Tree diversity reduces herbivory by forest insects

Biodiversity loss from plant communities is often acknowledged to affect primary production but little is known about effects on herbivores. We conducted a meta-analysis of a worldwide data set of 119 studies to compare herbivory in single-species and mixed forests. This showed a significant reduction of herbivory in more diverse forests but this varied with the host specificity of insects. In diverse forests, herbivory by oligophagous species was virtually always reduced, whereas the response of polyphagous species was variable. Further analyses revealed that the composition of tree mixtures may be more important than species richness per se because diversity effects on herbivory were greater when mixed forests comprised taxonomically more distant tree species, and when the proportion of non-host trees was greater than that of host trees. These findings provide new support for the role of biodiversity in ecosystem functioning across trophic levels.     

Tree species composition rather than diversity triggers associational resistance to the pine processionary moth

The reduction of insect herbivory is one of the services provided by tree diversity in forest ecosystems. While it is increasingly acknowledged that the compositional characteristics of tree species assemblages play a major role in triggering associational resistance to herbivores, underlying mechanisms are less well known. We addressed this question in the ORPHEE experiment by assessing pine processionary moth infestations (Thaumetopoea pityocampa) across a tree diversity gradient from pine monocultures to five species mixtures. We showed that tree species richness per se had no effect on the probability of attack by this pest. By contrast, the infestation rate was strongly dependent on plot composition. Mixtures of pines (Pinus pinaster) and birches (Betula pendula) were less prone to T. pityocampa infestations, whereas mixtures of pines and oaks (Quercus spp.) were more often attacked than pine monocultures. By taking into account the relative height of pines and associated broadleaved species, this effect could be explained by pine apparency. Pines were on average 343 ± 5 cm height. Birches, as fast growing trees, were slightly taller than pines (363 ± 6 cm), while oak trees were significantly smaller (74 ± 1 cm). Host trees of T. pityocampa were then partly hidden in mixtures of pines and birches but more apparent in mixtures with oaks. We suggest that reduced pine apparency disrupted visual cues used by female moths to select host trees prior to oviposition. This study highlights the need to take into account tree traits such as growth rate when selecting the tree species that have to be associated in order to improve forest resistance to pest insects.     

Vegetation and microclimatic edge effects in two mixed-mesophytic forest fragments

Forest edges are known to consist of microenvironments that may provide habitat for a different suite of species than forest interiors. Several abiotic attributes of the microenvironment may contribute to this change across the edge to center gradient (e.g., light, air temperature, soil moisture, humidity). Biotic components, such as seed dispersal, may also give rise to changes in species composition from forest edge to interior. We predicted that abiotic and biotic measures would correlate with distance from forest edge and would differ among aspects. To test these predictions, we measured abiotic and biotic variables on twelve 175 m transects in each of two 24 ha forest fragments in east-central Illinois that have remained in continuous isolation for upwards of 100 years. Both univariate and multivariate techniques were used to best describe the complex relationships among abiotic factors and between abiotic and biotic factors. Results indicate that microclimatic variables differ in the degree to and distance over which they show an edge effect. Relative humidity shows the widest edge, while light and soil moisture have the steepest gradients. Aspect influences are evidenced by the existence of more pronounced edge effects on south and west edges, except when these edges are protected by adjacent habitat. Edges bordered by agricultural fields have more extreme changes in microclimate than those bordered by trees. According to PCA results, species richness correlates well with microclimatic variation, especially light and soil moisture; however, in many cases species richness had a different depth of edge influence than either of these variables. The herbaceous plant community is heavily dominated by three species. Distributions of individual species as well as changes in plant community composition, estimated with a similarity index, indicate that competition may be influencing the response of the vegetation to the edge to interior gradient. This study indicates that edge effects must be considered when the size and potential buffering habitat of forest preserves are planned.     

