Wood performs several essential functions in plants, including mechanically supporting aboveground tissue, storing water and other resources, and transporting sap. Woody tissues are likely to face physiological, structural and defensive trade-offs. How a plant optimizes among these competing functions can have major ecological implications, which have been under-appreciated by ecologists compared to the focus they have given to leaf function. To draw together our current understanding of wood function, we identify and collate data on the major wood functional traits, including the largest wood density database to date (8412 taxa), mechanical strength measures and anatomical features, as well as clade-specific features such as secondary chemistry. We then show how wood traits are related to one another, highlighting functional trade-offs, and to ecological and demographic plant features (growth form, growth rate, latitude, ecological setting). We suggest that, similar to the manifold that tree species leaf traits cluster around the 'leaf economics spectrum', a similar 'wood economics spectrum' may be defined. We then discuss the biogeography, evolution and biogeochemistry of the spectrum, and conclude by pointing out the major gaps in our current knowledge of wood functional traits.

Towards a worldwide wood economics spectrum

The increasing availability of phylogenetic data, computing power and informatics tools has facilitated a rapid expansion of studies that apply phylogenetic data and methods to community ecology. Several key areas are reviewed in which phylogenetic information helps to resolve long-standing controversies in community ecology, challenges previous assumptions, and opens new areas of investigation. In particular, studies in phylogenetic community ecology have helped to reveal the multitude of processes driving community assembly and have demonstrated the importance of evolution in the assembly process. Phylogenetic approaches have also increased understanding of the consequences of community interactions for speciation, adaptation and extinction. Finally, phylogenetic community structure and composition holds promise for predicting ecosystem processes and impacts of global change. Major challenges to advancing these areas remain. In particular, determining the extent to which ecologically relevant traits are phylogenetically conserved or convergent, and over what temporal scale, is critical to understanding the causes of community phylogenetic structure and its evolutionary and ecosystem consequences. Harnessing phylogenetic information to understand and forecast changes in diversity and dynamics of communities is a critical step in managing and restoring the Earths biota in a time of rapid global change.

