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Elevational gradients affect the production of plant secondary metabolites through changes in both biotic and abiotic conditions. Previous studies have suggested both elevational increases and decreases in host-plant chemical defences. We analysed the correlation of alkaloids and polyphenols with elevation in a community of nine Ficus species along a continuously forested elevational gradient in Papua New Guinea. We sampled 204 insect species feeding on the leaves of these hosts and correlated their community structure to the focal compounds. Additionally, we explored species richness of folivorous mammals along the gradient. When we accounted for Ficus species identity, we found a general elevational increase in flavonoids and alkaloids. Elevational trends in non-flavonol polyphenols were less pronounced or showed non-linear correlations with elevation. Polyphenols responded more strongly to changes in temperature and humidity than alkaloids. The abundance of insect herbivores decreased with elevation, while the species richness of folivorous mammals showed an elevational increase. Insect community structure was affected mainly by alkaloid concentration and diversity. Although our results show an elevational increase in several groups of metabolites, the drivers behind these trends likely differ. Flavonoids may provide figs with protection against abiotic stressors. In contrast, alkaloids affect insect herbivores and may provide protection against mammalian herbivores and pathogens. Concurrent analysis of multiple compound groups alongside ecological data is an important approach for understanding the selective landscape that shapes plant defences.

Resource specialization is a key concept in ecology, but it is unexpectedly difficult to parameterize. Differences in resource availability, sampling effort and abundances preclude comparisons of incompletely sampled biotic interaction webs. Here, we extend the distance-based specialization index (DSI) that measures trophic specialization by taking resource phylogenetic relatedness and availability into account into a rescaled version, DSI*. It is a versatile metric of specialization that expands considerably the scope and applicability, hence the usefulness, of DSI. The new metric also accounts for differences in abundance and sampling effort of consumers, which enables robust comparisons among distinct guilds of consumers. It also provides an abundance threshold for the reliability of the metric for rare species, a very desirable property given the difficulty of assessing any aspect of rare species accurately. We apply DSI* to an extensive dataset on interactions between insect herbivores from four folivorous guilds and their host plants in Papua New Guinean rainforests. We demonstrate that DSI*, contrary to the original DSI, is largely independent of sample size and weakly and non-linearly related with several host specificity measures that do not adjust for plant phylogeny. Thus, DSI* provides further insights into host specificity patterns; moreover, it is robust to the number and phylogenetic diversity of plant species selected to be sampled for herbivores. DSI* can be used for a broad range of comparisons of distinct feeding guilds, geographical locations and ecological conditions. This is a key advance in elucidating the interaction structure and evolution of highly diversified systems.

A long-term goal in evolutionary ecology is to explain the incredible diversity of insect herbivores and patterns of host plant use in speciose groups like tropical Lepidoptera. Here, we used standardized food-web data, multigene phylogenies of both trophic levels and plant chemistry data to model interactions between Lepidoptera larvae (caterpillars) from two lineages (Geometridae and Pyraloidea) and plants in a species-rich lowland rainforest in New Guinea. Model parameters were used to make and test blind predictions for two hectares of an exhaustively sampled forest. For pyraloids, we relied on phylogeny alone and predicted 54% of species-level interactions, translating to 79% of all trophic links for individual insects, by sampling insects from only 15% of local woody plant diversity. The phylogenetic distribution of host-plant associations in polyphagous geometrids was less conserved, reducing accuracy. In a truly quantitative food web, only 40% of pair-wise interactions were described correctly in geometrids. Polyphenol oxidative activity (but not protein precipitation capacity) was important for understanding the occurrence of geometrids (but not pyraloids) across their hosts. When both foliar chemistry and plant phylogeny were included, we predicted geometrid-plant occurrence with 89% concordance. Such models help to test macroevolutionary hypotheses at the community level., (© 2017 The Author(s).)

