Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects.We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives. Geosphere-Biosphere Program (IGBP) and DIVERSITAS, the TRY database (TRY-not an acronym, rather a statement of sentiment; https ://www.try-db.org; Kattge et al., 2011) was proposed with the explicit assignment to improve the availability and accessibility of plant trait data for ecology and earth system sciences. The Max Planck Institute for Biogeochemistry (MPI-BGC) offered to host the database and the different groups joined forces for this community-driven program. Two factors were key to the success of TRY: the support and trust of leaders in the field of functional plant ecology submitting large databases and the long-term funding by the Max Planck Society, the MPI-BGC and the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, which has enabled the continuous development of the TRY database.
Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks.
Many examples of biased sex ratios are known in natural populations of plants. Proximal causes of these biases are gender diphasy (sex changing), differential mortality between male and female genets, differential rates of clonal growth (numbers of ramets per genet), and differential flowering (differences in flowering frequency or age to maturity). In the western North American shrub Oemleria cerasiformis we determined sex ratios for 60 natural populations and found an excess of males in 56 populations. The male bias was greatest in populations with little recent recruitment. Sampling of young and old plants indicated that males flowered at an earlier age than females, which led to a transient flowering bias in very young plants, and that the genet sex ratio was 1:1 in young mature plants but male biased in old plants, as a result of differential mortality. Examination of dead genets confirmed that mature females have higher mortality rates. Females also have greater reproductive effort and slower growth rates than males. The major cause of biased sex ratios in O. cerasiformis is greater mortality of female plants during their reproductive years, which probably arises directly or indirectly from their greater allocation to reproduction.
We examined the size, age, and spatial structure of trees in an old Engelmann spruce (Picea engelmannii)-subalpine fir (Abies lasiocarpa) forest based on four stem-mapped, 0.25 ha plots. Dendrochronological techniques were used on basal discs of 1,190 trees to reconstruct age and growth pattern, including dates of rapid growth increases. There were no obvious age cohorts or other evidence of past major disturbance. The abundance of both subalpine fir and spruce decreased rapidly with age, especially beyond the ages of 150 years. Very old trees were present, but rare. The best evidence from tree-ring width patterns for past disturbance was a period of release 100 years ago. However, few of the released trees grew into the canopy, which suggests a disturbance of low intensity. Patch dynamics and gap processes were not pronounced in the stand. Clumping was generally weak and only present at small spatial scales (<5 m) for live trees, and largely non-existent for dead trees; mortality was spatially random in this forest. Although spruce were sparse (5.1% of trees) in the forest relative to fir, which is consistent with predictions that fir will ultimately replace spruce in the absence of disturbance, coexistence seems more likely judging from the age structure and numbers of dead trees. In contrast to almost all spruce-fir forests studied previously, the stand we examined showed no record of major disturbances. Thus this stand falls at the limit of the range of dynamics - from disturbance-structured to near steady-state - encompassed in current thinking about forest ecosystems.
We used dendrochronological analysis of over 2000 trees in four 50 × 50 m plots to reconstruct the history and dynamics of a 330-year-old, fire-initiated spruce-fir forest. All lodgepole pine (Pinus contorta Dougl. ex Loud.), half of the canopy Engelmann spruce (Picea engelmannii Parry ex Engelm.), but less than 10% of the canopy subalpine fir (Abies lasiocarpa (Hook.) Nutt.) dated from the first 50 years of stand development. Tree-ring patterns of individual surviving trees showed no evidence of disturbance during the first 200 years after stand initiation; subsequently, episodes of disturbance are indicated by periods of release in understory fir. Although many fir owe their canopy position to release after disturbance, few canopy fir in the current stand established in response to either the stand-initiating event or subsequent partial disturbances. A seedling bank of long-lived fir appears critical to the dynamics of this forest. In contrast, establishment of almost all canopy spruce can be related to disturbance. This stand, although fire initiated, was structured primarily by a combination of partial disturbances and autogenic processes. We suspect that most old, fire-initiated stands in many forest regions are similarly structured and emphasize that the contribution of partial disturbances and autogenic processes should be fully assessed when examining their dynamics or managing such forests.
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