Kenneth J. Feeley

Assessing Evidence for a Pervasive Alteration in Tropical Tree Communities

In Amazonian tropical forests, recent studies have reported increases in aboveground biomass and in primary productivity, as well as shifts in plant species composition favouring fast-growing species over slow-growing ones. This pervasive alteration of mature tropical forests was attributed to global environmental change, such as an increase in atmospheric CO2 concentration, nutrient deposition, temperature, drought frequency, and/or irradiance. We used standardized, repeated measurements of over 2 million trees in ten large (16–52 ha each) forest plots on three continents to evaluate the generality of these findings across tropical forests. Aboveground biomass increased at seven of our ten plots, significantly so at four plots, and showed a large decrease at a single plot. Carbon accumulation pooled across sites was significant (+0.24 MgC ha−1 y−1, 95% confidence intervals [0.07, 0.39] MgC ha−1 y−1), but lower than reported previously for Amazonia. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. Over all ten plots, the fastest-growing quartile of species gained biomass (+0.33 [0.09, 0.55] % y−1) compared with the tree community as a whole (+0.15 % y−1); however, this significant trend was due to a single plot. Biomass of slow-growing species increased significantly when calculated over all plots (+0.21 [0.02, 0.37] % y−1), and in half of our plots when calculated individually. Our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Instead, they suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability. More long-term studies are necessary to clarify the contribution of global change to the functioning of tropical forests.

PREDICTING BIODIVERSITY CHANGE: OUTSIDE THE CLIMATE ENVELOPE, BEYOND THE SPECIES–AREA CURVE

Efforts to anticipate threats to biodiversity take the form of species richness predictions (SRPs) based on simple correlations with current climate and habitat area. We review the major approaches that have been used for SRP, species-area curves and climate envelopes, and suggest that alternative research efforts may provide more understanding and guidance for management. Extinction prediction suffers from a number of limitations related to data and the novelty of future environments. We suggest additional attention to (1) identification of variables related to biodiversity that are diagnostic and potentially more predictable than extinction, (2) constraints on species dispersal and reproduction that will determine population persistence and range shifts, including limited sources or potential immigrants for many regions, and (3) changes in biotic interactions and phenology. We suggest combinations of observational and experimental approaches within a framework available for ingesting heterogeneous data sources. Together, these recommendations amount to a shift in emphasis from prediction of extinction numbers to identification of vulnerabilities and leading indicators of change, as well as suggestions for surveillance tools needed to evaluate important variables and the experiments likely to provide most insight.

Analysis of avian communities in Lake Guri, Venezuela, using multiple assembly rule models

This study analyzed the distribution of resident, forest-interior bird species nesting on islands in Lake Guri, Venezuela using several different community assembly rule models. The models that were tested included Diamonds Assembly Rules, Size Structure, Guild Proportionality, Favored States, and Nestedness. It was determined that the species composition of the study communities was only weakly influenced by competition, but that competition did appear to limit the size similarity which is permissible for co-occurring species. There was no tendency for the relative proportion of species within guilds (i.e. insectivore, omnivore, nectivore and frugivore) to remain stable among the islands. When only the insectivorous and omnivorous species were analyzed (using feeding strata as the functional groups) there was some support for the guild proportionality hypothesis. This study found no support for Foxs Favored State hypothesis, possibly due to the overrepresentation of insectivores and omnivores in the species pool. The island communities exhibit a highly nested structure. This high degree of nestedness supports the hypothesis that the assemblages are more strongly determined by differential extinction vulnerability and selective species loss than by interspecific or inter-guild competition. Understanding patterns of community assembly and their underlying forces has important implications for conservation ecology and reserve design.

Biodiversity Conservation: The Key Is Reducing Meat Consumption

The consumption of animal-sourced food products by humans is one of the most powerful negative forces affecting the conservation of terrestrial ecosystems and biological diversity. Livestock production is the single largest driver of habitat loss, and both livestock and feedstock production are increasing in developing tropical countries where the majority of biological diversity resides. Bushmeat consumption in Africa and southeastern Asia, as well as the high growth-rate of per capita livestock consumption in China are of special concern. The projected land base required by 2050 to support livestock production in several megadiverse countries exceeds 30-50% of their current agricultural areas. Livestock production is also a leading cause of climate change, soil loss, water and nutrient pollution, and decreases of apex predators and wild herbivores, compounding pressures on ecosystems and biodiversity. It is possible to greatly reduce the impacts of animal product consumption by humans on natural ecosystems and biodiversity while meeting nutritional needs of people, including the projected 2-3 billion people to be added to human population. We suggest that impacts can be remediated through several solutions: (1) reducing demand for animal-based food products and increasing proportions of plant-based foods in diets, the latter ideally to a global average of 90% of food consumed; (2) replacing ecologically-inefficient ruminants (e.g. cattle, goats, sheep) and bushmeat with monogastrics (e.g. poultry, pigs), integrated aquaculture, and other more-efficient protein sources; and (3) reintegrating livestock production away from single-product, intensive, fossil-fuel based systems into diverse, coupled systems designed more closely around the structure and functions of ecosystems that conserve energy and nutrients. Such efforts would also impart positive impacts on human health through reduction of diseases of nutritional extravagance.