https://projects.au.dk/fileadmin/projects/circe/2009_Cavender_et_al_01.pdf

The merging of community ecology and phylogenetic biology

INTRODUCTION. Internal Organization of the Plant Body. Summary of Types of Cells and Tissues. General References. DEVELOPMENT OF THE SEED PLANT. The Embryo. From embryo to the Adult Plant. Apical Meristems and Their Derivatives. Differentiation, Specialization, and Morphogenesis. References. THE CELL. Cytoplasm. Nucleus. Plastids. Mitochondria. Microbodies. Vacuoles. Paramural Bodies. Ribosomes. Dictyosomes. Endoplasmic Reticulum. Lipid Globules. Microtubules. Ergastic Substances. References. CELL WALL. Macromolecular Components and Their Organization in the Wall. Cell Wall Layers. Intercellular Spaces. Pits, Primary Pit--Fields, and Plasmodesmata. Origin of Cell Wall During Cell Division. Growth of Cell Wall. References. PARENCHYMA AND COLLENCHYMA. Parenchyma. Collenchyma. References. SCLERENCHYMA. Sclereids. Fibers. Development of Sclereids and Fibers. References. EPIDERMIS. Composition. Developmental Aspects. Cell Wall. Stomata. Trichomes. References. XYLEM: GENERAL STRUCTURE AND CELL TYPES. Gross Structure of Secondary Xylem. Cell Types in the Secondary Xylem. Primary Xylem. Differentiation of Tracheary Elements. References. XYLEM: VARIATION IN WOOD STRUCTURE. Conifer Wood. Dicotyledon Wood. Some Factors in Development of Secondary Xylem. Identification of Wood. References. VASCULAR CAMBIUM. Organization of Cambium. Developmental Changes in the Initial Layer. Patterns and Causal Relations in Cambial Activity. References. PHLOEM. Cell Types. Primary Phloem. Secondary Phloem. References. PERIDERM. Structure of Periderm and Related Tissues. Development of Periderm. Outer Aspect of Bark in Relation to Structure. Lenticels. References. SECRETORY STRUCTURES. External Secretory Structures. Internal Secretory Structures. References. THE ROOT: PRIMARY STATE OF GROWTH. Types of Roots. Primary Structure. Development. References. THE ROOT: SECONDARY STATE OF GROWTH AND ADVENTITIOUS ROOTS. Common Types of Secondary Growth. Variations in Secondary Growths. Physiologic Aspects of Secondary Growth in Roots. Adventitious Roots. References. THE STEM: PRIMARY STATE OF GROWTH. External Morphology. Primary Structure. Development. References. THE STEM: SECONDARY GROWTH AND STRUCTURAL TYPES. Secondary Growth. Types of Stems. References. THE LEAF: BASIC STRUCTURE AND DEVELOPMENT. Morphology. Histology of Angiosperm Leaf. Development. Abscission. References. THE LEAF: VARIATIONS IN STRUCTURE. Leaf Structure and Environment. Dicotyledon Leaves. Monocotyledon Leaves. Gymnosperm Leaves. References. THE FLOWER: STRUCTURE AND DEVELOPMENT. Concept. Structure. Development. References. THE FLOWER: REPRODUCTIVE CYCLE. Microsporogenesis. Pollen. Male Gametophyte. Megasporogenesis. Female Gametophyte. Fertilization. References. THE FRUIT. Concept and Classification. The Fruit Wall. Fruit Types. Fruit Growths. Fruit Abscission. References. THE SEED. Concept and Morphology. Seed Development. Seed Coat. Nutrient Storage Tissues. References. EMBRYO AND SEEDLING. Mature Embryo. Development of Embryo. Classification of Embryos. Seedling. References. Glossary. Index.

Anatomy of Seed Plants

It is debated whether species-level differences in ecological strategy, which play a key role in much of coexistence theory, are important in structuring highly diverse communities. We examined the co-occurrence patterns of over 1100 tree species in a 25-hectare Amazonian forest plot in relation to field-measured functional traits. Using a null model approach, we show that co-occurring trees are often less ecologically similar than a niche-free (neutral) model predicts. Furthermore, we find evidence for processes that simultaneously drive convergence and divergence in key aspects of plant strategy, suggesting that at least two distinct niche-based processes are occurring. Our results show that strategy differentiation among species contributes to the maintenance of diversity in one of the most diverse tropical forests in the world.

https://science.sciencemag.org/content/sci/322/5901/580.full.pdf

Functional traits and niche-based tree community assembly in an Amazonian forest

Land-use change occurs nowhere more rapidly than in the tropics, where the imbalance between deforestation and forest regrowth has large consequences for the global carbon cycle. However, considerable uncertainty remains about the rate of biomass recovery in secondary forests, and how these rates are influenced by climate, landscape, and prior land use. Here we analyse aboveground biomass recovery during secondary succession in 45 forest sites and about 1,500 forest plots covering the major environmental gradients in the Neotropics. The studied secondary forests are highly productive and resilient. Aboveground biomass recovery after 20 years was on average 122 megagrams per hectare (Mg ha(-1)), corresponding to a net carbon uptake of 3.05 Mg C ha(-1) yr(-1), 11 times the uptake rate of old-growth forests. Aboveground biomass stocks took a median time of 66 years to recover to 90% of old-growth values. Aboveground biomass recovery after 20 years varied 11.3-fold (from 20 to 225 Mg ha(-1)) across sites, and this recovery increased with water availability (higher local rainfall and lower climatic water deficit). We present a biomass recovery map of Latin America, which illustrates geographical and climatic variation in carbon sequestration potential during forest regrowth. The map will support policies to minimize forest loss in areas where biomass resilience is naturally low (such as seasonally dry forest regions) and promote forest regeneration and restoration in humid tropical lowland areas with high biomass resilience.

