Landscape-level impact of tropical forest loss and fragmentation on bird occurrence in eastern Guatemala

Tropical forest destruction and fragmentation of habitat patches may reduce population persistence at the landscape level. Given the complex nature of simultaneously evaluating the effects of these factors on biotic populations, statistical presence/absence modelling has become an important tool in conservation biology. This study uses logistic regression to evaluate the independent effects of tropical forest cover and fragmentation on bird occurrence in eastern Guatemala. Logistic regression models were constructed for 10 species with varying response to habitat alteration. Predictive variables quantified forest cover, fragmentation and their interaction at three different radii (200, 500 and 1000 m scales) of 112 points where presence of target species was determined. Most species elicited a response to the 1000 m scale, which was greater than most species’ reported territory size. Thus, their presence at the landscape scale is probably regulated by extra-territorial phenomena, such as dispersal. Although proportion of forest cover was the most important predictor of species’ presence, there was strong evidence of area-independent and -dependent fragmentation effects on species presence, results that contrast with other studies from northernmost latitudes. Species’ habitat breadth was positively correlated with AIC model values, indicating a better fit for species more restricted to tropical forest. Species with a narrower habitat breadth also elicited stronger negative responses to forest loss. Habitat breadth is thus a simple measure that can be directly related to species’ vulnerability to landscape modification. Model predictive accuracy was acceptable for 4 of 10 species, which were in turn those with narrower habitat breadths.     

Landscape history, fragmentation, and patch occupancy: models for a forest bird with limited dispersal.

We developed and tested patch occupancy models for an endemic understory bird with limited dispersal ability, the Chucao Tapaculo (Scelorchilus rubecula), in two South American temperate rain forest landscapes that differed in levels and duration of forest loss. We assessed cover changes since 1961 in each landscape and surveyed patches for Chucao Tapaculo occupancy. We then developed incidence-based predictive models independently for each landscape and tested each model reciprocally in the alternative study area. We thereby assessed the domain of model applicability and identified those predictor variables with general effects and those that varied between the two landscapes. The two models were consistent regarding variable selection, and predictive accuracy of each model was high in the landscape where training data were collected. However, the models differed substantially in the magnitudes of effects related to patch size, with larger unoccupied patches observed in the landscape with the more advanced stage of fragmentation. Due to this discrepancy, each model performed poorly when applied to the alternative landscape, potentially reflecting the contrasting stages of habitat loss. Although it was impossible to dissociate effects of level and duration of forest loss, we viewed the landscapes as representing two extremes along a continuum of fragmentation, providing insights into potential trajectories for portions of the biome where deforestation is occurring. Further, our data suggest that static equilibrium models developed from distribution patterns in recently fragmented landscapes may overestimate persistence when used as a forecasting tool, or when extrapolated to alternative landscapes where fragmentation is more advanced. Thus, we suggest that landscapes used as standards for model building should be selected with caution. We recommend that distribution patterns be obtained from landscapes where fragmentation is advanced, preferably with histories of fragmentation long enough that time-delayed extinctions already would have occurred.     

Pre-reproductive survival in a tropical bird and its implications for avian life histories

The factors that affect survival until reproduction are essential to understanding the organization of life histories within and among species. Theory predicts, for example, that survival until reproduction influences the optimum level of reproductive investment by parents, which might partly explain prolonged parental care in species with high first-year survival. Tests and refinements of life-history theory have been hampered, however, by a lack of field-based estimates of pre-reproductive survival, especially for tropical species, which have been the subject of many comparative analyses. Tropical species are predicted to have higher first-year survival and delayed reproduction compared to Northern Hemisphere species. We estimated survival until reproduction, age at first reproduction, and sources of variation in juvenile survival in a Neotropical passerine, the Western Slaty-Antshrike (Thamnophilus atrinucha), in central Panama. We observed that fledged antshrikes had 76% survival through the dependent period and 48% survival to the age of 1 year; survival rate was lowest during the first week after leaving the nest. Timing of fledging within the breeding season, fledgling mass, and age at dispersal influenced survival, while sex of offspring and year did not. Individuals did not breed until two years of age, and post-fledging pre-reproductive survival was 41% of annual adult survival. High survival until reproduction in antshrikes balanced their low annual productivity, resulting in a stable population. Survival during the post-fledging period of dependence and the first year of independence in the Western Slaty-Antshrike exceeded estimates for Northern Hemisphere species. This difference appears to be associated with the extended post-fledging parental care, delayed dispersal, low costs of dispersal, and the less seasonal environment of antshrikes.     

