Browsing by Author "Terborgh, John W"

I examined recruitment patterns of multiple tree species in a western Amazonian floodplain forest at three ontogenetic stages: seed fall, seedling establishment, and sapling recruitment.

From analyzing a long-term seed rain dataset collected using a high-density array of seed traps, I confirmed that seed fall decreases sharply with increasing distance from fruiting trees, with disproportionately large contributions from a very small fraction of all trees. Patterns of seed fall, although idiosyncratic for individual species, tended to relate to dispersal syndrome. Intact seeds were found at significantly greater distances away from fruiting adults than ripe fruit and almost exclusively comprised the tail of the seed shadow for most species.

Saplings of all species examined recruited in areas of very low predicted seed density at significantly higher abundances than expected under a null hypothesis of "all seeds are equal". The value of a seed in terms of its potential to produce a sapling recruit - measured as sapling/seed ratio - initially increased greatly with increasing distance from reproductive conspecific adults and leveled off at farther distances, in almost all species.

A parallel experimental study employed >1000 individual seedlings of common tree species situated near and far from conspecific adults. Overall survival for all species pooled and for eight out of 11 individual species was significantly higher at sites located far from versus close to conspecific adults, with the study design controlling for seedling density at sites. Survival analysis based on multiple censuses revealed that a "distance effect" persisted and intensified over time, although the timing of onset of distance-related differential mortality differed amongst species. The role of host-specific invertebrate herbivores and microbial pathogens in causing seedling mortality near conspecific adults was confirmed by the use of mesh exclosures.

Overall, my results provide community-level support for the influence of distance-dependent processes on recruitment patterns. Seed dispersal appears critical for successful recruitment and undispersed seeds make a minimal contribution. When de-coupled from distance-dependence, effects of competition-based density-dependent processes on recruitment were weak or undetectable. I conclude that community-level tree recruitment processes and patterns in western Amazonian lowland rainforests that harbor intact floral and faunal assemblages conform closely to predictions of the Janzen-Connell hypothesis of tropical tree recruitment.

Amazonia contains the largest remaining tracts of undisturbed tropical forest on earth, and is thus critical to international nature conservation and carbon sequestration efforts. Amazonian forests are notoriously difficult to study, however, due to their species richness and inaccessibility. This has limited efforts to produce the accurate, high-resolution biodiversity maps needed for conservation and development. The aims of the research described here were to identify efficient solutions to the problems of tropical forest inventory; to use these methods to identify floristic patterns and their causes in western Amazonia; and propose new means to map floristic patterns in these forests.

Using tree inventories in the vicinity of Iquitos, Peru, I and a colleague systematically evaluated methods for rapid tropical forest inventory. Of these, inventory of particular taxonomic groups, or taxonomic scope inventory, was the most efficient, and was able to capture a majority of the pattern observed by traditional inventory techniques with one-fifth to one-twentieth the number of stems and species. Based on the success of this approach, I and colleagues specifically evaluated two plant groups, the Pteridophytes (ferns and fern allies) and the Melastomataceae (a family of shrubs and small trees), for use in rapid inventory. Floristic patterns based on inventories from either group were significantly associated with those based on the tree flora, and inventories of Pteridophytes in particular were in most cases able to capture the majority of floristic patterns identified by tree inventories. These findings indicate that Pteridophyte and Melastomataceae inventories are useful tools for rapid tropical forest inventory.

Using Pteridophyte and Melastomataceae inventories from 138 sites in northwestern Amazonia, combined with satellite data and soil sampling, I and colleagues studied the causes of vegetation patterns in western Amazonian forests. On the basis of these data, we identified a floristic discontinuity of at least 300km in northern Peru, corresponding to a 15-fold difference in soil cation concentrations and an erosion-generated geological boundary. On the basis of this finding, we assembled continent-scale satellite image mosaics, and used these to search for additional discontinuities in western Amazonia. These mosaics indicate a floristic and geological discontinuity of at least 1500km western Brasil, driven by similar erosional processes identified in our study area. We suggest that this represents a chemical and ecological boundary between western and central Amazonia.

Using a second network of 52 pteridophyte and soil inventories in northwestern Amazonia, we further studied the role of geology in generating floristic pattern. Consistent with earlier findings, we found that two widespread geological formations in western Amazonia differ eight-fold difference in soil cation concentrations and in a majority of their species. Difference in elevation, used as a surrogate for geological formation, furthermore explained up to one-third of the variation in plant species composition between these formations. Significant correlations between elevation, and cation concentrations and soil texture, confirmed that differences in species composition between these formations are driven by differences in soil properties. On the basis of these findings, we were able to use SRTM elevation data to accurately model species composition throughout our study area.

