. We conclude that selection pressure on prey colouration weakens considerably when naïve birds dominate in the community, because the survival advantages of aposematic colouration are temporarily lost for both the conspicuous and their neighbouring cryptic prey.

Predation and parasitism on herbivorous insects change in opposite directions in a latitudinal gradient crossing a boreal forest zone.

The latitudinal biotic interaction hypothesis (LBIH) predicts that the strength of various biotic interactions decreases from low to high latitudes. Inconsistency between studies testing this hypothesis may result from variations among different types of interactions and among study systems. Therefore, exploration of multiple interactions within one system would help to disentangle latitudinal patterns across individual interactions and to evaluate latitudinal changes in the overall impact of enemies on prey. We tested the prediction based on the LBIH that the pressure of natural enemies on herbivorous insects decreases with increase in latitude across the boreal forest zone. We also asked whether the impacts of major groups of these enemies exhibit similar latitudinal patterns and whether these patterns are consistent across study years. In 10 forest sites located from 60°N to 69°N in Northern Europe, each summer, from 2016 to 2019, we measured (a) mortality of three groups of leafmining insects caused by birds, ants, parasitoids and unknown factors, (b) bird attacks on caterpillar-shaped plasticine models and (c) birch foliar damage caused by defoliators and leafminers. Latitudinal patterns in both insect herbivory on birch and top-down pressure on herbivorous insects varied considerably and inconsistently among the four study years, so that only some of the year-specific correlations with latitude were statistically significant. Nevertheless, meta-analysis combining correlations across years, preys and enemies revealed general decreases in predation by birds (on both natural and model prey) and ants, but an increase in parasitism rates, from low to high latitudes. We found that the direction of latitudinal changes in the strength of biotic interactions was interaction-specific: predation and herbivory supported LBIH, whereas parasitism exhibited an opposite trend. Consequently, the overall impact of natural enemies on herbivorous insects did not change with latitude and was therefore an unlikely reason for the poleward decrease in herbivory observed in our gradient. Considerable among-year variation in the strength of the latitudinal patterns in all the studied interactions suggests that this variation is a widespread phenomenon.

Top-down factors contribute to differences in insect herbivory between saplings and mature trees in boreal and tropical forests.

Ontogenetic changes in herbivory are generally not consistent with ontogenetic changes in defensive traits of woody plants. This inconsistency suggests that other factors may affect ontogenetic trajectories in herbivory. We tested the hypothesis that top-down factors contribute to differences in foliar losses to insects between juvenile and mature trees in tropical and boreal forests. We used artificial caterpillars made of modelling clay to compare predation rates between saplings and mature trees of two common forest species, Siparuna guianensis in Brazil (tropical site) and Betula pubescens in Finland (boreal site). Leaf area losses to chewing insects in saplings were 2.5-fold higher than in mature trees in both species. Physical plant defences (measured as specific leaf area, SLA) did not differ between saplings and mature trees in the boreal forest, whereas in the tropical forest, SLA was greater in saplings than in mature trees. Attack rates on the model prey by birds were higher in the boreal forest, whereas attack rates by arthropod predators were higher in the tropical forest. Overall, predation rates on model prey were consistently higher on mature trees than on saplings at both sites, but in the boreal site, this pattern was primarily driven by birds, whereas in the tropical site, it was primarily driven by arthropod predators. We conclude that the effect of predation on herbivorous insects may considerably contribute to ontogenetic differences in herbivory, but the relative roles of different predatory groups and of top-down and bottom-up factors may vary between environments.

Ambient temperatures differently influence colour morphs of the leaf beetle Chrysomela lapponica: Roles of thermal melanism and developmental plasticity.