Insects tend to feed on related hosts. The phylogenetic composition of host plant communities thus plays a prominent role in determining insect specialization, food web structure, and diversity. Previous studies showed a high preference of insect herbivores for congeneric and confamilial hosts suggesting that some levels of host plant relationships may play more prominent role that others. We aim to quantify the effects of host phylogeny on the structure of quantitative plant-herbivore food webs. Further, we identify specific patterns in three insect guilds with different life histories and discuss the role of host plant phylogeny in maintaining their diversity. We studied herbivore assemblages in three temperate forests in Japan and the Czech Republic. Sampling from a canopy crane, a cherry picker and felled trees allowed a complete census of plant-herbivore interactions within three 0·1 ha plots for leaf chewing larvae, miners, and gallers. We analyzed the effects of host phylogeny by comparing the observed food webs with randomized models of host selection. Larval leaf chewers exhibited high generality at all three sites, whereas gallers and miners were almost exclusively monophagous. Leaf chewer generality dropped rapidly when older host lineages (5-80 myr) were collated into a single lineage but only decreased slightly when the most closely related congeneric hosts were collated. This shows that leaf chewer generality has been maintained by feeding on confamilial hosts while only a few herbivores were shared between more distant plant lineages and, surprisingly, between some congeneric hosts. In contrast, miner and galler generality was maintained mainly by the terminal nodes of the host phylogeny and dropped immediately after collating congeneric hosts into single lineages. We show that not all levels of host plant phylogeny are equal in their effect on structuring plant-herbivore food webs. In the case of generalist guilds, it is the phylogeny of deeper plant lineages that drives the food web structure whereas the terminal relationships play minor roles. In contrast, the specialization and abundance of monophagous guilds are affected mainly by the terminal parts of the plant phylogeny and do not generally reflect deeper host phylogeny., (© 2017 The Authors. Journal of Animal Ecology © 2017 British Ecological Society.)

We studied a community of frugivorous Lepidoptera in the lowland rainforest of Papua New Guinea. Rearing revealed 122 species represented by 1,720 individuals from 326 woody plant species. Only fruits from 52% (171) of the plant species sampled were attacked. On average, Lepidoptera were reared from 1 in 89 fruits and a kilogram of fruit was attacked by 1.01 individuals. Host specificity of Lepidoptera was notably low: 69% (33) of species attacked plants from >1 family, 8% (4) fed on single family, 6% (3) on single genus and 17% (8) were monophagous. The average kilogram of fruits was infested by 0.81 individual from generalist species (defined here as feeding on >1 plant genus) and 0.07 individual from specialist species (feeding on a single host or congeneric hosts). Lepidoptera preferred smaller fruits with both smaller mesocarp and seeds. Large-seeded fruits with thin mesocarp tended to host specialist species whereas those with thick, fleshy mesocarp were often infested with both specialist and generalist species. The very low incidence of seed damage suggests that pre-dispersal seed predation by Lepidoptera does not play a major role in regulating plant populations via density-dependent mortality processes outlined by the Janzen-Connell hypothesis.

Plant-insect food webs tend to be dominated by interactions resulting from diffuse co-evolution between plants and multiple lineages of herbivores rather than by reciprocal co-evolution and co-cladogenesis. Plants therefore require defence strategies effective against a broad range of herbivore species. In one extreme, plants could develop a single universal defence effective against all herbivorous insects, or tailor-made strategies for each herbivore species. The evolution and ecology of plant defence has to be studied with entire insect assemblages, rather than small subsets of pairwise interactions. The present study examines whether specialists and generalists in three coexisting insect lineages, forming the leaf-chewing guild, respond uniformly to plant phylogeny, secondary metabolites, nutrient content and mechanical antiherbivore defences of their hosts, thus permitting universal plant defence strategies against specialized and generalist folivorous insects from various taxa. The extensive data on folivorous assemblages comprising three insect orders and 193 species are linked with plant phylogeny, secondary chemistry (salicylates, flavonoids and tannins), leaf morphological traits [specific leaf area (SLA) and trichome coverage], nutrient (C : N) content and growth form of eight willow (Salix) and one aspen (Populus) species growing in sympatry. Generalists responded to overall host plant chemistry and trichomes, whilst specialists responded to host plant phylogeny and secondary metabolites that are unique to willows and that are capable of being utilized as an antipredator protection. We did not find any significant impact of other plant traits, that is SLA, C : N ratio, flavonoids, tannins and growth form, on the composition of leaf-chewing communities. Our results show that the response to plant traits is differential among specialists and generalists. This finding constrains the ability of plants to develop defensive traits universally effective against herbivores and may lead to diversification of plant defensive mechanisms into several complementary syndromes, required for effective protection against generalists and specialists from multiple insect taxa comprising most leaf-chewing assemblages. These results point to the necessity of broad studies of plant-herbivore interactions, across multiple insect taxa and guilds., (© 2015 The Authors. Journal of Animal Ecology © 2015 British Ecological Society.)

