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Premise of research. Herbivore damage is generally higher in the sun than in the shade. The influence of shade tolerance on herbivory has been mainly related to the fact that light-demanding plant species show lower defense investments—and hence higher herbivory—than shade-tolerant species. The pattern of increased herbivory with increasing light availability may take different forms, depending on the drivers of herbivore damage across the light gradient: (1) herbivory pressure alone, (2) herbivory pressure together with plant resistance associated with shade tolerance, or (3) the two drivers seen in scenario 2 added to a differential effect of light availability on plant resistance depending on shade tolerance. Elucidating the specific patterns behind the general pattern of increased herbivory with increasing light availability may contribute to understanding and aid prediction of herbivory patterns in forest communities under natural or anthropogenic disturbance. Methodology. In a field study with 15 tree species distributed across the light gradient in a temperate rainforest (1.6%–26.2% canopy openness), we evaluated the relationship between canopy openness and seedling herbivory, with 755 seedlings surveyed for herbivore damage. Specifically, we evaluated the consistency between the observed herbivory–canopy openness relationship and the three scenarios described above. Pivotal results. We found steeper slopes for the herbivory–canopy openness relationship in tree species with lower shade tolerance; that is, we verified scenario 3. Conclusions. Shade tolerance determined the slope of the herbivory–canopy openness relationship across the light gradient. This finding expands our understanding of the recognized influence of shade tolerance on the interplay between herbivory and the light environment. [ABSTRACT FROM AUTHOR]

Despite the ubiquitous presence of fungal endophytes in woody plants, mechanisms underlying variation in foliar fungal endophyte communities are poorly understood. Given that endophytes in woody plants are predominantly horizontally transmitted, fungal endophyte colonization of foliar tissues is likely to be influenced by plant resistance traits. Here, we evaluated the association between leaf resistance traits and colonization and community composition of horizontally transmitted endophytes (HTE) in 10 dominant trees species in a temperate rainforest in Southern Chile. Because resistance traits might restrict colonization of HTE, it was hypothesized that host trees with similar leaf resistance traits would show similar HTE communities.We measured leaf traits with reported roles in plant resistance against pathogens and/or herbivores: cell wall polysaccharides, leaf toughness, flavonoids, anthocyanins, terpenoids and chitinases. We also determined variation in fungal read counts, number of operational taxonomic units (OTUs), diversity and community structure across tree species. Foliar fungal endophyte communities were characterized using Illumina MiSeq sequencing of tagged amplicons of the ITS1 region.Fungal read counts were significantly different among host trees, while number of OTUs and diversity were similar among them. Multivariate analyses showed that community composition of endophytic fungi significantly differed among hosts, and that variation in cell wall, flavonoids, anthocyanins and terpenoids was associated with differences in HTE communities. We found that host trees sharing leaf resistance traits harbour similar HTE communities. Fungal read counts decreased with increasing content of cell wall polysaccharides, while endophyte diversity decreased with increasing anthocyanin levels. Fungal read counts, in contrast, were positively associated with terpenoids. Additionally, culture‐based data indicated that some HTE strains inhibited the growth of the common fungal pathogen Botrytis cinerea.Synthesis. Overall, we found that leaf resistance traits may influence fungal endophyte colonization and community composition in tree species, likely through deterrent effects of structural and chemical traits. We suggest that endophytic fungi that overcome resistance traits might play a role in plant protection. [ABSTRACT FROM AUTHOR]

There is limited evidence regarding the adaptive value of plant functional traits in contrasting light environments. It has been suggested that changes in these traits in response to light availability can increase herbivore susceptibility. We tested the adaptive value of plant functional traits linked with carbon gain in contrasting light environments and also evaluated whether herbivores can modify selection on these traits in each light environment. In a temperate rainforest, we examined phenotypic selection on functional traits in seedlings of the pioneer tree Aristotelia chilensis growing in sun (canopy gap) and shade (forest understory) and subjected to either natural herbivory or herbivore exclusion. We found differential selection on functional traits depending on light environment. In sun, there was positive directional selection on photosynthetic rate and relative growth rate (RGR), indicating that selection favors competitive ability in a high-resource environment. Seedlings with high specific leaf area (SLA) and intermediate RGR were selected in shade, suggesting that light capture and conservative resource use are favored in the understory. Herbivores reduced the strength of positive directional selection acting on SLA in shade. We provide the first demonstration that natural herbivory rates can change the strength of selection on plant ecophysiological traits, that is, attributes whose main function is resource uptake. Research addressing the evolution of shade tolerance should incorporate the selective role of herbivores. [ABSTRACT FROM AUTHOR]

