Paul V. A. Fine

The merging of community ecology and phylogenetic biology

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.

Phylogenetic beta diversity: linking ecological and evolutionary processes across space in time

A key challenge in ecological research is to integrate data from different scales to evaluate the ecological and evolutionary mechanisms that influence current patterns of biological diversity. We build on recent attempts to incorporate phylogenetic information into traditional diversity analyses and on existing research on beta diversity and phylogenetic community ecology. Phylogenetic beta diversity (phylobetadiversity) measures the phylogenetic distance among communities and as such allows us to connect local processes, such as biotic interactions and environmental filtering, with more regional processes including trait evolution and speciation. When combined with traditional measures of beta diversity, environmental gradient analyses or ecological niche modelling, phylobetadiversity can provide significant and novel insights into the mechanisms underlying current patterns of biological diversity.

THE GROWTH-DEFENSE TRADE-OFF AND HABITAT SPECIALIZATION BY PLANTS IN AMAZONIAN FORESTS

Tropical forests include a diversity of habitats, which has led to specialization in plants. Near Iquitos, in the Peruvian Amazon, nutrient-rich clay forests surround nutrient-poor white-sand forests, each harboring a unique composition of habitat specialist trees. We tested the hypothesis that the combination of impoverished soils and herbivory creates strong natural selection for plant defenses in white-sand forest, while rapid growth is favored in clay forests. Recently, we reported evidence from a reciprocal-transplant experiment that manipulated the presence of herbivores and involved 20 species from six genera, including phylogenetically independent pairs of closely related white-sand and clay specialists. When protected from herbivores, clay specialists exhibited faster growth rates than white-sand specialists in both habitats. But, when unprotected, white-sand specialists outperformed clay specialists in white-sand habitat, and clay specialists outperformed white-sand specialists in clay habitat. Here we test further the hypothesis that the growth defense trade-off contributes to habitat specialization by comparing patterns of growth, herbivory, and defensive traits in these same six genera of white-sand and clay specialists. While the probability of herbivore attack did not differ between the two habitats, an artificial defoliation experiment showed that the impact of herbivory on plant mortality was significantly greater in white-sand forests. We quantified the amount of terpenes, phenolics, leaf toughness, and available foliar protein for the plants in the experiment. Different genera invested in different defensive strategies, and we found strong evidence for phylogenetic constraint in defense type. Overall, however, we found significantly higher total defense investment for white-sand specialists, relative to their clay specialist congeners. Furthermore, herbivore resistance consistently exhibited a significant trade-off against growth rate in each of the six phylogenetically independent species-pairs. These results confirm theoretical predictions that a trade-off exists between growth rate and defense investment, causing white-sand and clay specialists to evolve divergent strategies. We propose that the growth-defense trade-off is universal and provides an important mechanism by which herbivores govern plant distribution patterns across resource gradients.

Global patterns of leaf mechanical properties

Leaf mechanical properties strongly influence leaf lifespan, plant-herbivore interactions, litter decomposition and nutrient cycling, but global patterns in their interspecific variation and underlying mechanisms remain poorly understood. We synthesize data across the three major measurement methods, permitting the first global analyses of leaf mechanics and associated traits, for 2819 species from 90 sites worldwide. Key measures of leaf mechanical resistance varied c. 500-800-fold among species. Contrary to a long-standing hypothesis, tropical leaves were not mechanically more resistant than temperate leaves. Leaf mechanical resistance was modestly related to rainfall and local light environment. By partitioning leaf mechanical resistance into three different components we discovered that toughness per density contributed a surprisingly large fraction to variation in mechanical resistance, larger than the fractions contributed by lamina thickness and tissue density. Higher toughness per density was associated with long leaf lifespan especially in forest understory. Seldom appreciated in the past, toughness per density is a key factor in leaf mechanical resistance, which itself influences plant-animal interactions and ecosystem functions across the globe.

The invasibility of tropical forests by exotic plants

Current research efforts to understand the relative invasibility of different plant communities have mostly ignored tropical forests. Only a few studies have treated invasive species in tropical forests, and recent worldwide analyses have not provided clear predictions concerning the relative invasibility of tropical forests. In this review, the extent to which exotic species have invaded tropical forests is summarized and four leading hypotheses to explain the apparently low frequency of invading plants in tropical forests are evaluated. In general, it is found that invasibility positively correlates with human disturbance, and that undisturbed tropical forests harbour few exotic species. To date, there is no evidence to attribute the low invasibility of undisturbed tropical forests to either their high species diversity or their high diversity of functional types. Instead, the low occurrence of exotic species in most tropical forests is most likely due to the fact that the great majority of exotic species that are transported to tropical countries lack specific life history traits, most importantly shade tolerance, necessary for successful invasion of undisturbed tropical forests. Unfortunately, this situation could change in the future with the expected increase in the plantation forestry of high-grade timber combined with common forestry practices that favour the cultivation of exotic species.