Understanding variation in resource specialization is important for progress on issues that include coevolution, community assembly, ecosystem processes, and the latitudinal gradient of species richness. Herbivorous insects are useful models for studying resource specialization, and the interaction between plants and herbivorous insects is one of the most common and consequential ecological associations on the planet. However, uncertainty persists regarding fundamental features of herbivore diet breadth, including its relationship to latitude and plant species richness. Here, we use a global dataset to investigate host range for over 7,500 insect herbivore species covering a wide taxonomic breadth and interacting with more than 2,000 species of plants in 165 families. We ask whether relatively specialized and generalized herbivores represent a dichotomy rather than a continuum from few to many host families and species attacked and whether diet breadth changes with increasing plant species richness toward the tropics. Across geographic regions and taxonomic subsets of the data, we find that the distribution of diet breadth is fit well by a discrete, truncated Pareto power law characterized by the predominance of specialized herbivores and a long, thin tail of more generalized species. Both the taxonomic and phylogenetic distributions of diet breadth shift globally with latitude, consistent with a higher frequency of specialized insects in tropical regions. We also find that more diverse lineages of plants support assemblages of relatively more specialized herbivores and that the global distribution of plant diversity contributes to but does not fully explain the latitudinal gradient in insect herbivore specialization.

Understanding global drivers of changes in species interactions is vital, both in terms of improving our knowledge of fundamental large-scale ecology, and in order to predict and protect against human-driven changes to ecosystems. Here we present the LifeWebs project, which aims to collate existing data on interaction networks in order to understand their large-scale patterns. Initially we will collate data relating to plant–herbivore interactions, and this article is a call for contributions. We outline the kinds of data in which we are interested, the procedure for making contributions, and what contributors can expect in terms of co-authorship on subsequent papers and access to the database for conducting their own analyses.

Insect herbivores have the potential to consume large amounts of plant tissue in tropical forests, but insectivorous vertebrates effectively control their abundances, indirectly increasing plant fitness accordingly. Despite several studies already sought understanding of the top-down effects on arthropod community structure and herbivory, such studies of trophic cascades in old tropics are underrepresented, and little attention was paid to top-down forces in various habitats. Therefore, we examine how flying insectivorous vertebrates (birds and bats) impact arthropods and, consequently, affect herbivore damage of leaves in forest habitats in Papua New Guinea. In a 3-month long predator exclosure experiment conducted at four study sites across varying elevation and successional stage, we found that vertebrate predators reduced arthropod density by ∼52%. In addition, vertebrate predators decreased the mean body size of arthropods by 26% in leaf chewers and 47% in non-herbivorous arthropods but had only a small effect on mesopredators and sap suckers. Overall, the exclusion of vertebrate predators resulted in a ~ 41% increase in leaf damage. Our results, across different types of tropical forests in Papua New Guinea, demonstrate that flying vertebrate insectivores have a crucial impact on plant biomass, create a selective pressure on larger and non-predatory prey individuals and they prey partition with mesopredators. [ABSTRACT FROM AUTHOR]

