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.

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.

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.

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.

Foliar stable isotope ratios of carbon and nitrogen in boreal forest plants exposed to long-term pollution from the nickel-copper smelter at Monchegorsk, Russia.

Long-term exposure to primary air pollutants, such as sulphur dioxide (SO2) and nitrogen oxides (NOx), alters the structure and functions of forest ecosystems. Many biochemical and biogeochemical processes discriminate against the heavier isotopes in a mixture; thus, the values of δ13C and δ15N (i.e. the ratio of stable isotopes 13C to 12C and that of 15 N to 14 N, respectively) may give insights into changes in ecosystem processes and identify the immediate drivers of these changes. We studied sources of variation in the δ13C and δ15N values in the foliage of eight boreal forest C3 plants at 10 sites located at the distance of 1-40 km from the Monchegorsk nickel-copper smelter in Russia. From 1939‒2019, this smelter emitted over 14,000,000 metric tons (t) of SO2, 250,000 t of metals, primarily nickel and copper, and 140,000 t of NOx. The δ13C value in evergreen plants and the δ15N value in all plants increased near the smelter independently of the plant mycorrhizal type. We attribute the pollution-related increase in the foliar δ13C values of evergreen species mainly to direct effects of SO2 on stomatal conductance, in combination with pollution-related water stress, which jointly override the potential opposite effect of increasing ambient CO2 concentration on δ13C values. Stomatal uptake of NOx and root uptake of 15N-enriched organic N compounds and NH4+ may explain the increased foliar δ15N values and elevated foliar N concentrations, especially in the evergreen trees (Pinus sylvestris), close to Monchegorsk, where the soil inorganic N supply is reduced due to the impact of long-term SO2 and heavy metal emissions on plant biomass. We conclude that, despite the uncertainties in interpreting δ13C and δ15N responses to pollution, the Monchegorsk smelter has imposed and still imposes a great impact on C and N cycling in the surrounding N-limited subarctic forest ecosystems.

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.

Changes in Biomass and Diversity of Soil Macrofauna along a Climatic Gradient in European Boreal Forests.

Latitudinal gradients allow insights into the factors that shape ecosystem structure and delimit ecosystem processes, particularly climate. We asked whether the biomass and diversity of soil macrofauna in boreal forests change systematically along a latitudinal gradient spanning from 60° N to 69° N. Invertebrates (3697 individuals) were extracted from 400 soil samples (20 × 20 cm, 30 cm depth) collected at ten sites in 2015-2016 and then weighed and identified. We discovered 265 species living in soil and on the soil surface; their average density was 0.486 g d·w·m-2. The species-level diversity decreased from low to high latitudes. The biomass of soil macrofauna showed no latitudinal changes in early summer but decreased towards the north in late summer. This variation among study sites was associated with the decrease in mean annual temperature by ca 5 °C and with variation in fine root biomass. The biomass of herbivores and fungivores decreased towards the north, whereas the biomass of detritivores and predators showed no significant latitudinal changes. This variation in latitudinal biomass patterns among the soil macrofauna feeding guilds suggests that these guilds may respond differently to climate change, with poorly understood consequences for ecosystem structure and functions.

Climate shapes the spatiotemporal variation in color morph diversity and composition across the distribution range of Chrysomela lapponica leaf beetle.

Color polymorphism offers rich opportunities for studying the eco-evolutionary mechanisms that drive the adaptations of local populations to heterogeneous and changing environments. We explored the color morph diversity and composition in a Chrysomela lapponica leaf beetle across its entire distribution range to test the hypothesis that environmental and climatic variables shape spatiotemporal variation in the phenotypic structure of a polymorphic species. We obtained information on 13 617 specimens of this beetle from museums, private collections, and websites. These specimens (collected from 1830-2020) originated from 959 localities spanning 33° latitude, 178° longitude, and 4200 m altitude. We classified the beetles into five color morphs and searched for environmental factors that could explain the variation in the level of polymorphism (quantified by the Shannon diversity index) and in the relative frequencies of individual color morphs. The highest level of polymorphism was found at high latitudes and altitudes. The color morphs differed in their climatic requirements; composition of colour morphs was independent of the geographic distance that separated populations but changed with collection year, longitude, mean July temperature and between-year temperature fluctuations. The proportion of melanic beetles, in line with the thermal melanism hypothesis, increased with increasing latitude and altitude and decreased with increasing climate seasonality. Melanic morph frequencies also declined during the past century, but only at high latitudes and altitudes where recent climate warming was especially strong. The observed patterns suggest that color polymorphism is especially advantageous for populations inhabiting unpredictable environments, presumably due to the different climatic requirements of coexisting color morphs.

