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Climate affects neighbour-induced changes in leaf chemical defences and tree diversity-herbivory relationships.

ABSTRACT: Associational resistance theory predicts that insect herbivory decreases with increasing tree diversity in forest ecosystems. However, the generality of this effect and its underlying mechanisms are still debated, particularly since evidence has accumulated that climate may influence the direction and strength of the relationship between diversity and herbivory.We quantified insect leaf herbivory and leaf chemical defences (phenolic compounds) of silver birch Betula pendula in pure and mixed plots with different tree species composition across 12 tree diversity experiments in different climates. We investigated whether the effects of neighbouring tree species diversity on insect herbivory in birch, that is, associational effects, were dependent on the climatic context, and whether neighbour-induced changes in birch chemical defences were involved in associational resistance to insect herbivory.We showed that herbivory on birch decreased with tree species richness (i.e. associational resistance) in colder environments but that this relationship faded as mean annual temperature increased.Birch leaf chemical defences increased with tree species richness but decreased with the phylogenetic distinctiveness of birch from its neighbours, particularly in warmer and more humid environments.Herbivory was negatively correlated with leaf chemical defences, particularly when birch was associated with closely related species. The interactive effect of tree diversity and climate on herbivory was partially mediated by changes in leaf chemical defences.Our findings confirm that tree species diversity can modify the leaf chemistry of a focal species, hence its quality for herbivores. They further stress that such neighbour-induced changes are dependent on climate and that tree diversity effects on insect herbivory are partially mediated by these neighbour-induced changes in chemical defences.

SUBMITTER: Poeydebat C

PROVIDER: S-EPMC7610350 | biostudies-literature |

REPOSITORIES: biostudies-literature

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Project description:The diversity of plant neighbors commonly results in direct, bottom-up effects on herbivore ability to locate their host, and in indirect effects on herbivores involving changes in plant traits and a top-down control by their enemies. Yet, the relative contribution of bottom-up and top-down forces remains poorly understood. We also lack knowledge on the effect of abiotic constraints such as summer drought on the strength and direction of these effects. We measured leaf damage on pedunculate oak (Quercus robur), alone or associated with birch, pine or both in a long-term tree diversity experiment (ORPHEE), where half of the plots were irrigated while the other half remained without irrigation and received only rainfall. We tested three mechanisms likely to explain the effects of oak neighbors on herbivory: (1) Direct bottom-up effects of heterospecific neighbors on oak accessibility to herbivores, (2) indirect bottom-up effects of neighbors on the expression of leaf traits, and (3) top-down control of herbivores by predators. Insect herbivory increased during the growth season but was independent of neighbor identity and irrigation. Specific leaf area, leaf toughness, and thickness varied with neighbor identity while leaf dry matter content or C:N ratio did not. When summarized in a principal component analysis (PCA), neighbor identity explained 87% of variability in leaf traits. PCA axes partially predicted herbivory. Despite greater rates of attack on dummy caterpillars in irrigated plots, avian predation, and insect herbivory remained unrelated. Our study suggests that neighbor identity can indirectly influence insect herbivory in mixed forests by modifying leaf traits. However, we found only partial evidence for these trait-mediated effects and suggest that more attention should be paid to some unmeasured plant traits such as secondary metabolites, including volatile organic compounds, to better anticipate the effects of climate change on plant-insect interactions in the future.

Project description:The degree to which plant pathogen infestation occurs in a host plant is expected to be strongly influenced by the level of species diversity among neighbouring host and non-host plant species. Since pathogen infestation can negatively affect host plant performance, it can mediate the effects of local biodiversity on ecosystem functioning.We tested the effects of tree diversity and the proportion of neighbouring host and non-host species with respect to the foliar fungal pathogens of Tilia cordata and Quercus petraea in the Kreinitz tree diversity experiment in Germany. We hypothesized that fungal pathogen richness increases while infestation decreases with increasing local tree diversity. In addition, we tested whether fungal pathogen richness and infestation are dependent on the proportion of host plant species present or on the proportion of particular non-host neighbouring tree species.Leaves of the two target species were sampled across three consecutive years with visible foliar fungal pathogens on the leaf surface being identified macro- and microscopically. Effects of diversity among neighbouring trees were analysed: (i) for total fungal species richness and fungal infestation on host trees and (ii) for infestation by individual fungal species.We detected four and five fungal species on T. cordata and Q. petraea, respectively. High local tree diversity reduced (i) total fungal species richness and infestation of T. cordata and fungal infestation of Q. petraea and (ii) infestation by three host-specialized fungal pathogen species. These effects were brought about by local tree diversity and were independent of host species proportion. In general, host species proportion had almost no effect on fungal species richness and infestation. Strong effects associated with the proportion of particular non-host neighbouring tree species on fungal species richness and infestation were, however, recorded.Synthesis. For the first time, we experimentally demonstrated that for two common forestry tree species, foliar fungal pathogen richness and infestation depend on local biodiversity. Thus, local tree diversity can have positive impacts on ecosystem functioning in managed forests by decreasing the level of fungal pathogen infestation.