The extent of edge effects in fragmented landscapes: Insights from satellite measurements of tree cover

Due to deforestation, intact tropical forest areas are increasingly transformed into a mixture of remaining forest patches and human modified areas. These forest fragments suffer from edge effects, which cause changes in ecological and ecosystem processes, undermining habitat quality and the offer of ecosystem services. Even though detailed and long term studies were developed on the topic of edge effects at local scale, understanding edge effect characteristics in fragmented forests on larger scales and finding indicators for its impact is crucial for predicting habitat loss and developing management options. Here we evaluate the spatial and temporal dimensions of edge effects in large areas using remote sensing. First we executed a neighborhood pixel analysis in 11 LANDSAT Tree Cover (LTC) scenes (180 × 185 km each, 8 in the tropics and 3 in temperate forested areas) using tree cover as an indicator of habitat quality and in relation to edge distance. Second, we executed a temporal analysis of LTC in a smaller area in the Brazilian Amazon forest where one larger forest fragment (25,890 ha) became completely fragmented in 5 years. Our results show that for all 11 scenes pixel neighborhood variation of LTC is much higher in the vicinity of forest edges, becoming lower towards the forest interior. This analysis suggests a maximum distance for edge effects and can indicate the location of unaffected core areas. However, LTC patterns in relation to fragment edge distance vary according to the analyzed region, and maximum edge distance may differ according to local conditions. Our temporal analysis illustrates the change in tree cover patterns after 5 years of fragmentation, becoming on average lower close to the edge (between 50 and 100 m). Although it is still unclear which are the main causes of LTC edge variability within and between regions, LANDSAT Tree Cover could be used as an accessible and efficient discriminator of edge and interior forest habitats in fragmented landscapes, and become invaluable for deriving qualitative spatial and temporal information of ecological and ecosystem processes.     

Edge and shape effects on ant (Hymenoptera: Formicidae) species richness and composition in forest fragments

In this paper, we tested four hypotheses relative to edge and shape effects on ant communities: (i) forest edges have lower species richness than the remnant core; (ii) species richness increases with distance from the edge; (iii) irregularly shaped remnants have lower species richness than more regular remnants; (iv) there is a higher similarity of species composition between edge and core in irregular than in regular remnants. We sampled litter ant communities on the edge and core of ten remnants, in Viçosa, Minas Gerais, Brazil. Species richness was larger at the forest core than at the edges, although did not increase with distance from the edge. Species richness did not vary with shape complexity. The similarity of species composition between edge and core showed a decreasing trend with remnant area, and did not vary with shape complexity. The observed differences of species richness between forest core and edge may be due to higher harshness of edges, caused by environmental changes. The absence of relationship between species richness and distance from the edge might indicate the range of edge effects, which would be smaller than the smallest distance of core sampled. Therefore, edges would affect litter-dwelling ant species richness in a distance smaller than 50 m. The observation of species composition allowed us to notice an effect of fragmentation that would not be noticed if we were considering only species richness. Edge may serve as step to generalist species, which may use it to colonise forest remnants. Furthermore, small remnants are more colonisation-prone by such species, and have a more homogeneous species composition than large remnants.     

Wood decomposition is more strongly controlled by temperature than by tree species and decomposer diversity in highly species rich subtropical forests

While the number of studies on the role of biodiversity on ecosystem functioning is steadily increasing, a key component of biogeochemical cycling in forests, dead wood decay, has been largely neglected. It remains widely unknown whether and how dead wood decay is affected by diversity loss in forests. We studied the hierarchical effects of tree species diversity on wood decay rates in a subtropical forest landscape in southeast China via its influence on fungal OTU richness and invertebrate diversity using piecewise structural equation models. The experiment was conducted in natural forest plots that span a wide gradient of tree species diversity embedded in a heterogeneous topography. To account for interactions between macro‐invertebrates and fungi, that potentially modify the influence of tree biodiversity and climate on dead wood decay, we compared a macro‐invertebrate exclusion treatment with a control treatment that allowed access to all types of decomposers. Diversity effects of trees on wood decay rates were mostly negative and mediated by the diversity of macro‐invertebrates. However, the effects of tree species diversity or fungal OTU richness and macro‐invertebrate diversity on wood decay rates were comparatively weak. Temperature affected decay rates positively and had the strongest influence in all treatments. While the exclusion of macro‐invertebrates did not lead to a reduction of wood decay rates, our results suggest that they may however have a mediating role in the process. In the presence of invertebrates the predictability of wood decay rates was higher and we observed a tendency of a stronger temperature control. Our results suggest that there is evidence for diversity effects on wood decomposition, but the temperature control is still more important. Thus, an increase in mean annual temperature will increase carbon and nutrient turnover through wood decomposition in subtropical forest irrespective of biotic composition.     

