Miguel A. Zavala

Rate of tree carbon accumulation increases continuously with tree size

Forests are major components of the global carbon cycle, providing substantial feedback to atmospheric greenhouse gas concentrations. Our ability to understand and predict changes in the forest carbon cycle—particularly net primary productivity and carbon storage—increasingly relies on models that represent biological processes across several scales of biological organization, from tree leaves to forest stands. Yet, despite advances in our understanding of productivity at the scales of leaves and stands, no consensus exists about the nature of productivity at the scale of the individual tree, in part because we lack a broad empirical assessment of whether rates of absolute tree mass growth (and thus carbon accumulation) decrease, remain constant, or increase as trees increase in size and age. Here we present a global analysis of 403 tropical and temperate tree species, showing that for most species mass growth rate increases continuously with tree size. Thus, large, old trees do not act simply as senescent carbon reservoirs but actively fix large amounts of carbon compared to smaller trees; at the extreme, a single big tree can add the same amount of carbon to the forest within a year as is contained in an entire mid-sized tree. The apparent paradoxes of individual tree growth increasing with tree size despite declining leaf-level and stand-level productivity can be explained, respectively, by increases in a tree’s total leaf area that outpace declines in productivity per unit of leaf area and, among other factors, age-related reductions in population density. Our results resolve conflicting assumptions about the nature of tree growth, inform efforts to undertand and model forest carbon dynamics, and have additional implications for theories of resource allocation and plant senescence.

The effects of phenotypic plasticity and local adaptation on forecasts of species range shifts under climate change

Species are the unit of analysis in many global change and conservation biology studies; however, species are not uniform entities but are composed of different, sometimes locally adapted, populations differing in plasticity. We examined how intraspecific variation in thermal niches and phenotypic plasticity will affect species distributions in a warming climate. We first developed a conceptual model linking plasticity and niche breadth, providing five alternative intraspecific scenarios that are consistent with existing literature. Secondly, we used ecological niche-modeling techniques to quantify the impact of each intraspecific scenario on the distribution of a virtual species across a geographically realistic setting. Finally, we performed an analogous modeling exercise using real data on the climatic niches of different tree provenances. We show that when population differentiation is accounted for and dispersal is restricted, forecasts of species range shifts under climate change are even more pessimistic than those using the conventional assumption of homogeneously high plasticity across a species range. Suitable population-level data are not available for most species so identifying general patterns of population differentiation could fill this gap. However, the literature review revealed contrasting patterns among species, urging greater levels of integration among empirical, modeling and theoretical research on intraspecific phenotypic variation.

Jack-of-all-trades effects drive biodiversity-ecosystem multifunctionality relationships in European forests

There is considerable evidence that biodiversity promotes multiple ecosystem functions (multifunctionality), thus ensuring the delivery of ecosystem services important for human well-being. However, the mechanisms underlying this relationship are poorly understood, especially in natural ecosystems. We develop a novel approach to partition biodiversity effects on multifunctionality into three mechanisms and apply this to European forest data. We show that throughout Europe, tree diversity is positively related with multifunctionality when moderate levels of functioning are required, but negatively when very high function levels are desired. For two well-known mechanisms, ‘complementarity and ‘selection, we detect only minor effects on multifunctionality. Instead a third, so far overlooked mechanism, the ‘jack-of-all-trades effect, caused by the averaging of individual species effects on function, drives observed patterns. Simulations demonstrate that jack-of-all-trades effects occur whenever species effects on different functions are not perfectly correlated, meaning they may contribute to diversity–multifunctionality relationships in many of the worlds ecosystems.

Architecture of Iberian canopy tree species in relation to wood density, shade tolerance and climate

Tree architecture has important consequences for tree performance as it determines resource capture, mechanical stability and dominance over competitors. We analyzed architectural relationships between stem and crown dimensions for 13 dominant Iberian canopy tree species belonging to the Pinaceae (six Pinus species) and Fagaceae (six Quercus species and Fagus sylvatica) and related these architectural traits to wood density, shade tolerance and climatic factors. Fagaceae had, compared with Pinaceae, denser wood, saplings with wider crowns and adults with larger maximal crown size but smaller maximal height. In combination, these traits enhance light acquisition and persistence in shaded environments; thus, contributing to their shade tolerance. Pinaceae species, in contrast, had low-density wood, allocate more resources to the formation of the central trunk rather than to branches and attained taller maximal heights, allowing them to grow rapidly in height and compete for light following disturbances; thus, contributing to their high light requirements. Wood density had a strong relationship with tree architecture, with dense-wooded species having smaller maximum height and wider crowns, probably because of cheaper expansion costs for producing biomechanically stable branches. Species from arid environments had shorter stems and shallower crowns for a given stem diameter, probably to reduce hydraulic path length and assure water transport. Wood density is an important correlate of variation in tree architecture between species and the two dominant families, with potentially large implications for their resource foraging strategies and successional dynamics.

Stand dynamics and tree coexistence in an analytical structured model: the role of recruitment.

Understanding the mechanisms of coexistence and niche partitioning in plant communities is a central question in ecology. Current theories of forest dynamics range between the so-called neutral theories which assume functional equivalence among coexisting species to forest simulators that explain species assemblages as the result of tradeoffs in species individual strategies at several ontogenetic stages. Progress in these questions has been hindered by the inherent difficulties of developing analytical size-structured models of stand dynamics. This precludes examination of the relative importance of each mechanism on tree coexistence. In previous simulation and analytical studies emphasis has been given to interspecific differences at the sapling stage, and less so to interspecific variation in seedling recruitment. In this study we develop a partial differential equation model of stand dynamics in which competition takes place at the recruitment stage. Species differ in their size-dependent growth rates and constant mortality rates. Recruitment is described as proportional to the basal area of conspecifics, to account for fecundity and seed supply per unit of basal area, and is corrected with a decreasing function of species specific basal area to account for competition. We first analyze conditions for population persistence in monospecific stands and second we investigate conditions of coexistence for two species. In the monospecific case we found a stationary stand structure based on an inequality between mortality rate and seed supply. In turn, intra-specific competition does not play any role on the asymptotic extinction or population persistence. In the two-species case we found that coexistence can be attained when the reciprocal negative effect on recruitment follows a given relation with respect to intraspecific competition. Specifically a tradeoff between recruitment potential (i.e. shade tolerance or predation avoidance) and fecundity or growth rate. This is to our knowledge the first study that describes coexistence mechanisms in an analytical size-structured model in terms of competitive differences at the regeneration state.

Revealing patterns of local species richness along environmental gradients with a novel network tool

How species richness relates to environmental gradients at large extents is commonly investigated aggregating local site data to coarser grains. However, such relationships often change with the grain of analysis, potentially hiding the local signal. Here we show that a novel network technique, the “method of reflections”, could unveil the relationships between species richness and climate without such drawbacks. We introduced a new index related to potential species richness, which revealed large scale patterns by including at the local community level information about species distribution throughout the dataset (i.e., the network). The method effectively removed noise, identifying how far site richness was from potential. When applying it to study woody species richness patterns in Spain, we observed that annual precipitation and mean annual temperature explained large parts of the variance of the newly defined species richness, highlighting that, at the local scale, communities in drier and warmer areas were potentially the species richest. Our method went far beyond what geographical upscaling of the data could unfold, and the insights obtained strongly suggested that it is a powerful instrument to detect key factors underlying species richness patterns, and that it could have numerous applications in ecology and other fields.