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.

Decline of Eulia ministrana (Lepidoptera: Tortricidae) in polluted habitats is not accompanied by phenotypic stress responses.

Environmental pollution is currently identified as one of the major drivers of rapid decline of insect populations, and this finding has revitalized interest in insect responses to pollution. We tested the hypothesis that the pollution-induced decline of insect populations can be predicted from phenotypic stress responses expressed as morphological differences between populations inhabiting polluted and unpolluted sites. We explored populations of the brassy tortrix Eulia ministrana in subarctic forests along an environmental disturbance gradient created by long-lasting severe impacts of aerial emissions of the copper-nickel smelter in Monchegorsk, northwestern Russia. We used pheromone traps to measure the population densities of this leafrolling moth and to collect specimens for assessment of three morphological stress indices: size, forewing melanization, and fluctuating asymmetry in wing venation. Wing length of E. ministrana increased by 10%, and neither forewing melanization nor fluctuating asymmetry changed from the unpolluted forest to the heavily polluted industrial barren. However, the population density of E. ministrana decreased 5 to 10 fold in the same pollution gradient. Thus, none of the studied potential morphological stress indicators signaled vulnerability of E. ministrana to environmental pollution and/or to pollution-induced environmental disturbance. We conclude that insect populations can decline without any visible signs of stress. The use of morphological proxies of insect fitness to predict the consequences of human impact on insect populations is therefore risky until causal relationships between these proxies and insect abundance are deciphered.

Parasitoids indicate major climate-induced shifts in arctic communities.

Climatic impacts are especially pronounced in the Arctic, which as a region is warming twice as fast as the rest of the globe. Here, we investigate how mean climatic conditions and rates of climatic change impact parasitoid insect communities in 16 localities across the Arctic. We focus on parasitoids in a widespread habitat, Dryas heathlands, and describe parasitoid community composition in terms of larval host use (i.e., parasitoid use of herbivorous Lepidoptera vs. pollinating Diptera) and functional groups differing in their closeness of host associations (koinobionts vs. idiobionts). Of the latter, we expect idiobionts-as being less fine-tuned to host development-to be generally less tolerant to cold temperatures, since they are confined to attacking hosts pupating and overwintering in relatively exposed locations. To further test our findings, we assess whether similar climatic variables are associated with host abundances in a 22 year time series from Northeast Greenland. We find sites which have experienced a temperature rise in summer while retaining cold winters to be dominated by parasitoids of Lepidoptera, with the reverse being true for the parasitoids of Diptera. The rate of summer temperature rise is further associated with higher levels of herbivory, suggesting higher availability of lepidopteran hosts and changes in ecosystem functioning. We also detect a matching signal over time, as higher summer temperatures, coupled with cold early winter soils, are related to high herbivory by lepidopteran larvae, and to declines in the abundance of dipteran pollinators. Collectively, our results suggest that in parts of the warming Arctic, Dryas is being simultaneously exposed to increased herbivory and reduced pollination. Our findings point to potential drastic and rapid consequences of climate change on multitrophic-level community structure and on ecosystem functioning and highlight the value of collaborative, systematic sampling effort.

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.

Biases in estimation of insect herbivory from herbarium specimens.

Information regarding plant damage by insects in the past is essential to explore impacts of climate change on herbivory. We asked whether insect herbivory measured from herbarium specimens reflects the levels of herbivory occurring in nature at the time of herbarium sampling. We compared herbivory measurements between herbarium specimens collected by botany students and ecological samples collected simultaneously by the authors by a method that minimized unconscious biases, and asked herbarium curators to select one of two plant specimens, which differed in leaf damage, for their collections. Both collectors and curators generally preferred specimens with lesser leaf damage, but the strength of this preference varied among persons. In addition, the differences in measured leaf damage between ecological samples and herbarium specimens varied among plant species and increased with the increase in field herbivory. Consequently, leaf damage in herbarium specimens did not correlate with the actual level of herbivory. We conclude that studies of herbarium specimens produce biased information on past levels of herbivory, because leaf damage measured from herbarium specimens not only underestimates field herbivory, but it is not proportional to the level of damage occurring in nature due to multiple factors that cannot be controlled in data analysis.

Climate warming leads to decline in frequencies of melanic individuals in subarctic leaf beetle populations.