Drivers of understorey biomass: tree species identity is more important than richness in a young forest

林下生物量影响因素：幼龄林树种特性比丰富度更重要 生物多样性与生态系统功能的正相关关系已被广泛报道,其主要来源于对草原生态系统的研究。然而,该结论并不一定适用于更复杂的环境,例如具有不同垂直层次的森林。举例而言,已有研究表明上层乔木树种丰富度与林下生产力降低有关。树种丰富度是否会通过增加(由于生境异质性)或降低(通过增强竞争)资源的可利用性进而影响林下生产力,以及林下生产力是否受树种特性的影响更大,这些影响机制都可能会随着时间的推移而改变。此外,研究还表明,丰富度-生产力关系随着环境背景的变化而改变。本研究利用可以操控树种丰富度的实验林场研究了这些不同垂直层位里的时间和环境动态。在中国亚热带森林生物多样性与生态系统功能(BEF-China)研究计划的框架下,我们在3年时间里沿树种丰富度梯度反复采集林下生物量样本,研究了不同环境处理中树种丰富度、树种特性和时间对林下生物量的影响。尽管我们发现乔木层特性对林下生物量有显著和一致的影响,但是树种丰富度对后者却不具有这种影响。另外,在森林结构层之间,可能并不存在单一的、具有普遍性的上层乔木树种丰富度与林下生产力的相关关系,并且与上层乔木相关的环境因素(如透光率)对林下生产力的贡献程度会随着时间而变化。总体而言,我们的结果表明,在研究森林结构层之间的关系时应将时间动态变化考虑在内。     

Bottom‐up and top‐down effects of tree species diversity on leaf insect herbivory

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.     

Temporal stability of aquatic food webs: partitioning the effects of species diversity, species composition and enrichment

Theory predicts that species diversity can enhance stability of community‐level biomass while simultaneously decreasing population‐level stability. Enrichment can theoretically destabilize communities but effects may become weaker with increasing diversity because of the inclusion of consumer‐resistant prey. Few experiments using direct manipulations of species diversity have tested these predictions. We used laboratory‐based aquatic food webs to examine the effects of species composition, diversity and enrichment on temporal variability of population‐ and community‐level biomass. We found weak effects of enrichment on population‐ and community‐level stability. However, diversity enhanced community‐level stability while species composition had no influence. In contrast, composition effects outweighed diversity effects when stability was measured at the population level. We found no negative effects of diversity on population‐level stability, in opposition to theory. Our results indicate that diversity can enhance stability in multitrophic systems, but effects vary with the scale of biological organization at which stability is measured.     

Functional identity is the main driver of diversity effects in young tree communities

Two main effects are proposed to explain biodiversity–ecosystem functioning relationships: niche complementarity and selection effects. Both can be functionally defined using the functional diversity (FD) and functional identity (FI) of the community respectively. Herein, we present results from the first tree diversity experiment that separated the effect of selection from that of complementarity by varying community composition in high‐density plots along a gradient of FD, independent of species richness and testing for the effects of FD and community weighted means of traits (a proxy for FI) on stem biomass increment (a proxy for productivity). After 4 years of growth, most mixtures did not differ in productivity from the averages of their respective monocultures, but some did overyield significantly. Those positive diversity effects resulted mostly from selection effects, primarily driven by fast‐growing deciduous species and associated traits. Net diversity effect did not increase with time over 4 years.     

Climate change‐driven extinctions of tree species affect forest functioning more than random extinctions

Aim

Climate change affects forest functioning not only through direct physiological effects such as modifying photosynthesis and growing season lengths, but also through indirect effects on community composition related to species extinctions and colonizations. Such indirect effects remain poorly explored in comparison with the direct ones. Biodiversity–ecosystem functioning (BEF) studies commonly examine the effects of species loss by eliminating species randomly. However, species extinctions caused by climate change will depend on the species’ vulnerability to the new environmental conditions, thus occurring in a specific, non‐random order. Here, we evaluated whether successive tree species extinctions, according to their vulnerability to climate change, impact forest functions differently than random species losses.

