Direct and indirect effects of climate on richness drive the latitudinal diversity gradient in forest trees.

Climate is widely recognised as an important determinant of the latitudinal diversity gradient. However, most existing studies make no distinction between direct and indirect effects of climate, which substantially hinders our understanding of how climate constrains biodiversity globally. Using data from 35 large forest plots, we test hypothesised relationships amongst climate, topography, forest structural attributes (stem abundance, tree size variation and stand basal area) and tree species richness to better understand drivers of latitudinal tree diversity patterns. Climate influences tree richness both directly, with more species in warm, moist, aseasonal climates and indirectly, with more species at higher stem abundance. These results imply direct limitation of species diversity by climatic stress and more rapid (co-)evolution and narrower niche partitioning in warm climates. They also support the idea that increased numbers of individuals associated with high primary productivity are partitioned to support a greater number of species.

Persistent effects of fragmentation on tropical rainforest canopy structure after 20 yr of isolation.

Assessing the persistent impacts of fragmentation on aboveground structure of tropical forests is essential to understanding the consequences of land use change for carbon storage and other ecosystem functions. We investigated the influence of edge distance and fragment size on canopy structure, aboveground woody biomass (AGB), and AGB turnover in the Biological Dynamics of Forest Fragments Project (BDFFP) in central Amazon, Brazil, after 22+ yr of fragment isolation, by combining canopy variables collected with portable canopy profiling lidar and airborne laser scanning surveys with long-term forest inventories. Forest height decreased by 30% at edges of large fragments (>10 ha) and interiors of small fragments (<3 ha). In larger fragments, canopy height was reduced up to 40 m from edges. Leaf area density profiles differed near edges: the density of understory vegetation was higher and midstory vegetation lower, consistent with canopy reorganization via increased regeneration of pioneers following post-fragmentation mortality of large trees. However, canopy openness and leaf area index remained similar to control plots throughout fragments, while canopy spatial heterogeneity was generally lower at edges. AGB stocks and fluxes were positively related to canopy height and negatively related to spatial heterogeneity. Other forest structure variables typically used to assess the ecological impacts of fragmentation (basal area, density of individuals, and density of pioneer trees) were also related to lidar-derived canopy surface variables. Canopy reorganization through the replacement of edge-sensitive species by disturbance-tolerant ones may have mitigated the biomass loss effects due to fragmentation observed in the earlier years of BDFFP. Lidar technology offered novel insights and observational scales for analysis of the ecological impacts of fragmentation on forest structure and function, specifically aboveground biomass storage.

Evidence for a recent increase in forest growth.

Forests and their soils contain the majority of the earth's terrestrial carbon stocks. Changes in patterns of tree growth can have a huge impact on atmospheric cycles, biogeochemical cycles, climate change, and biodiversity. Recent studies have shown increases in biomass across many forest types. This increase has been attributed to climate change. However, without knowing the disturbance history of a forest, growth could also be caused by normal recovery from unknown disturbances. Using a unique dataset of tree biomass collected over the past 22 years from 55 temperate forest plots with known land-use histories and stand ages ranging from 5 to 250 years, we found that recent biomass accumulation greatly exceeded the expected growth caused by natural recovery. We have also collected over 100 years of local weather measurements and 17 years of on-site atmospheric CO(2) measurements that show consistent increases in line with globally observed climate-change patterns. Combined, these observations show that changes in temperature and CO(2) that have been observed worldwide can fundamentally alter the rate of critical natural processes, which is predicted by biogeochemical models. Identifying this rate change is important to research on the current state of carbon stocks and the fluxes that influence how carbon moves between storage and the atmosphere. These results signal a pressing need to better understand the changes in growth rates in forest systems, which influence current and future states of the atmosphere and biosphere.

Amazon forest carbon dynamics predicted by profiles of canopy leaf area and light environment.

Tropical forest structural variation across heterogeneous landscapes may control above-ground carbon dynamics. We tested the hypothesis that canopy structure (leaf area and light availability) - remotely estimated from LiDAR - control variation in above-ground coarse wood production (biomass growth). Using a statistical model, these factors predicted biomass growth across tree size classes in forest near Manaus, Brazil. The same statistical model, with no parameterisation change but driven by different observed canopy structure, predicted the higher productivity of a site 500 km east. Gap fraction and a metric of vegetation vertical extent and evenness also predicted biomass gains and losses for one-hectare plots. Despite significant site differences in canopy structure and carbon dynamics, the relation between biomass growth and light fell on a unifying curve. This supported our hypothesis, suggesting that knowledge of canopy structure can explain variation in biomass growth over tropical landscapes and improve understanding of ecosystem function.

Mycorrhizal type influences plant density dependence and species richness across 15 temperate forests.