Project description:1.Insect herbivory can strongly affect ecosystem processes, and its relationship with plant diversity is a central topic in biodiversity-functioning research. However, very little is known about this relationship from complex ecosystems dominated by long-lived individuals, such as forests, especially over gradients of high plant diversity.2.We analysed insect herbivory on saplings of 10 tree and shrub species across 27 forest stands differing in age and tree species richness in an extraordinarily diverse subtropical forest ecosystem in China. We tested whether plant species richness significantly influences folivory in these highly diverse forests or whether other factors play a more important role at such high levels of phytodiversity.3.Leaf damage was assessed on 58 297 leaves of 1284 saplings at the end of the rainy season in 2008, together with structural and abiotic stand characteristics.4.Species-specific mean damage of leaf area ranged from 3% to 16%. Herbivory increased with plant species richness even after accounting for potentially confounding effects of stand characteristics, of which stand age-related aspects most clearly covaried with herbivory. Intraspecific density dependence or other abiotic factors did not significantly influence overall herbivory across forest stands.5.Synthesis.The positive herbivory-plant diversity relationship indicates that effects related to hypotheses of resource concentration, according to which a reduction in damage by specialized herbivores might be expected as host plant concentration decreases with increasing plant diversity, do not seem to be major determinants for overall herbivory levels in our phytodiverse subtropical forest ecosystem. We discuss the potential role of host specificity of dominant herbivores, which are often expected to show a high degree of specialization in many (sub)tropical forests. In the forest system we studied, a much higher impact of polyphagous species than traditionally assumed might explain the observed patterns, as these species can profit from a broad dietary mix provided by high plant diversity. Further testing is needed to experimentally verify this assumption.

Project description:Bivariate relationships between plant tissue nutrient concentration have largely been studied across broad environmental scales regardless of their covariation with soil and climate. Comparing leaf and branch wood concentrations of C, Ca, K, Mg, N, Na, and P for trees growing in tropical forests in Amazonia and Australia we found that the concentrations of most elements varied with sampling location, but with foliar and branch woody tissues varying from site to site in different ways. Using a Mixed Effect Model (MEM) approach it was further found that relationships between branch and leaf concentrations within individual plots differed in terms of both slope and/or significance to the ordinary least squares (OLS) estimates for most elements. Specifically, using MEM we found that within plots only K and Mg were correlated across organs, but with the K cross-organ intercept estimates varying significantly between sites. MEM analyses further showed that within-plot wood density variations were also negatively related to wood K and Na, suggesting a potentially important role for these cations in water transport and/or storage in woody tissues. The OLS method could not detect significant correlations in any of the above cases. By contrast, although Ca, N, and P leaf and wood tissue concentrations showed similar patterns when individual elements were compared across sites, MEM analyses suggested no consistent association within sites. Thus, for all these three elements, strong within-tree scaling relationships were inferred when data were analyzed across sites using OLS, even though there was no relationship within individual sites. Thus (as for Ca, N, and P) not only can a pooling of data across sites result in trait (co)variations attributable to the environment potentially being incorrectly attributed solely to the species and/or individual (the so-called "ecological fallacy"), but in some cases (as was found here for K and Na) the opposite can also sometimes occur with significant within-site covariations being obscured by large site-site variations. We refer to the latter phenomenon as "environmental obfuscation."

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Climate affects neighbour-induced changes in leaf chemical defences and tree diversity-herbivory relationships.

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