Location

Eleven temperate forests across a gradient of climatic conditions in central Europe.

Methods

We simulated tree community dynamics with a forest succession model to study the impact of species loss on the communities’ aboveground biomass, productivity and temporal stability. Tree species were removed from the local pool (1) randomly, and according to (2) their inability to be recruited under a warmer climate or (3) their increased mortality under drier conditions.

Results

Results showed that non‐random species loss (i.e., based on their vulnerability to warmer or drier conditions) changed forest functioning at a different rate, and sometimes direction, than random species loss. Furthermore, directed extinctions, unlike random, triggered tipping points along the species loss process where forest functions were strongly impacted. These tipping points occurred after fewer extinctions in forests located in the coldest areas, where ecosystem functioning relies on fewer species.

Main conclusions

We showed that the extinction of species in a deterministic and mechanistically motivated order, in this case the species vulnerability to climate change, strengthens the selection effect of diversity on ecosystem functioning. BEF studies exploring the impact of species loss on ecosystem functioning using random extinctions thus possibly underestimate the potential effect of biodiversity loss when driven by a directional force, such as climate change.

     

Environment is more relevant than spatial structure as a driver of regional variation in tropical tree community richness and composition

Background: Understanding how factors related to environment and geographical distance explain community variation allows insights about how ecological niche and neutral processes control tropical community assembly.

Aims: Quantify how variation in regional tree community richness and composition in a humid tropical forest across a mountain chain are related to niche and putative neutral processes.

Methods: We used a variation partitioning routine based on Redundancy Analysis to model tropical tree community richness and composition within three distinct elevation belts, as a function of environment and spatial structure, using data from 32 studies in the Serra do Mar Range, south-eastern Brazil.

Results: Environmental effects were greater than spatial structure effects to explain community variation in the three elevation belts. There was a trend of decreasing spatial structure effects while environmental effects remained constant from lower to higher elevations. Patterns were congruent for species richness and composition.

Conclusions: We suggest that on tropical mountains, niche-related processes are equally relevant for tropical forest community assembly at all elevations, while neutral processes become weaker towards higher elevations. Determining if this trend is a consequence of the greater heterogeneity of environmental conditions associated with higher elevations in tropical mountainous terrain remains an important area of research.     

Leaf litter decomposition of sweet chestnut is affected more by oomycte infection of trees than by water temperature

Riparian forests are subjected to multiple disturbances, such as tree diseases caused by invasive pathogens, whose consequences on stream functioning are unknown. We assessed the impact of Phytophthora cinnamomi infection, and interactions with temperature, on microbial decomposition of Castanea sativa leaves. Leaves from healthy, symptomatic and highly symptomatic trees were incubated in the laboratory at 13 and 18 °C for 64 d. Infection significantly increased polyphenolic concentration and leaf toughness, reducing leaf decomposition and microbial respiration rates irrespective of temperature. Aquatic hyphomycete communities differed significantly in leaves from highly symptomatic trees. Fungal biomass was highest at 18 °C, irrespective of tree health status. None of the parameters were influenced by the tree health status × temperature interaction, suggesting that temperature rise may not synergistically increase the cross-ecosystem effects caused by P. cinnamomi in streams where litter decomposition is microbial-driven. Infection by P. cinnamomi alters the nutritional quality of leaves affecting the functioning of aquatic ecosystems.     