Recent studies suggest that the mycorrhizal type associated with tree species is an important trait influencing ecological processes such as response to environmental conditions and conspecific negative density dependence (CNDD). However, we lack a general understanding of how tree mycorrhizal type influences CNDD strength and the resulting patterns of species abundance and richness at larger spatial scales. We assessed 305 species across 15 large, stem-mapped, temperate forest dynamics plots in Northeastern China and North America to explore the relationships between tree mycorrhizal type and CNDD, species abundance, and species richness at a regional scale. Tree species associated with arbuscular mycorrhizal (AM) fungi showed a stronger CNDD and a more positive relationship with species abundance than did tree species associated with ectomycorrhizal (ECM) fungi. For each plot, both basal area and stem abundance of AM tree species was lower than that of ECM tree species, suggesting that AM tree species were rarer than ECM tree species. Finally, ECM tree dominance showed a negative effect on plant richness across plots. These results provide evidence that tree mycorrhizal type plays an important role in influencing CNDD and species richness, highlighting this trait as an important factor in structuring plant communities in temperate forests.

Chemical novelty facilitates herbivore resistance and biological invasions in some introduced plant species.

Ecological release from herbivory due to chemical novelty is commonly predicted to facilitate biological invasions by plants, but has not been tested on a community scale. We used metabolomics based on mass spectrometry molecular networks to assess the novelty of foliar secondary chemistry of 15 invasive plant species compared to 46 native species at a site in eastern North America. Locally, invasive species were more chemically distinctive than natives. Among the 15 invasive species, the more chemically distinct were less preferred by insect herbivores and less browsed by deer. Finally, an assessment of invasion frequency in 2,505 forest plots in the Atlantic coastal plain revealed that, regionally, invasive species that were less preferred by insect herbivores, less browsed by white-tailed deer, and chemically distinct relative to the native plant community occurred more frequently in survey plots. Our results suggest that chemically mediated release from herbivores contributes to many successful invasions.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Hülsmann and Hartig suggest that ecological mechanisms other than specialized natural enemies or intraspecific competition contribute to our estimates of conspecific negative density dependence (CNDD). To address their concern, we show that our results are not the result of a methodological artifact and present a null-model analysis that demonstrates that our original findings-(i) stronger CNDD at tropical relative to temperate latitudes and (ii) a latitudinal shift in the relationship between CNDD and species abundance-persist even after controlling for other processes that might influence spatial relationships between adults and recruits.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Chisholm and Fung claim that our method of estimating conspecific negative density dependence (CNDD) in recruitment is systematically biased, and present an alternative method that shows no latitudinal pattern in CNDD. We demonstrate that their approach produces strongly biased estimates of CNDD, explaining why they do not detect a latitudinal pattern. We also address their methodological concerns using an alternative distance-weighted approach, which supports our original findings of a latitudinal gradient in CNDD and a latitudinal shift in the relationship between CNDD and species abundance.

Canopy gradients in leaf functional traits for species that differ in growth strategies and shade tolerance.

In temperate deciduous forests, vertical gradients in leaf mass per area (LMA) and area-based leaf nitrogen (Narea) are strongly controlled by gradients in light availability. While there is evidence that hydrostatic constraints on leaf development may diminish LMA and Narea responses to light, inherent differences among tree species may also influence leaf developmental and morphological response to light. We investigated vertical gradients in LMA, Narea and leaf carbon isotope composition (δ13C) for three temperate deciduous species (Carpinus caroliniana Walter, Fagus grandifolia Ehrh., Liriodendron tulipifera L.) that differed in growth strategy (e.g., indeterminate and determinate growth), shade tolerance and leaf area to sapwood ratio (Al:As). Leaves were sampled across a broad range of light conditions within three vertical layers of tree crowns to maximize variation in light availability at each height and to minimize collinearity between light and height. All species displayed similar responses to light with respect to Narea and δ13C, but not for LMA. Light was more important for gradients in LMA for the shade-tolerant (C. caroliniana) and -intolerant (L. tulipifera) species with indeterminate growth, and height (e.g., hydrostatic gradients) and light were equally important for the shade-tolerant (F. grandifolia) species with determinate growth. Fagus grandifolia had a higher morphological plasticity in response to light, which may offer a competitive advantage in occupying a broader range of light conditions throughout the canopy. Differences in responses to light and height for the taller tree species, L. tulipifera and F. grandifolia, may be attributed to differences in growth strategy or Al:As, which may alter morphological and functional responses to light availability. While height was important in F. grandifolia, height was no more robust in predicting LMA than light in any of the species, confirming the strong role of light availability in determining LMA for temperate deciduous species.

Plant diversity increases with the strength of negative density dependence at the global scale.