We asked whether ambient temperatures can affect morph frequencies within a subarctic population of the polymorphic leaf beetle Chrysomela lapponica through thermal melanism and/or developmental plasticity. Body temperature increased faster in beetles of dark morph than in beetles of light morph under exposure to artificial irradiation. Dark males ran faster than light males in both field and laboratory experiments, and this difference decreased with increasing ambient air temperature, from significant at 10 °C to non-significant at 20 °C and 26 °C. On cold days (6-14 °C), significantly more dark males than light males were found on their host plants in copula (40.8% and 27.3% respectively); on warm days (15-22 °C) this difference disappeared. Light females produced twice as many eggs as dark females; this difference did not depend on the ambient temperature. The proportion of dark morphs in the progenies of pairs with one dark parent was twice as high as that in the progenies of pairs in which both parents were light, and this proportion was greater when larvae developed at low (10 and 15 °C) than at high (20 and 25 °C) temperatures. We conclude that low temperatures may increase the frequencies of dark morphs in C. lapponica populations due to both the mating advantages of dark males over light males and developmental plasticity. Variation in frequencies of low-fecund dark morphs in the population, caused by among-year differences in temperature together with density-dependent selection, may contribute to the evolutionary dynamics of the colour polymorphism and may influence abundance fluctuations in these leaf beetle populations.

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.

Strategies of chemical anti-predator defences in leaf beetles: is sequestration of plant toxins less costly than de novo synthesis?

The evolution of defensive traits is driven both by benefits gained from protection against enemies and by costs of defence production. We tested the hypothesis that specialisation of herbivores on toxic host plants, accompanied by the ability to acquire plant defensive compounds for herbivore defence, is favoured by the lower costs of sequestration compared to de novo synthesis of defensive compounds. We measured physiological costs of chemical defence as a reduction in larval performance in response to repeated removal of secretions (simulating predator attack) and compared these costs between five species synthesising defences de novo and three species sequestering salicylic glucosides (SGs) from their host plants. Experiments simulating low predator pressure revealed no physiological costs in terms of survival, weight and duration of development in any of study species. However, simulation of high predation caused reduction in relative growth rate in Chrysomela lapponica larvae producing autogenous defences more frequently, than in larvae sequestering SGs. Still meta-analysis of combined data showed no overall difference in costs of autogenous and sequestered defences. However, larvae synthesising their defences de novo demonstrated secretion-conserving behaviour, produced smaller amounts of secretions, replenished them at considerably lower rates and employed other types of defences (regurgitation, evasion) more frequently when compared to sequestering larvae. These latter results provide indirect evidence for biosynthetic constraints for amounts of defensive secretions produced de novo, resulting in low defence effectiveness. Lifting these constraints by sequestration may have driven some leaf beetle lineages toward sequestration of plant allelochemicals as the main defensive strategy.

Sap-feeding insects on forest trees along latitudinal gradients in northern Europe: a climate-driven patterns.

Knowledge of the latitudinal patterns in biotic interactions, and especially in herbivory, is crucial for understanding the mechanisms that govern ecosystem functioning and for predicting their responses to climate change. We used sap-feeding insects as a model group to test the hypotheses that the strength of plant-herbivore interactions in boreal forests decreases with latitude and that this latitudinal pattern is driven primarily by midsummer temperatures. We used a replicated sampling design and quantitatively collected and identified all sap-feeding insects from four species of forest trees along five latitudinal gradients (750-1300 km in length, ten sites in each gradient) in northern Europe (59 to 70°N and 10 to 60°E) during 2008-2011. Similar decreases in diversity of sap-feeding insects with latitude were observed in all gradients during all study years. The sap-feeder load (i.e. insect biomass per unit of foliar biomass) decreased with latitude in typical summers, but increased in an exceptionally hot summer and was independent of latitude during a warm summer. Analysis of combined data from all sites and years revealed dome-shaped relationships between the loads of sap-feeders and midsummer temperatures, peaking at 17 °C in Picea abies, at 19.5 °C in Pinus sylvestris and Betula pubescens and at 22 °C in B. pendula. From these relationships, we predict that the losses of forest trees to sap-feeders will increase by 0-45% of the current level in southern boreal forests and by 65-210% in subarctic forests with a 1 °C increase in summer temperatures. The observed relationships between temperatures and the loads of sap-feeders differ between the coniferous and deciduous tree species. We conclude that climate warming will not only increase plant losses to sap-feeding insects, especially in subarctic forests, but can also alter plant-plant interactions, thereby affecting both the productivity and the structure of future forest ecosystems.