Down by the riverside: Riparian edge effects on three monkey species in a fragmented Costa Rican forest

Rivers represent natural edges in forests, serving as transition zones between landscapes. Natural edge effects are important to study to understand how intrinsic habitat variations affect wildlife as well as the impact of human-induced forest fragmentation. We examined the influence of riparian and anthropogenic edge on mantled howler, white-faced capuchin, Central American spider monkeys, and vegetation structure at La Suerte Biological Research Station (abbreviated as LSBRS), Costa Rica. We predicted lower monkey encounter rate, tree species richness, and median dbh at both edge types compared to interior and that monkeys would show species-specific responses to edge based on size and diet. We expected large, folivorous–frugivorous howler monkeys and small, generalist capuchins would be found at increased density in forest edge, while large, frugivorous spider monkeys would be found at decreased density in forest edge. We conducted population and vegetation surveys along interior, riparian, and anthropogenic edge transects at LSBRS and used GLMM to compare vegetation and monkey encounter rate. Tree species richness and median dbh were higher in forest interior than anthropogenic edge zones. Although spider monkey encounter rate did not vary between forest edges and interior, howler monkeys were encountered at highest density in riparian edge, while capuchins were encountered at highest density in anthropogenic edge. Our results indicate that diverse forest edges have varying effects on biota. Vegetation was negatively affected by forest edges, while monkey species showed species-specific edge preferences. Our findings suggest that riparian zones should be prioritized for conservation in Neotropical forests.     

Early subtropical forest growth is driven by community mean trait values and functional diversity rather than the abiotic environment

While functional diversity (FD) has been shown to be positively related to a number of ecosystem functions including biomass production, it may have a much less pronounced effect than that of environmental factors or species‐specific properties. Leaf and wood traits can be considered particularly relevant to tree growth, as they reflect a trade‐off between resources invested into growth and persistence. Our study focussed on the degree to which early forest growth was driven by FD, the environment (11 variables characterizing abiotic habitat conditions), and community‐weighted mean (CWM) values of species traits in the context of a large‐scale tree diversity experiment (BEF‐China). Growth rates of trees with respect to crown diameter were aggregated across 231 plots (hosting between one and 23 tree species) and related to environmental variables, FD, and CWM, the latter two of which were based on 41 plant functional traits. The effects of each of the three predictor groups were analyzed separately by mixed model optimization and jointly by variance partitioning. Numerous single traits predicted plot‐level tree growth, both in the models based on CWMs and FD, but none of the environmental variables was able to predict tree growth. In the best models, environment and FD explained only 4 and 31% of variation in crown growth rates, respectively, while CWM trait values explained 42%. In total, the best models accounted for 51% of crown growth. The marginal role of the selected environmental variables was unexpected, given the high topographic heterogeneity and large size of the experiment, as was the significant impact of FD, demonstrating that positive diversity effects already occur during the early stages in tree plantations.     

Quality or quantity: is nutrient transfer driven more by symbiont identity and productivity than by symbiont abundance?

By forming symbiotic interactions with microbes, many animals and plants gain access to the products of novel metabolic pathways. We investigated the transfer of symbiont-derived carbon and nitrogen to the sponges Aplysina cauliformis, Aplysina fulva, Chondrilla caribensis, Neopetrosia subtriangularis and Xestospongia bocatorensis, all of which host abundant microbial populations, and Niphates erecta, which hosts a sparse symbiont community. We incubated sponges in light and dark bottles containing seawater spiked with ¹³C- and ¹⁵N-enriched inorganic compounds and then measured ¹³C and ¹⁵N enrichment in the microbial (nutrient assimilation) and sponge (nutrient transfer) fractions. Surprisingly, although most sponges hosting abundant microbial communities were more enriched in ¹³C than N. erecta, only N. subtriangularis was more enriched in ¹⁵N than N. erecta. Although photosymbiont abundance varied substantially across species, ¹³C and ¹⁵N enrichment was not significantly correlated with photosymbiont abundance. Enrichment was significantly correlated with the ratio of gross productivity to respiration (P:R), which varied across host species and symbiont phylotype. Because irradiance impacts P:R ratios, we also incubated A. cauliformis in ¹³C-enriched seawater under different irradiances to determine whether symbiont carbon fixation and transfer are dependent on irradiance. Carbon fixation and transfer to the sponge host occurred in all treatments, but was greatest at higher irradiances and was significantly correlated with P:R ratios. Taken together, these results demonstrate that nutrient transfer from microbial symbionts to host sponges is influenced more by host–symbiont identities and P:R ratios than by symbiont abundance.