Theory predicts that higher biodiversity in the tropics is maintained by specialized interactions among plants and their natural enemies that result in conspecific negative density dependence (CNDD). By using more than 3000 species and nearly 2.4 million trees across 24 forest plots worldwide, we show that global patterns in tree species diversity reflect not only stronger CNDD at tropical versus temperate latitudes but also a latitudinal shift in the relationship between CNDD and species abundance. CNDD was stronger for rare species at tropical versus temperate latitudes, potentially causing the persistence of greater numbers of rare species in the tropics. Our study reveals fundamental differences in the nature of local-scale biotic interactions that contribute to the maintenance of species diversity across temperate and tropical communities.

A general model of intra-annual tree growth using dendrometer bands.

Tree growth is an important indicator of forest health, productivity, and demography. Knowing precisely how trees' grow within a year, instead of across years, can lead to a finer understanding of the mechanisms that drive these larger patterns. The growing use of dendrometer bands in research forests has only rarely been used to measure growth at resolutions finer than yearly, but intra-annual growth patterns can be observed from dendrometer bands using precision digital calipers and weekly measurements. Here we present a workflow to help forest ecologists fit growth models to intra-annual measurements using standard optimization functions provided by the R platform. We explain our protocol, test uncertainty in parameter estimates with respect to sample sizes, extend the optimization protocol to estimate robust lower and upper annual diameter bounds, and discuss potential challenges to optimal fits. We offer R code to implement this workflow. We found that starting values and initial optimization routines are critical to fitting the best functional forms. After using a bounded, broad search method, a more focused search algorithm obtained consistent results. To estimate starting and ending annual diameters, we combined the growth function with early and late estimates of beginning and ending growth. Once we fit the functions, we present extension algorithms that estimate periodic reductions in growth, total growth, and present a method of controlling for the shifting allocation to girth during the growth season. We demonstrate that with these extensions, an analysis of growth response to weather (e.g., the water available to a tree) can be derived in a way that is comparable across trees, years, and sites. Thus, this approach, when applied across broader data sets, offers a pathway to build inference about the effects of seasonal weather on growth, size- and light-dependent patterns of growth, species-specific patterns, and phenology.

Forest Canopy Stratification-Is It Useful?

It has long been recognized that the forest canopy has a complex structure that is significant for environmental interactions, regeneration, growth, and biotic habitat. Not only is the structure variously complex, but also there are many ways to conceptualize that complexity. Yet the persistent theme when considering the structure of canopies continues to be that of stratification: whether structural units are arranged in layers above the ground. We examined the use of the terms "stratification," "layering," and others in connection with canopy structure and found they had various meanings (often only implied) that were difficult to reconcile and to measure. We applied the definitions to the structure of a single, well-studied canopy located in Virginia, U.S.A., and found they failed to define consistently and clearly the presence, number, or location of strata. Additionally, we found the concept had limitations related to scale dependence, point of reference, and spatial averaging. Thus, asserting that a forest is stratified or naming the number of layers generally provided no guide to its structure. We propose alternative ways of conceptualizing and studying the forest canopy that avoid most of the problems associated with stratification. Among these are direct measurement and mapping of structural and environmental variables that have clear potential connections with canopy functions and viewing the distribution of structures or environmental conditions within the canopy as ecological gradients.

Canopy light transmittance in Douglas-fir--western hemlock stands.

We measured vertical and horizontal variation in canopy transmittance of photosynthetically active radiation in five Pseudotsuga menziesii (Mirb.) Franco-Tsuga heterophylla (Raf.) Sarg. (Douglas-fir-western hemlock) stands in the central Cascades of southern Washington to determine how stand structure and age affect the forest light environment. The shape of the mean transmittance profile was related to stand height, but height of mean maximum transmittance was progressively lower than maximum tree height in older stands. The vertical rate of attenuation declined with stand age in both the overstory and understory. A classification of vertical light zones based on the mean and variance of transmittance showed a progressive widening of the bright (low variance and high mean) and transition (high variance and rapid vertical change) zones in older stands, whereas the dim zone (low variance and mean) narrowed. The zone of maximum canopy surface area in height profiles, estimated by inversion of transmittance profiles, changed from relatively high in the canopy in most young stands ("top-heavy") to lower in the canopy in older stands ("bottom-heavy"). In the understory, all stands had similar mean transmittances, but the spatial scale of variation increased with stand age and increasing crown size. The angular distribution of openness was similar in all stands, though the older stands were less open at all angles than the younger stands. Understory openness was generally unrelated to transmittance in the canopy above. Whole-canopy leaf area indices, estimated using three methods of inverting light measurements, showed little correspondence across methods. The observed patterns in light environment are consistent with structural changes occurring during stand development, particularly the diversification of crowns, the creation of openings of various sizes and the elaboration of the outer canopy surface. The ensemble of measurements has potential use in distinguishing forests of differing ages that have similar stature.