Evidence for a Time‐Integrated Species‐Area Effect on the Latitudinal Gradient in Tree Diversity

The greater area of tropical forest biomes has been proposed as a factor that drives the latitudinal gradient in species diversity by modulating speciation and extinction rates. But speciation and extinction are processes that operate over millions of years, so an adequate test of area’s contribution to diversity patterns must take into consideration that biome areas have changed through time in response to climate. Here we correlate estimates of current tree species diversity with a composite parameter integrating area over geological time for each continent’s tropical, temperate, and boreal biomes. We find significant positive correlations between current tree diversity and area‐time for periods since the Eocene, Oligocene, and Miocene, which we take as evidence for a time‐integrated species‐area effect on current patterns of species richness across biomes. These results contribute to explanations for why most lineages have tropical origins and why tropical forests are more diverse than extratropical forests.

THE CONTRIBUTION OF EDAPHIC HETEROGENEITY TO THE EVOLUTION AND DIVERSITY OF BURSERACEAE TREES IN THE WESTERN AMAZON

Abstract —Environmental heterogeneity in the tropics is thought to lead to specialization in plants and thereby contribute to the diversity of the tropical flora. We examine this idea with data on the habitat specificity of 35 western Amazonian species from the genera Protium, Crepidospermum, and Tetragastris in the monophyletic tribe Protieae (Burseraceae) mapped on a molecular‐based phylogeny. We surveyed three edaphic habitats that occur throughout terra firme Amazonia: white‐sand, clay, and terrace soils in eight forests across more than 2000 km in the western Amazon. Twenty‐six of the 35 species were found to be associated with only one of three soil types, and no species was associated with all three habitats; this pattern of edaphic specialization was consistent across the entire region. Habitat association mapped onto the phylogenetic tree shows association with terrace soils to be the probable ancestral state in the group, with subsequent speciation events onto clay and white‐sand soils. The repeated gain of clay association within the clade likely coincides with the emergence of large areas of clay soils in the Miocene deposited during the Andean uplift. Character optimizations revealed that soil association was not phylogenetically clustered for white‐sand and clay specialists, suggesting repeated independent evolution of soil specificity is common within the Protieae. This phylogenetic analysis also showed that multiple cases of putative sister taxa with parapatric distributions differ in their edaphic associations, suggesting that edaphic heterogeneity was an important driver of speciation in the Protieae in the Amazon basin.

Global Gradients in Vertebrate Diversity Predicted by Historical Area-Productivity Dynamics and Contemporary Environment

A novel hierarchical framework integrates the effects of time, area, productivity, and temperature at their respective relevant scales and successfully predicts the latitudinal gradient in global vertebrate diversity.

A floristic study of the white-sand forests of Peru.

Abstract Tropical forests occurring on white-sand soils have a unique structure and are famous for their endemism. Yet, no comprehensive floristic study has ever been undertaken in white-sand forests in the western Amazon. Here, we present the results of floristic inventories from 16 plots in seven sites from the Peruvian Amazon to investigate diversity, species composition, and endemism in white-sand forests. We compare our results to a large data set from terra firme forests from more fertile soils in the same region. We found that white-sand forest plots have extremely low average species diversity (41.5 species per 0.1-ha plot) and that white-sand plots have significantly different species composition from terra firme plots. We classify 114 species as endemic to white sand, with another 21 species that can be considered facultative specialists or cryptic endemics. These endemics and specialists are extremely dominant, accounting for more than 83% of the total number of stems surveyed in white-sand forest plots. We place our results in the context of the role of environmental heterogeneity influencing patterns of species diversity and the conservation of Amazonian forests.

Environmental factors predict community functional composition in Amazonian forests

Summary 1. The consequences of biodiversity loss for ecosystem services largely depend on the functional identities of extirpated species. However, poor descriptions of spatial patterns of community functional composition across landscapes hamper accurate predictions, particularly in highly diverse tropical regions. Therefore, understanding how community functional composition varies across environmental gradients remains an important challenge. 2. We sampled 15 functional traits in 800 Neotropical tree species across 13 forest plots representative of the broad climatic and soil gradients encompassed by three widespread lowland forest habitats (terra firme forests on clay-rich soils, seasonally flooded forests and white-sand forests) at opposite ends of Amazonia (Peru and French Guiana). We combined univariate and multivariate approaches to test the magnitude and predictability of environmental filtering on community leaf and wood functional composition. 3. Directional shifts in community functional composition correlated with environmental changes across the 13 plots, with denser leaves, stems and roots in forests occurring in environments with limited water and soil-nutrient availability. Critically, these relationships allowed us to accurately predict the functional composition of 61 additional forest plots from environmental data alone. 4. Synthesis. Environmental filtering consistently shapes the functional composition of highly diverse tropical forests at large scales across the terra firme, seasonally flooded and white-sand forests of lowland Amazonia. Environmental factors drive and allow the prediction of variation in community functional composition among habitat types in Amazonian forests.