Intraspecific diversity buffers populations from deleterious impacts of environmental change. Nevertheless, the consequences of climate warming for phenotypic and genetic diversity within populations and species remain poorly understood. The goal of our study was to explore among-year variations in the phenotypic structure of populations and their relationships with climate variability and population dynamics. We analysed multiyear (1992-2018) data on colour morph frequencies within populations of the leaf beetle, Chrysomela lapponica, from multiple sites in the Kola Peninsula (northwestern Russia). We observed a strong decline in the proportion of dark (melanic) morphs among overwintered beetles during the study period; this decline was consistent across all study sites. Using model selection procedures, we explained declines in the dark morph of overwintered beetles by increases in minimum spring (May-June) daily temperatures. Other climatic characteristics, pollution load, and beetle population density were unrelated to variation in colour morph frequencies. Among newly emerged beetles (August), dark morph frequencies also decreased with an increase in average spring temperatures, but were unrelated to mean temperatures during the larval development period (July). These results suggest that the two-fold decline in dark morph frequencies during the past 26 years has been driven by the 2.5 °C increase in spring temperatures, most likely because dark males lose the mating advantages over light males that they obtain during cold springs. The continued loss of dark morphs and related decrease in within-population diversity may render leaf beetle populations more vulnerable to future environmental changes, in particular to those expressed in extreme weather fluctuations. Our study demonstrates that declines in within-population diversity are already underway in subarctic areas, and that these declines are likely driven by climate warming.

Opposite latitudinal patterns for bird and arthropod predation revealed in experiments with differently colored artificial prey.

The strength of biotic interactions is generally thought to increase toward the equator, but support for this hypothesis is contradictory. We explored whether predator attacks on artificial prey of eight different colors vary among climates and whether this variation affects the detection of latitudinal patterns in predation. Bird attack rates negatively correlated with model luminance in cold and temperate environments, but not in tropical environments. Bird predation on black and on white (extremes in luminance) models demonstrated different latitudinal patterns, presumably due to differences in prey conspicuousness between habitats with different light regimes. When attacks on models of all colors were combined, arthropod predation decreased, whereas bird predation increased with increasing latitude. We conclude that selection for prey coloration may vary geographically and according to predator identity, and that the importance of different predators may show contrasting patterns, thus weakening the overall latitudinal trend in top-down control of herbivorous insects.

Temperature and herbivory, but not pollution, affect fluctuating asymmetry of mountain birch leaves: Results of 25-year monitoring around the copper­nickel smelter in Monchegorsk, northwestern Russia.

Fluctuating asymmetry (FA), which is defined as the magnitude of the random deviations from a symmetrical shape, reflects developmental instability and is commonly assumed to increase under environmental and genetic stress. We monitored the leaf FA of mountain birch, Betula pubescens subsp. czerepanovii, from 1993 to 2017 in individually marked trees at 21 sites around the copper­nickel smelter at Monchegorsk, and we then analysed the results with respect to spatial and temporal variation in pollution, climate and background insect herbivory. Responses of leaf FA to different stressors were stressor specific: FA did not correlate with pollution load, it decreased significantly with an increase in June air temperature and it increased slightly but significantly with an increase in the previous-year leaf damage due to defoliating and leafmining insects. Our findings suggest that climate warming is unlikely to impose stress on the explored mountain birch populations, but even small increases in insect herbivory may adversely affect birch trees. However, these conclusions, since they are based on an observational study, should be viewed as tentative until confirmed by controlled experiments. We also demonstrated that the use of non-blinded measurements, which are prone to confirmation bias, was the primary reason for the earlier report of an increase in birch leaf FA near the Monchegorsk smelter. We hope that our findings will promote a wide use of blinded methods in ecological research and that they will contribute to debunking the myth that plant leaf FA consistently increases with increases in environmental pollution.

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.

Local Insect Damage Reduces Fluctuating Asymmetry in Next-year's Leaves of Downy Birch.

Insect herbivory imposes stress on host plants. This stress may cause an increase in leaf fluctuating asymmetry (FA), which is defined as the magnitude of the random deviations from a symmetrical leaf shape. We tested the hypothesis that differences in leaf FA among individual shoots of downy birch, Betula pubescens, are at least partly explained by local damage caused by insects in the previous year. Unexpectedly, we found that in the year following the damage imposed by miners, leafrollers and defoliators, damaged birch shoots produced leaves with lower FAs compared to shoots from the same tree that had not been damaged by insects. This effect was consistent among the different groups of insects investigated, but intra-species comparisons showed that statistical significance was reached only in shoots that had been damaged by the birch leaf roller, Deporaus betulae. The detected decrease in leaf FA in the year following the damage agrees with the increases in shoot performance and in antiherbivore defence. The present results indicate that within-plant variation in leaf FA may have its origin in previous-year damage by insects, and that FA may influence the current-year's distribution of herbivory.

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.