THE EFFECTS OF HERBIVORE DENSITY ON SOIL NUTRIENTS AND TREE GROWTH IN TROPICAL FOREST FRAGMENTS

The role of herbivores in nutrient cycling in tropical forest ecosystems re- mains poorly understood. This study investigates several aspects of nutrient cycling along a gradient in herbivore (Alouatta seniculus, red howler monkey) density among small landbridge islands in Lago Guri, Venezuela. Specifically, two contrasting hypotheses were addressed: (1) herbivores increase the availability of soil nutrients and subsequently primary productivity, and (2) herbivores decrease nutrient availability and primary productivity because they increase the dominance of non-preferred, nutrient-poor, tree species. Although C:N increased with herbivore density, the annual increase in basal area (an indicator of aboveground productivity) increased with herbivore density. According to an analysis of the tree communities on the study islands, herbivory may also be causing a shift in the tree community toward non-preferred species, and thus, over a longer time scale, both nutrient availability and productivity are expected to decline. The influence of herbivores on nutrient cycling and plant productivity has important implications for conservation and rates of carbon sequestration in tropical forests.

The role of gap phase processes in the biomass dynamics of tropical forests.

The responses of tropical forests to global anthropogenic disturbances remain poorly understood. Above-ground woody biomass in some tropical forest plots has increased over the past several decades, potentially reflecting a widespread response to increased resource availability, for example, due to elevated atmospheric CO2 and/or nutrient deposition. However, previous studies of biomass dynamics have not accounted for natural patterns of disturbance and gap phase regeneration, making it difficult to quantify the importance of environmental changes. Using spatially explicit census data from large (50 ha) inventory plots, we investigated the influence of gap phase processes on the biomass dynamics of four ‘old-growth’ tropical forests (Barro Colorado Island (BCI), Panama; Pasoh and Lambir, Malaysia; and Huai Kha Khaeng (HKK), Thailand). We show that biomass increases were gradual and concentrated in earlier-phase forest patches, while biomass losses were generally of greater magnitude but concentrated in rarer later-phase patches. We then estimate the rate of biomass change at each site independent of gap phase dynamics using reduced major axis regressions and ANCOVA tests. Above-ground woody biomass increased significantly at Pasoh (+0.72% yr−1) and decreased at HKK (−0.56% yr−1) independent of changes in gap phase but remained stable at both BCI and Lambir. We conclude that gap phase processes play an important role in the biomass dynamics of tropical forests, and that quantifying the role of gap phase processes will help improve our understanding of the factors driving changes in forest biomass as well as their place in the global carbon budget.

Thermophilization of adult and juvenile tree communities in the northern tropical Andes

Significance Understanding how species respond to climate change is crucial to the development of effective conservation strategies. We found directional and systematic changes in the composition of both adult and juvenile tree species in tropical north Andean forests at rates consistent with concurrent temperature increases, supporting the hypothesis of upward species migrations resulting from global warming. Our results indicate that compositional shifts occur primarily via range retractions. This suggests elevated risk for species extinctions and local biodiversity loss with ongoing warming. Other environmental filters, such as species-soil adaptations, as well as anthropogenic disturbances (e.g., habitat loss and hunting) may limit future species migrations and decrease the ability of forest communities and their constituent species to respond to climate change. Climate change is expected to cause shifts in the composition of tropical montane forests towards increased relative abundances of species whose ranges were previously centered at lower, hotter elevations. To investigate this process of “thermophilization,” we analyzed patterns of compositional change over the last decade using recensus data from a network of 16 adult and juvenile tree plots in the tropical forests of northern Andes Mountains and adjacent lowlands in northwestern Colombia. Analyses show evidence that tree species composition is strongly linked to temperature and that composition is changing directionally through time, potentially in response to climate change and increasing temperatures. Mean rates of thermophilization [thermal migration rate (TMR), °C⋅y−1] across all censuses were 0.011 °C⋅y−1 (95% confidence interval = 0.002–0.022 °C⋅y−1) for adult trees and 0.027 °C⋅y−1 (95% confidence interval = 0.009–0.050 °C⋅y−1) for juvenile trees. The fact that thermophilization is occurring in both the adult and juvenile trees and at rates consistent with concurrent warming supports the hypothesis that the observed compositional changes are part of a long-term process, such as global warming, and are not a response to any single episodic event. The observed changes in composition were driven primarily by patterns of tree mortality, indicating that the changes in composition are mostly via range retractions, rather than range shifts or expansions. These results all indicate that tropical forests are being strongly affected by climate change and suggest that many species will be at elevated risk for extinction as warming continues.