I argue that Amazonian forests are partitioned into large-area units on the basis of geological formations and their edaphic properties. This finding has implications for both the ecology and evolution of these forests, and suggests that conservation strategies be implemented on a region-by-region basis. Fortunately, the methods described here provide a means for generating accurate and detailed maps of floristic patterns in these vast and remote forests.

This thesis examines the dynamics of cloud forest bird communities along an altitudinal gradient on the eastern slopes of the Ecuadorian Andes. Specifically, I examined the seasonality of breeding, investigated a novel tool for documenting altitudinal migrations, and compared diets among groups of hummingbirds. In Chapter 2, I compared the prevalence of breeding condition in mist-netted birds among elevations and months. Overall, birds breed less seasonally at lower latitudes, but there is substantial variation in the timing of breeding, which varies with both abiotic conditions like precipitation (Hau 2004; Tye 1992), photoperiod (Hau et al. 1998) and temperature (Wikelski et al. 2000), and the biotic variation in plant phenology (Komdeur 1996), and insect abundance (Poulin et al. 1992). Several of these biotic (e.g. canopy height, biological diversity) and abiotic (e.g. temperature, pressure) factors vary along elevational gradients. I compared the percentage of birds captured in breeding condition along an altitudinal transect in eastern Ecuador, and found that breeding is more seasonal at higher altitudes. There was a marked increase in breeding birds during Sept-Nov at higher elevations, but I found no such "breeding season" at lower elevations.

I also examined a novel methodology for tracing annual altitudinal migrations which takes advantage of the natural variation in deuterium abundance from the base to the peaks of the Andes (Chapter 3). Local migrations by birds in the tropics pose conservation problems, in part because the movements themselves are difficult to document. There is a theoretical relationship between Deuterium (or 18O) signature and elevation, because of fractionation during precipitation events and evapotranspiration. A previous study had suggested that if a bird had more or less deuterium in its tissues than theory would suggest, such discrepancy might be used to identify altitudinal migrants. Unfortunately, when I refined the methodology, I found that the variation within species and sites was too great to allow such applications.

In Chapter 4 I shifted my focus to comparing diet among hummingbirds. Hummingbirds rely on the sugars in nectar to meet their high metabolic requirements, but most nectars are extremely low in nitrogen. As a result, the birds must also consume arthropods to meet their protein requirements. In many hummingbird species, males use nectar resources differently from females. I hypothesized that the different genders might also differ in their intake of arthropods, because females have higher nitrogen requirements for breeding. I used stable nitrogen isotopic analysis of feathers and blood to demonstrate that females feed at higher trophic levels than males and adults at higher levels than juveniles, respectively. Feathers from female Coeligena torquata (Collared Inca) showed significantly higher 15N levels (one-tailed t20=1.73, p<.05) than males. The difference between genders in Heliodoxa leadbeateri (Violet-fronted brilliant) was smaller (one-tailed t16=1.63, p=.06). δ15N was significantly lower for juveniles (mean = 6.34, SD = 2.10) than for adults (mean = 7.53, SD = 1.24). It appears that females captured during the breeding season were also feeding at higher trophic levels than those captured outside of the breeding season, although the sample sizes were small. Finally, I also found a slight but unanticipated effect of elevation on δ15N values in feathers.

Ecologists have long sought to uncover the mechanisms behind large-scale, macroecological patterns in the distribution and abundance of species. Macroecological patterns are often attributed to the dynamics of dispersal (e.g. dispersal limitation or widespread dispersal). However, few studies actually measure dispersal to determine if dispersal rates are commensurate with the observed macroecological patterns. In this dissertation, I use population genetic analyses across many species to obtain community-level estimates of dispersal rates for two different ecological systems: birds on islands and trees in tropical rainforests. These independent estimates of dispersal then allow me to determine if macroecological patterns in these two systems can be attributed to dispersal dynamics.