The community composition of insect herbivores is largely shaped by host–plant phylogeny and functional traits. These effects differ between herbivores due at least in part to varying levels of diet specialisation. Distinguishing between herbivores with differing specialisation is therefore necessary to understand the roles of host phylogeny and traits in shaping complex insect communities.We surveyed the complete plant–caterpillar food web in a 0.2 ha plot of a lowland cool‐temperate deciduous forest. We measured leaf nutrients, physical traits and polyphenols of the hostplants and analysed the effects of leaf traits and phylogeny on Lepidoptera caterpillars with differing levels of host specialisation.We sampled 190 species from 16 433 individual caterpillars on 20 host plant species. These included 102 species of specialists using confamilial hostplant species, 17 species of conservative generalists using allofamilial but phylogenetically close hostplant species and 71 species of non‐conservative generalists using phylogenetically distant hostplant species.The species composition of non‐conservative generalists associated with the polyphenol protein precipitation capacity and overall leaf trait dissimilarity; conservative generalists were weakly associated with polyphenol oxidative activity. In contrast, the composition of specialist assemblages showed no correlation with leaf traits.Our results demonstrate that host phylogeny and traits play variable roles in structuring communities of herbivores, based on the phylogenetic specialisation of herbivores. Understanding the factors that shape the community structures of various herbivores traditionally classified as 'generalists' is important as they account for a large proportion of herbivore species while showing differential responses to traits of hosts. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Both the abiotic environment and the composition of animal and plant communities change with elevation. For mutualistic species, these changes are expected to result in altered partner availability, and shifts in context-dependent benefits for partners. To test these predictions, we assessed the network structure of terrestrial ant-plant mutualists and how the benefits to plants of ant inhabitation changed with elevation in tropical forest in Papua New Guinea. At higher elevations, ant-plants were rarer, species richness of both ants and plants decreased, and the average ant or plant species interacted with fewer partners. However, networks became increasingly connected and less specialized, more than could be accounted for by reductions in ant-plant abundance. On the most common ant-plant, ants recruited less and spent less time attacking a surrogate herbivore at higher elevations, and herbivory damage increased. These changes were driven by turnover of ant species rather than by within-species shifts in protective behaviour. We speculate that reduced partner availability at higher elevations results in less specialized networks, while lower temperatures mean that even for ant-inhabited plants, benefits are reduced. Under increased abiotic stress, mutualistic networks can break down, owing to a combination of lower population sizes, and a reduction in context-dependent mutualistic benefits. [ABSTRACT FROM AUTHOR]

Throughout the course of their evolution, plants have acquired a wide range of chemical and mechanical defenses to protect against herbivores. Ehrlich & Raven's coevolutionary theory suggests that this diversification of defensive traits is driven by the strong impact of novel traits on insect herbivores. However, the impact of plant defenses on insects is difficult to compare between related plant species due to variation in environmental and biotic conditions. We standardized these factors as far as possible by analyzing the effects of chemical and mechanical defensive traits on insects in a local community of 11 Salicaceae species growing in sympatry, and their leaf-chewing herbivores. Defensive traits (salicylates, flavonoids, tannins, trichomes, and leaf toughness) were generally not inter-correlated, with the exception of a negative correlation between salicylates and trichomes. The content of salicylates, a novel group of defensive metabolites in the Salicaceae, was correlated with low herbivore diversity and high host specificity. Despite these effects, the phylogeny of the studied species shows loss of salicylates in some Salix species instead of their further diversification. This could be due to salicylates not decreasing the overall abundance of herbivores, despite accounting for up to 22% of the dry leaf mass and therefore being costly. The defense of low-salicylate willow species is thus probably maintained by other defensive traits, such as trichomes. Our study shows that the balance between costs and benefits of defensive traits is not necessarily in favor of novel compounds and illustrates a process, which may lead to the reduction in a defensive trait. [ABSTRACT FROM AUTHOR]