Patterns of bird functional diversity on land-bridge island fragments.

The loss of species diversity due to habitat fragmentation has been extensively studied. In contrast, the impacts of habitat fragmentation on functional diversity remains relatively poorly understood. We conducted bird functional diversity studies on a set of 41 recently isolated land-bridge islands in the Thousand Island Lake, China. We analysed differences in bird species richness and a recently developed suite of complementary functional diversity indices (FRic, volume of functional space occupied; FEve, evenness of abundance distribution in the functional trait space; FDiv, divergence in the distribution of abundance in the trait volume) across different gradients (island area and isolation). We found no correlations between FRic and FEve or FEve and FDiv, but negative correlations between FRic and FDiv. As predicted, island area accounted for most of the variation in bird species richness, whereas isolation explained most of the variation in species evenness (decreasing species evenness with increasing isolation). Functional diversity appears to be more strongly influenced by habitat filtering as opposed to limiting similarity. More specifically, across all islands, both FRic and FEve were significantly lower than expected for randomly assembled communities, but FDiv showed no clear patterns. FRic increased with island area, FEve decreased with island area and FDiv showed no clear patterns. Our finding that FEve decreases with island area at TIL may indicate low functional stability on such islands, and as such large islands and habitat patches may deserve extra attention and/or protection. These results help to demonstrate the importance of considering the effects of fragmentation on functional diversity in habitat management and reserve design plans.

Determinants of plant species richness and patterns of nestedness in fragmented landscapes: Evidence from land-bridge islands

Land-bridge islands formed by dam construction are considered to be “experimental” systems for studying the effects of habitat loss and fragmentation, offering many distinct advantages over terrestrial fragments. The Thousand Island Lake in Southeast China is one such land-bridge system with more than 1000 islands. Based on a field survey of vascular plant richness on 154 land-bridge islands during 2007–2008, we examined the effects of island and landscape attributes on plant species richness and patterns of species nestedness. We also examined the different responses of plant functional groups (classified according to growth form and shade tolerance) to fragmentation. We found that island area explained the greatest amount of variation in plant species richness. Island area and shape index positively affected species diversity and the degree of nestedness exhibited by plant communities while the perimeter to area ratio of the islands had a negative effect. Shade-tolerant plants were the most sensitive species group to habitat fragmentation. Isolation negatively affected the degree of nestedness in herb and shade-intolerant plants including species with various dispersal abilities in the fragmented landscape. Based on these results, we concluded that the effects of habitat loss and fragmentation on overall species richness depended mostly on the degree of habitat loss, but patterns of nestedness were generated from different ecological mechanisms due to species-specific responses to different characteristics of habitat patches.

Ancient human disturbances may be skewing our understanding of Amazonian forests

Significance The Amazon harbors thousands of species and plays a vital role in the Earth’s climate and carbon cycles. Much of what we know about the Amazon is based on censuses of only a small number of forest inventory plots, an even smaller number of which are censused repeatedly and used to study forest dynamics and carbon fluxes. The effects of ancient human impacts have never been properly assessed or accounted for in studies of Amazonian plots. New spatial analyses show that plots significantly oversample areas with high abundances of archaeological evidence of past human activities. This suggests that our interpretations of the Amazon’s structure, composition, and function are based disproportionately on forests still reflecting the legacies of past human disturbances. Although the Amazon rainforest houses much of Earth’s biodiversity and plays a major role in the global carbon budget, estimates of tree biodiversity originate from fewer than 1,000 forest inventory plots, and estimates of carbon dynamics are derived from fewer than 200 recensus plots. It is well documented that the pre-European inhabitants of Amazonia actively transformed and modified the forest in many regions before their population collapse around 1491 AD; however, the impacts of these ancient disturbances remain entirely unaccounted for in the many highly influential studies using Amazonian forest plots. Here we examine whether Amazonian forest inventory plot locations are spatially biased toward areas with high probability of ancient human impacts. Our analyses reveal that forest inventory plots, and especially forest recensus plots, in all regions of Amazonia are located disproportionately near archaeological evidence and in areas likely to have ancient human impacts. Furthermore, regions of the Amazon that are relatively oversampled with inventory plots also contain the highest values of predicted ancient human impacts. Given the long lifespan of Amazonian trees, many forest inventory and recensus sites may still be recovering from past disturbances, potentially skewing our interpretations of forest dynamics and our understanding of how these forests are responding to global change. Empirical data on the human history of forest inventory sites are crucial for determining how past disturbances affect modern patterns of forest composition and carbon flux in Amazonian forests.