Changes in the background losses of woody plant foliage to insects during the past 60 years: are the predictions fulfilled?

The existing scenarios generally predict that herbivory will increase with climate warming. An analysis of the published data on the background foliar losses of woody plants to insects in natural ecosystems across the globe from 1952 to 2013 provided no support for this hypothesis. We detected no temporal trend in herbivory within the temperate climate zone and a significant decrease in herbivory in the tropics. From 1964 to 1990, herbivory in the tropics was 39% higher than in the temperate region, but these differences disappeared by the beginning of the 2000s. Thus, environmental changes have already disturbed one of the global ecological patterns--the decrease in herbivory with latitude--by affecting ecosystem processes differently in tropical and temperate climate zones.

Inter- and Intrapopulation Variability in the Composition of Larval Defensive Secretions of Willow-Feeding Populations of the Leaf Beetle Chrysomela lapponica.

We explored the inter- and intrapopulation variability in the larval defensive chemistry of the leaf beetle Chrysomela lapponica with respect to the salicylic glycoside (SG) content of its host species. Secretions of larvae from three populations associated in nature with SG-poor willows contained nearly twice as many components and 40-fold higher concentrations of autogenously produced isobutyrates and 2-methylbutyrates than secretions of larvae from three populations associated with SG-rich willows, which in turn had 200-fold higher concentrations of host-derived salicylaldehyde. Reciprocal transfer experiments showed that the larvae from populations associated with SG-rich willows did not produce appreciable amounts of butyrates on either SG-rich or SG-poor willows, while populations feeding on several SG-poor willow species retained the ability for efficient sequestration of SGs, along with their ability to produce high amounts of isobutyrates and 2-methylbutyrates. Only the populations associated with SG-poor willows demonstrated among-family variation in the composition of defensive secretion and differential responses of individual families to willows with alternative SG levels, which can be seen as the prerequisites for shifting to novel hosts. These non-specialized populations show a dual defensive strategy, which corresponds to the ancestral state of this species, while populations that fully depend on host-derived toxins (feeding on SG-rich willows) or have lost the ability to produce salicylaldehyde (feeding on birches) are most deviant from the ancestral state. The results indicate that defensive strategies may differ between populations within a species, and suggest that this variation reduces extinction risks and maintains the high ecological diversity and wide distribution of C. lapponica.

Predation risk estimated on live and artificial insect prey follows different patterns.

Models mimicking prey organisms are increasingly used in ecological studies, including testing fundamental ecological and evolutionary theories. The general consensus is that predation risk estimated on artificial models may not quantitatively correspond to predation pressure on live prey, but it still can be used in various comparisons. We tested whether the use of live and artificial prey reveals the same patterns of variation in predation risk. We exposed live prey (blowfly larvae and puparia) and plasticine models of blowfly puparia in two boreal forest sites, both openly and in ant- and bird-exclusion treatments, and we quantified attacks by both avian and invertebrate predators. Bird attack rates were always higher on live puparia than on their plasticine models, but the magnitude of this difference declined from 8.4-fold in early summer to 2-fold in mid- and late-summer. We attribute these changes to different responses to prey by experienced adult birds that dominate the bird communities in early summer versus explorative juvenile birds that are abundant later in the season. Invertebrate daily predation rates on maggots decreased from 56% in early summer to 28% in late summer, but invertebrate attacks on plasticine models showed no seasonal changes. Overall, invertebrate predation on maggots was 67-fold greater than their predation on models. Observations showed that wood ants did not attack plasticine models and did not leave on them any damage marks. Estimates based on artificial prey indicate a much greater role of bird predation than invertebrate predation, while estimates based on live prey suggest the opposite pattern. Thus, using live and artificial prey may lead to different conclusions about relative importance of different predator groups in a locality. Moreover, for both avian and invertebrate predators, predation risk based on artificial and live prey shows different seasonal changes and may potentially demonstrate different spatial patterns.