Climbing plants are a key component of rainforests, but mechanistic approaches to their distribution and abundance are scarce. In a southern temperate rainforest, we addressed whether the dominance of climbing plants across light environments is associated with the expression of ecophysiological traits. In mature forest and canopy gaps, we measured leaf size, specific leaf area, photosynthetic rate, and dark respiration in six of the most abundant woody vines. Mean values of traits and their phenotypic change (%) between mature forest and canopy gaps were predictor variables. Leaf size and specific leaf area were not significantly associated with climbing plant dominance. Variation in gas-exchange traits between mature forest and canopy gaps explained, at least partly, the dominance of climbers in this forest. A greater increase in photosynthetic rate and a lower increase in dark respiration rate when canopy openings occur were related to the success of climbing plant species. Dominant climbers showed a strategy of maximizing exploitation of resource availability but minimizing metabolic costs. Results may reflect phenotypic plasticity or genetic differentiation in ecophysiological traits between light environments. It is suggested that the dominant climbers in this temperate rainforest would be able to cope with forest clearings due to human activities. [ABSTRACT FROM AUTHOR]

Light requirements and functional strategies of plants to cope with light heterogeneity in the field have a strong influence on community structure and dynamics. Shade intolerant plants often show a shade avoidance strategy involving a phytochrome-mediated stem elongation in response to changes in red : far red ratio, while shade-tolerant plants typically harvest light very efficiently. We measured plant size, stem diameter, internode and leaf lengths in randomly chosen saplings of 11 woody species differing in their shade tolerance in both a secondary forest and an old-growth temperate evergreen rainforest in southern Chile. We also recorded the irradiance spectrum and the diffuse and direct light availabilities at each sampling point. Significant differences were found for the mean light environment of the saplings of each species, which also differed in basal stem diameter, internode length and leaf length, but not in plant height. Both plant slenderness (plant height/stem diameter) and mean internode length increased with increasing light availability, but no relationship was found between any of these two traits and red : far red ratio. The change in plant slenderness with light availability was of lesser magnitude with increasing shade tolerance of the species, while internode change with light availability increased with increasing shade tolerance of the species. Shade tolerators afford higher costs (thicker stems and plants), which render more biomechanically robust plants, and respond more to the light environment in a trait strongly influencing light interception (internode length) than shade intolerant species. By contrast, less shade-tolerant plants afforded higher risks with a plastic response to escape from the understorey by making thinner plants that were biomechanically weaker and poorer light interceptors. Thus, species differing in their shade tolerances do differ in their plastic responses to light. Our results contribute to explain plant coexistence in heterogeneous light environments by improving our mechanistic understanding of species responses to light. [ABSTRACT FROM AUTHOR]

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment. [ABSTRACT FROM AUTHOR]

Optimal defence theory (ODT) predicts that, whereas high risk of herbivory should select for high constitutive levels of defence, induced defences should be more advantageous in environments with a low probability of herbivory. In the present field study, conducted on the Azteca– Cecropia ant–plant system in a Neotropical rainforest, we evaluated whether the constitutive and induced ant defence of leaves are directly and inversely related to an estimate of herbivory risk, respectively. To assess the constitutive level of Azteca defence in Cecropia obtusifolia trees, we recorded the number of ants patrolling undamaged leaves. To evaluate the induced level of Azteca defence, the same leaves were subjected to simulated herbivory by punching circular holes in them. We recorded the maximum number of ants patrolling the damaged leaves from 2 to 15 min after damage. Past herbivory (% defoliation of old leaves) was assumed to indicate a risk of herbivory. Regression analyses showed that, whereas the constitutive level of ant patrolling was positively associated with the magnitude of herbivory on old leaves, there was a negative association between the magnitude of induced ant defence and past herbivory. These preliminary results lend support to ODT. [ABSTRACT FROM AUTHOR]