Biomass resilience of Neotropical secondary forests

Wood density is a crucial variable in carbon accounting programs of both secondary and old-growth tropical forests. It also is the best single descriptor of wood: it correlates with numerous morphological, mechanical, physiological, and ecological properties. To explore the extent to which wood density could be estimated for rare or poorly censused taxa, and possible sources of variation in this trait, we analyzed regional, taxonomic, and phylogenetic variation in wood density among 2456 tree species from Central and South America. Wood density varied over more than one order of magnitude across species, with an overall mean of 0.645 g/cm3. Our geographical analysis showed significant decreases in wood density with increasing altitude and significant differences among low-altitude geographical regions: wet forests of Central America and western Amazonia have significantly lower mean wood density than dry forests of Central and South America, eastern and central Amazonian forests, and the Atlantic forests of Brazil; and eastern Amazonian forests have lower wood densities than the dry forests and the Atlantic forest. A nested analysis of variance showed that 74% of the species-level wood density variation was explained at the genus level, 34% at the Angiosperm Phylogeny Group (APG) family level, and 19% at the APG order level. This indicates that genus-level means give reliable approximations of values of species, except in a few hypervariable genera. We also studied which evolutionary shifts in wood density occurred in the phylogeny of seed plants using a composite phylogenetic tree. Major changes were observed at deep nodes (Eurosid 1), and also in more recent divergences (for instance in the Rhamnoids, Simaroubaceae, and Anacardiaceae). Our unprecedented wood density data set yields consistent guidelines for estimating wood densities when species-level information is lacking and should significantly reduce error in Central and South American carbon accounting programs.

Regional and phylogenetic variation of wood density across 2456 Neotropical tree species.

El aliso (Alnus acuminata, Betulaceae) es un arbol pionero que domina bosques riparios de montana en los Andes orientales. Estudios iniciales atribuyeron la presencia de estos bosques a las tipicas condiciones microclimaticas de cauces de rios, pero luego estos fueron vinculados al regimen periodico de disturbios de rios de montana. Para comprender la dinamica de estos bosques, analizamos la estructura de edades y el arreglo espacial de arboles en dos parcelas sobre el rio Potrero y otra parcela sobre el rio La Horqueta, en la provincia de Tucuman, noroeste argentino. Mediante barrenado intensivo y reconstruccion de edades identificamos cohortes de arboles con un rango maximo de edades de ocho anos. El mapeo de arboles y el indice de autocorrelacion espacial I de Moran indicaron que las cohortes de arboles estaban agrupadas en espacio, proporcionando evidencia adicional de que los parches coetaneos de A. acuminata resultan de eventos de regeneracion masiva luego de crecientes catastroficas. Evaluamos la asociacion entre anos de inicio de cohortes y registros regionales de descarga maxima de rios por medio de aleatorizaciones y encontramos evidencia debil de que los establecimientos ocurren luego de crecientes devastadoras que eliminan la vegetacion preexistente y originan sitios desnudos propicios para la regeneracion del aliso. Las fechas de cicatrices de crecientes en arboles del rio Potrero preceden por un ano al inicio de la cohorte mas reciente y dan respaldo adicional a esta conexion. A pesar de que estudios previos encontraron una correlacion positiva entre eventos de El Nino y promedios de precipitacion en el noroeste argentino, nuestros datos no revelan una asociacion significativa entre El Nino y caudales maximos anuales en rios de la provincia de Tucuman. Se requieren mas estudios para identificar los factores que originan crecientes devastadoras y cohortes de A. acuminata

/pdf/don-t-camp-beside-the-river-structure-and-dynamics-of-andean-4z0ihjcj95.pdf

Tomás A. Easdale

Papers

Regional and phylogenetic variation of wood density across 2456 Neotropical tree species.

Don't camp beside the river: structure and dynamics of Andean alder (Alnus acuminata) forests affected by river floods, northwestern Argentina