A shift in the composition of the herbivore guild in the invasive range is expected to select for plants with a higher competitive ability, a lower regrowth capacity and a lower investment in defence. We show here that parallel evolution took place in three geographically distinct invasive regions that differed significantly in climatic conditions. This makes it most likely that indeed the shifts in herbivore guilds were causal to the evolutionary changes. We studied competitive ability and regrowth of invasive and native Jacobaea vulgaris using an intraspecific competition set-up with and without herbivory. Without herbivores invasive genotypes have a higher competitive ability than native genotypes. The invasive genotypes were less preferred by the generalist Mamestra brassicae but more preferred by the specialist Tyria jacobaeae, consequently their competitive ability was significantly increased by the first and reduced by the latter. Invasive genotypes showed a lower regrowth ability in both herbivore treatments. [ABSTRACT FROM AUTHOR]

Humans are altering the global distributional ranges of plants, while their co-evolved herbivores are frequently left behind. Native herbivores often colonise non-native plants, potentially reducing invasion success or causing economic loss to introduced agricultural crops. We developed a predictive model to forecast novel interactions and verified it with a data set containing hundreds of observed novel plant-insect interactions. Using a food network of 900 native European butterfly and moth species and 1944 native plants, we built an herbivore host-use model. By extrapolating host use from the native herbivore-plant food network, we accurately forecasted the observed novel use of 459 non-native plant species by native herbivores. Patterns that governed herbivore host breadth on co-evolved native plants were equally important in determining non-native hosts. Our results make the forecasting of novel herbivore communities feasible in order to better understand the fate and impact of introduced plants. [ABSTRACT FROM AUTHOR]

Phylogenetic ecology has identified patterns of diversity in communities that may find explanation in trophic interactions, and yet there have been few attempts to directly relate such patterns among trophic levels. Density-dependent processes involving pests and pathogens, for example, have been invoked to account for plant community phylogenetic patterns, but relatively little is known about how plant relatedness might affect community structure at other trophic levels. We examined the degree to which the abundance of herbivores in a rain forest community is explained by the phylogeny and functional traits of host plants. We destructively sampled all stems ≥5 cm diameter in two 1-ha plots of New Guinea primary and secondary lowland forest to test predicted relationships between herbivore abundance and plant resources. We analyzed per-tree caterpillar and leaf miner abundance, total leaf biomass (kg), percentage of immature foliage, specific leaf area (cm2/g), leaf nitrogen content (percentage of dry mass), and presence of exudates in the context of a plant community phylogeny estimated from DNA barcodes. Apart from nitrogen content and exudates, neither plant resources nor herbivore abundance showed evidence of phylogenetic signal in our community sample. The plant traits we measured could account for only 30% and 16% of variation in caterpillar and leaf miner abundance, respectively, among individual trees. Leaf biomass was a stronger predictor of herbivore abundance than either resource quality (leaf nitrogen content) or palatability (percentage of immature foliage, specific leaf area). The primary importance of resource quantity was also observed at the plant species level in analyses of species means and phylogenetic generalized least-squares regression. Plant relatedness did not account for much variation in herbivore abundance, but significant effects of exudates and leaf nitrogen content on caterpillar abundance illustrate how conserved traits at one trophic level may influence community-wide patterns at another. [ABSTRACT FROM AUTHOR]

1. Beta diversity of plant–herbivore food webs, defined as turnover of trophic interactions between the food webs, represents a potentially useful extension of traditional studies of plant and herbivore beta diversity as it integrates spatial turnover of plant and herbivore species with changes in herbivore host plant preferences. 2. Beta diversity of plant–herbivore food webs can be partitioned into four components, corresponding to the turnover of plant–herbivore interactions due to change in (i) both plant and herbivore species, (ii) plant, but not herbivore, species, (iii) herbivore, but not plant, species, and (iv) herbivore host preferences between food webs. These components can be quantified using a range of existing beta diversity indices. 3. Relative magnitude of plant and herbivore beta diversity of plant–herbivore food webs can be analysed as an outcome of herbivore (i) host specificity, (ii) ability to track host plant populations by dispersal, and (iii) response to environmental conditions, competitors and enemies. The relative importance of these factors in tropical ecosystems remains to be determined. [ABSTRACT FROM AUTHOR]