There is increasing evidence that geographic and climatic clines drive the patterns of plant defence allocation and defensive strategies. We quantified early growth rate and both constitutive and inducible chemical defences of 18 Pinaceae species in a common greenhouse environment and assessed their defensive allocation with respect to each species' range across climatic gradients spanning 31° latitude and 2300 m elevation. Constitutive defences traded-off with induced defences, and these defensive strategies were associated with growth rate such that slow-growing species invested more in constitutive defence, whereas fast-growing species invested more in inducible defence. The position of each pine species along this trade-off axis was in turn associated with geography; moving poleward and to higher elevations, growth rate and inducible defences decreased, while constitutive defence increased. These geographic patterns in plant defence were most strongly associated with variation in temperature. Climatic and geographical clines thus act as drivers of defence profiles by mediating the constraints imposed by trade-offs, and this dynamic underlays global patterns of defence allocation.

Summary1. Production of antiherbivore chemical defences is generally assumed to be costly in terms of fitness, although some studies have failed to detect such costs. A convincing explanation is that the expression of fitness costs depends on environmental conditions such as nutrient availability. 2. We performed a greenhouse experiment with 33 half-sib families in order to study the phenotypic plasticity of constitutive and methyl jasmonate-induced chemical defences to soil phosphorus (P) availability, the existence of genetic trade-offs (costs) between growth and the production of those defences and the extent to which P availability may modulate the expression of those costs. 3. We measured some proxies of vegetative fitness (primary growth, secondary growth and total biomass), plant reserves (soluble sugars and starch) and the concentration of quantitative chemical defences (diterpene content in the stem, total polyphenolics and condensed tannins in the needles). 4. Phosphorus availability had a considerable effect, both on the allocation of resources to constitutive and induced defences and on the expression of vegetative costs associated with those chemical defences. Constitutive investment in chemical defences was greater under P-limited conditions for all studied traits. Inducibility of foliar phenolic compounds was greater under P-limited conditions, and it was strongly constrained under high P availability. Availability of P did not affect the inducibility of stem diterpenes. 5. All defensive traits showed significant genetic variation, with different levels of genetic control in constitutive and induced modes, and genetic variation in their inducibility. We found significant negative genetic correlations (i.e. trade-offs) between growth and defensive investment, but costs of chemical defences emerged only in P-limited conditions. Vegetative costs of constitutive defences were detected for stem diterpenes but not for needle phenolics, while costs of induced defences were found for leaf phenolics but not for stem diterpenes. 6. Synthesis. Our results indicate that P availability controls the production of chemical defences in this pine species, influencing the resource allocation to constitutive defences, the inducibility of those defences and the emergence of related vegetative costs. Phosphorus availability thus appears as a major driver in the evolution of pine resistance to insects and a potential factor in maintaining genetic variation in defences.

Classic research on elevational gradients in plant-herbivore interactions holds that insect herbivore pressure is stronger under warmer, less seasonal climates characteristic of low elevations, and that this in turn selects for increased defence in low-(relative to high-) elevation plants. However, recent work has questioned this paradigm, arguing that it overly simplifies the ecological complexity in which plant-insect herbivore interactions are embedded along elevational gradients. Numerous biotic and abiotic factors vary with elevation, and their simultaneous influences are the focus of current work on elevational gradients in insect herbivory and plant defences. The present review 1) synthesizes current knowledge on elevational gradients in plant-insect herbivore interactions; 2) critically analyses research gaps and highlights recent advances that contribute to filling these gaps; and 3) outlines new research opportunities to uncover underlying mechanisms and build towards a unified theory on elevational gradients. We conclude that the next generation of studies should embrace community complexity -including multi-trophic dynamics and the multivariate nature of plant defence -and to do so by combining observational data, manipulative experiments and emerging analytical tools.