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.

Sources of variation in plant responses to belowground insect herbivory: a meta-analysis.

Growing interest in belowground herbivory and the remarkable diversity of the accumulated information on this topic inspired us to quantitatively explore the variation in the outcomes of individual studies. We conducted a meta-analysis of 85 experimental studies reporting the effects of root-feeding insect herbivores (36 species) on plants (75 species). On average, belowground herbivory led to a 36.3% loss of root biomass, which was accompanied by a reduction in aboveground growth (-16.3%), photosynthesis (-11.7%) and reproduction (-15.5%). The effects of root herbivory on aboveground plant characteristics were significant in agricultural and biological control studies, but not in studies of natural systems. Experiments conducted in controlled environments yielded larger effects on plants than field experiments, and infestation experiments resulted in more severe effects than removal studies employing natural levels of herbivory. Simulated root herbivory led to greater aboveground growth reductions than similar root loss imposed by insect feeding. External root chewers caused stronger detrimental effects than sap feeders or root borers; specialist herbivores imposed milder adverse effects on plants than generalists. Woody plants suffered from root herbivory more than herbaceous plants, although root loss was similar in these two groups. Evergreen woody plants responded to root herbivory more strongly than deciduous woody plants, and grasses suffered from root herbivory more than herbs. Environmental factors such as drought, poor nutrient supply, among-plant competition, and aboveground herbivory increased the adverse effects of root damage on plants in an additive manner. In general, plant tolerance to root herbivores is lower than tolerance to defoliating aboveground herbivores.

Elevational changes in insect herbivory on woody plants in six mountain ranges of temperate Eurasia: Sources of variation.

Current theory predicts that the intensity of biotic interactions, particularly herbivory, decreases with increasing latitude and elevation. However, recent studies have revealed substantial variation in both the latitudinal and elevational patterns of herbivory. This variation is often attributed to differences in study design and the type of data collected by different researchers. Here, we used a similar sampling protocol along elevational gradients in six mountain ranges, located at different latitudes within temperate Eurasia, to uncover the sources of variation in elevational patterns in insect herbivory on woody plant leaves. We discovered a considerable variation in elevational patterns among different mountain ranges; nevertheless, herbivory generally decreased with increasing elevation at both the community-wide and individual plant species levels. This decrease was mostly due to openly living defoliators, whereas no significant association was detected between herbivory and elevation among insects living within plant tissues (i.e., miners and gallers). The elevational decrease in herbivory was significant for deciduous plants but not for evergreen plants, and for tall plants but not for low-stature plants. The community-wide herbivory increased with increases in both specific leaf area and leaf size. The strength of the negative correlation between herbivory and elevation increased from lower to higher latitudes. We conclude that despite the predicted overall decrease with elevation, elevational gradients in herbivory demonstrate considerable variation, and this variation is mostly associated with herbivore feeding habits, some plant traits, and latitude of the mountain range.

Diversity but Not Overall Abundance of Moths and Butterflies (Insecta: Lepidoptera) Decreases around Two Arctic Polluters.