In chapter two, I explore the contrasting macroecological patterns of two groups of Lesser Antillean birds. The groups' differing macroecological patterns could be due to differences in dispersal, but other authors have advocated different mechanisms. Population genetic analyses show that the two groups do differ significantly in rates of inter-island dispersal, indicating that dispersal dynamics can explain their contrasting macroecological patterns. In chapter three, I turn my attention to tropical tree communities. In contrast to studies of birds on islands, studies of trees in tropical rainforests may suffer from misidentification of individuals in the field. Using a phylogenetic approach, I determine errors rates in identification, and then assess the effect of these errors on macroecological patterns and other ecological analyses of tropical tree communities. I find that error rates are substantial, but that they have little effect on macroecological patterns. In contrast, species-level ecological analyses can be dramatically affected by these errors.

In chapter four, I return to the influence of dispersal on macroecological patterns, this time in tropical tree communities. One notable macroecological pattern in Amazonian tree communities is a high correlation in the relative abundances of species shared across communities, which could indicate high rates of dispersal between communities. However, population genetic analyses show that dispersal is severely limited between communities. Thus, some factor besides dispersal, such as differences in competitive ability or susceptibility to disease, must be driving species to achieve similar relative abundances in geographically separated communities. In contrast, I show that dispersal limitation is the likely cause of another macroecological pattern frequently observed in tropical tree communities: the decline in the compositional similarity of communities with distance. However, this is not steady-state dispersal limitation in an equilibrium framework as is conventionally thought. Instead, the dispersal limitation appears to be historical in nature, which implies a heretofore unnoticed role for historical contingency in the assembly of Amazonian tree communities.

Rapid urbanization threatens the survival of native wildlife species worldwide. In order to fully grasp the implications of the ongoing growth of urban areas on biodiversity, conservationists need to be able to quantify the response patterns of a wide range of different species to the expansion of urban and suburban land use. In this study, we set up two road-based transects across gradients of urbanization and habitat loss in the diverse longleaf pine forests of the Sandhills region of North Carolina, USA. With funding provided by the NC Wildlife Resources Commission, we drove the transects repeatedly at night in the field seasons of 2006-2008, tallying all vertebrate animals encountered (live or dead). The first transect (driven in all three years; 75 km long) ran from the urban areas of Southern Pines and Pinehurst down to the remote and relatively pristine habitats associated with the state-owned Sandhills Gamelands. The second transect (driven in 2007 and partially in 2008; 69 km long) began at the terminus of the first transect in the Gamelands, and then stretched down to the urban zones of Hamlet and Rockingham.

A total of 4900 vertebrate animals representing 69 species were observed on or near the road routes after driving a total of 16,625 km. This total includes 592 nightjars (ground-nesting nocturnal birds; e.g. whip-poor-wills) that we heard while driving the transects. In addition, in 2007 we surveyed for the nightjars and for quail (a high-priority game species that also nests on the ground) using 75 point count locations evenly distributed along the northern road route.

Regression tree analysis (a robust, nonparametric technique with minimal assumptions) was used to model the animal observation rates for a given 1 km road segment or point count as a function of various habitat variables measured within corresponding buffer zones for each segment. We also modeled snake and bird encounter rates as a function of mesopredator mammal observations.

Our results reveal that amphibian, snake, and ground-nesting bird observation rates are negatively associated with increasing levels of traffic and impervious surface. Conversely, mesopredator mammals (and domestic cats in particular) responded slightly positively to increasing urbanization, and negatively to protected area coverage. Both ground-nesting birds and snakes showed signs of negative correlations with mesopredator encounter rates, although these trends were not always significant due to high variability in the mesopredator data.

In order to try and confirm the results of the regression tree analyses, we also used a multivariate ordination approach (non-metric multidimensional scaling) to visualize the integrated community structure of all of the major vertebrate groups we observed in the Sandhills. The ordinations revealed that while the snake, ground-nesting bird, and amphibian groups were similar to each other in terms of their avoidance of urban conditions, the cats and native mesopredator species actually seemed to define widely divergent axes of community variation. Cats in particular were separated from the other groups on 2 out of 3 axes of the species-space ordination. Still, as we noted above for the regression tree models, it is difficult to sort out with our correlative data set whether cats and other mesopredators truly played an independent role in structuring and/or depleting the other wildlife guilds along our route. More experimental approaches are recommended for trying to resolve whether overabundant predators or road mortality and inappropriate habitat are more to blame for the much reduced encounter rates we observed for the snakes, birds, and amphibians in urban areas. Future studies will also be needed to confirm the logical assumption that road encounter rates provide a reasonably accurate index of the relative abundance of the different animal groups along the survey routes.