A rich body of theory has been developed to predict the effects of plant diversity on communities at higher trophic levels and the mechanisms underpinning such effects. However, there are currently a number of key gaps in knowledge that have hindered the development of a predictive framework of plant diversity effects on consumers. For instance, we still know very little about how the magnitude of plant trait variation (e.g. intraspecific vs. inter-specific), as well as the identity and combined effects of plant, herbivore and natural enemy traits, mediate plant diversity effects on consumers. Moreover, the fine-scale mechanisms (e.g. changes in consumer behaviour or recruitment responses) underlying such diversity effects in many cases remain elusive or have been overlooked. In addition, most studies of plant diversity effects on associated consumers have been developed under a static, unidirectional (bottom-up) framework of effects on herbivores and predators without taking into account the potential for dynamic feedbacks across trophic levels. Here we seek to address these key gaps in knowledge as well as to capitalize on recent advances and emerging frameworks in plant biodiversity research. In doing so, we provide new insights as well as recommendations which will stimulate new research and advance this field of study. IntroductionThe consequences of plant intra-specific and inter-specific diversity on associated faunas have been the focus of much research over the last decade (e.g. [1 ,2-8,9 ]). Studies have found that plant diversity positively influences arthropod diversity and abundance [3,4,8,10], and alters plant-arthropod and arthropod-arthropod interactions [3,7,11,12]. These findings emphasize that conserving and manipulating plant diversity in natural and managed systems, respectively, is crucial for maintaining ecosystem function [13][14][15].A rich body of theory has been developed to predict the effects of plant diversity on communities at higher trophic levels ([16-22], see Box 1). Despite this vast collection of theory behind plant diversity effects on associated faunas and the large number of empirical studies conducted thus far, formal evaluations of the mechanisms behind the observed patterns have been developed in natural communities (but see [20] for e.g. in agricultural systems). In addition, there are also a number of key gaps in knowledge that have hindered the development of a predictive framework of plant diversity effects on higher trophic levels ( Figure 1). For example, we generally ignore how the magnitude of variation in plant traits (e.g. interspecific vs. intra-specific diversity) or the identity (including independent and interactive effects of multiple traits) of plant traits determines such effects. Similarly, many studies have lacked an explicit evaluation of the influence of herbivore traits such as diet breadth, mobility and feeding behaviour, and the underlying mechanisms for diversity effects on consumer abundance or behaviour remain elusive (e.g. effects of diversit...

Ecological research conducted over the past five decades has shown that increasing tree species richness at forest stands can improve tree resistance to insect pest damage. However, the commonality of this finding is still under debate. In this review, we provide a quantitative assessment (i.e., a meta-analysis) of tree diversity effects on insect herbivory and discuss plausible mechanisms underlying the observed patterns. We provide recommendations and working hypotheses that can serve to lay the groundwork for research to come. Based on more than 600 study cases, our quantitative review indicates that insect herbivory was, on average, lower in mixed forest stands than in pure stands, but these diversity effects were contingent on herbivore diet breadth and tree species composition. In particular, tree species diversity mainly reduced damage of specialist insect herbivores in mixed stands with phylogenetically distant tree species. Overall, our findings provide essential guidance for forest pest management. Expected final online publication date for the Annual Review of Entomology, Volume 66 is January 11, 2020. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

1. Ecological theory indicates that warmer and more stable climates should result in stronger biotic interactions. Therefore, plant species growing at lower elevations and experiencing greater herbivore pressure should invest in higher levels of defences than those at higher elevations. Nonetheless, there are a number of studies that have found no effect of elevational gradients on plant defensive traits. Several factors might explain the lack of consistency for the altitude-defence relationships, including (i) the reduction of all defensive traits into one measure of resistance; (ii) not considering plant defence as the simultaneous expression of several defensive traits; and (iii) not considering the relative influence of biotic (e.g. herbivory) and abiotic (e.g. climate and soil conditions) factors associated with the ecological gradient.2. Here, we present a comprehensive test of the effects of elevation and its associated biotic and abiotic factors on the individual and simultaneous expression of constitutive direct and indirect defences and their inducibility (i.e. expression of defences after herbivore attack). Specifically, we estimated climatic and soil variables and measured herbivore damage and constitutive and jasmonic acid-induced glucosinolate levels in the leaves as a proxy for direct defences, and volatile emission as a proxy for indirect defences in 16 Cardamine species naturally growing along the steep elevational gradient of the Alps. 3. Within a phylogenetic comparative framework, we found that species growing at lower elevations invested more in the simultaneous inducibility of both direct and indirect defences, whereas species growing at higher elevations invested more in constitutive direct defences. Although we found strong elevational gradients in herbivory and climatic and soil variables, these biotic and abiotic factors only partially explained elevational patterns in plant defences. 4. Synthesis. These results highlight that the complex regulation of multiple defence traits strongly vary across elevational gradients and build towards a better understanding of the multiple mechanisms underlying trait evolution and species interactions along ecological gradients.

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

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