Alarming reports on the rapid decline of insects during the past decades call for the exploration of potential drivers of this process. Here, we test the hypothesis that the overall abundance and diversity of moths and butterflies (Lepidoptera) decrease under the impact of industrial pollution in the fragile arctic environment. For this purpose, experienced collectors netted adult Lepidoptera at five tundra sites located 0.5 to 45.3 km from the ore-roasting plant in Zapolyarnyy and at five forest sites located 1.4 to 37.8 km from the copper-nickel smelter at Nikel, in the Murmansk region of Russia. The analysis of the 100 samples collected from 2003 to 2008 and containing 2312 individuals of 122 species revealed that the diversity of Lepidoptera declined significantly near both of these polluters due to both decreases in species richness and changes in the abundance of individual species, whereas the overall abundance of moths and butterflies was independent of the pollution load. These patterns did not differ between Nikel and Zapolyarnyy, and they were consistent with patterns previously found near the copper-nickel smelter at Monchegorsk. The abundances of Lepidoptera species showed variable changes along pollution gradients, from significantly negative to significantly positive, but individual species showed similar density changes around these three polluters. Disproportional increases in the abundance of a few pollution-tolerant species change the community structure and explain why the overall abundance of moths and butterflies does not decline even in localities experiencing extreme loads of sulphur dioxide and heavy metals.

Insect herbivory increases from forest to alpine tundra in Arctic mountains.

Current theory holds that the intensity of biotic interactions decreases with increases in latitude and elevation; however, empirical data demonstrate great variation in the direction, strength, and shape of elevational changes in herbivory. The latitudinal position of mountains may be one important source of this variation, but the acute shortage of data from polar mountains hampers exploration of latitude effects on elevational changes in herbivory. Here, we reduce this knowledge gap by exploring six elevation gradients located in three Arctic mountain ranges to test the prediction that a decrease in herbivory occurs with increasing elevation from forest to alpine tundra. Across the 10 most abundant evergreen and deciduous woody plant species, relative losses of foliage to insect herbivores were 2.2-fold greater at the highest elevations (alpine tundra) than in mid-elevation birch woodlands or low-elevation coniferous forests. Plant quality for herbivores (quantified by specific leaf area) significantly decreased with elevation across all studied species, indicating that bottom-up factors were unlikely to shape the observed pattern in herbivory. An experiment with open-top chambers established at different elevations showed that even a slight increase in ambient temperature enhances herbivory in Arctic mountains. Therefore, we suggest that the discovered increase in herbivory with elevation is explained by higher temperatures at the soil surface in open habitats above the tree line compared with forests at lower elevations. This explanation is supported by the significant difference in elevational changes in herbivory between low and tall plants: herbivory on low shrubs increased fourfold from forest to alpine sites, while herbivory on trees and tall shrubs did not change with elevation. We suggest that an increase in herbivory with an increase in elevation is typical for high-latitude mountains, where inverse temperature gradients, especially at the soil surface, are common. Verification of this hypothesis requires further studies of elevational patterns in herbivory at high latitudes.

Recovery of moth and butterfly (Lepidoptera) communities in a polluted region following emission decline.

Environmental pollution is one of the major drivers of the present-day decline in global biodiversity. However, the links between the effects of industrial pollution on insect communities and the underlying species-specific responses remain poorly understood. We explored the spatial pattern in insect communities by analysing 581 samples of moths and butterflies (containing 25,628 individuals of 345 species) collected along a strong pollution gradient in subarctic Russia, and we recorded temporal changes in these communities during the pollution decline that occurred from 1992 to 2006. In the 1990s, the diversity of the Lepidoptera community was positively correlated with the distance from the copper-nickel smelter at Monchegorsk. The overall abundance of Lepidoptera did not change along the pollution gradient, although the abundance of many species decreased with increasing pollution. The responses of each individual species to pollution were associated with its life history traits. The abundances of monophagous species that fed inside live plant tissues and hibernated as imagoes or pupae were not affected by pollution, whereas the abundances of oligophagous and polyphagous species that fed externally on plants and hibernated as larvae generally declined near the smelter. Substantial decreases in aerial emissions from the smelter between 1992 and 2006 resulted in an increase in the diversity of moths and butterflies in severely polluted habitats, whereas their overall abundance did not change. This recovery of the Lepidoptera community occurred due to the reappearance of rare species that had been previously extirpated by pollution and was observed despite the lack of any signs of recovery of the vegetation in the heavily polluted sites. We conclude that the recovery trajectories of insect communities following emission control can be predicted from studies of their changes along spatial pollution gradients by using space-for-time substitution.