Habitat fragmentation and effects of herbivore (howler monkey) abundances on bird species richness

Habitat fragmentation can alter herbivore abundances, potentially causing changes in the plant community that can propagate through the food web and eventually influence other important taxonomic groups such as birds. Here we test the relationship between the density of red howler monkeys (Alouatta seniculus) and bird species richness on a large set of recently isolated land-bridge islands in Lago Guri, Venezuela (n = 29 islands). Several of these islands host relict populations of howler monkeys at densities up to more than 30 times greater than those on the mainland. These "hyperabundant" herbivores previously have been shown to have a strong positive influence on aboveground plant productivity. We predicted that this should lead to a positive, indirect effect of howler monkey density on bird species richness. After accounting for passive sampling (the tendency for species richness to be positively associated with island area, regardless of differences in habitat quality) we found a significant positive correlation between howler monkey density and bird species richness. A path analysis incorporating data on tree growth rates from a subset of islands (n = 9) supported the hypothesis that the effect of howler monkeys on the resident bird communities is indirect and is mediated through changes in plant productivity and habitat quality. These results highlight the potential for disparate taxonomic groups to be related through indirect interactions and trophic cascades.     

Satellite detection of bird communities in tropical countryside.

The future of biodiversity hinges partly on realizing the potentially high conservation value of human-dominated countryside. The characteristics of the countryside that promote biodiversity preservation remain poorly understood, however, particularly at the fine scales at which individual farmers tend to make land use decisions. To address this problem, we explored the use of a rapid remote sensing method for estimating bird community composition in tropical countryside, using a two-step process. First, we asked how fine-grained variation in land cover affected community composition. Second, we determined whether the observed changes in community composition correlated with three easily accessible remote sensing metrics (wetness, greenness, and brightness), derived from performing a tasseled-cap transformation on a Landsat Enhanced Thematic Mapper Plus image. As a comparison, we also examined whether the most commonly used remote sensing indicator in ecology, the Normalized Difference Vegetation Index (NDVI), correlated with community composition. We worked within an agricultural landscape in southern Costa Rica, where the land comprised a complex and highly heterogeneous mosaic of remnant native vegetation, pasture, coffee cultivation, and other crops. In this region, we selected 12 study sites (each < 60 ha) that encompassed the range of available land cover possibilities in the countryside. Within each site, we surveyed bird communities within all major land cover types, and we conducted detailed field mapping of land cover. We found that the number of forest-affiliated species increased with forest cover and decreased with residential area across sites. Conversely, the number of agriculture-affiliated species using forest increased with land area devoted to agricultural and residential uses. Interestingly, we found that the wetness and brightness metrics predicted the number of forest- and agriculture-affiliated species within a site as well as did detailed field-generated maps of land cover. In contrast, NDVI and the closely correlated greenness metric did not correlate with land cover or with bird communities. Our study shows the strong potential of the tasseled-cap transformation as a tool for assessing the conservation value of countryside for biodiversity.     

Poverty and corruption compromise tropical forest reserves.

We used the global fire detection record provided by the satellite-based Moderate Resolution Imaging Spectroradiometer (MODIS) to determine the number of fires detected inside 823 tropical and subtropical moist forest reserves and for contiguous buffer areas 5, 10, and 15 km wide. The ratio of fire detection densities (detections per square kilometer) inside reserves to their contiguous buffer areas provided an index of reserve effectiveness. Fire detection density was significantly lower inside reserves than in paired, contiguous buffer areas but varied by five orders of magnitude among reserves. The buffer: reserve detection ratio varied by up to four orders of magnitude among reserves within a single country, and median values varied by three orders of magnitude among countries. Reserves tended to be least effective at reducing fire frequency in many poorer countries and in countries beset by corruption. Countries with the most successful reserves include Costa Rica, Jamaica, Malaysia, and Taiwan and the Indonesian island of Java. Countries with the most problematic reserves include Cambodia, Guatemala, Paraguay, and Sierra Leone and the Indonesian portion of Borneo. We provide fire detection density for 3964 tropical and subtropical reserves and their buffer areas in the hope that these data will expedite further analyses that might lead to improved management of tropical reserves.     

Functional nonredundancy of elephants in a disturbed tropical forest

Conservation efforts are often motivated by the threat of global extinction. Yet if conservationists had more information suggesting that extirpation of individual species could lead to undesirable ecological effects, they might more frequently attempt to protect or restore such species across their ranges even if they were not globally endangered. Scientists have seldom measured or quantitatively predicted the functional consequences of species loss, even for large, extinction‐prone species that theory suggests should be functionally unique. We measured the contribution of Asian elephants (Elephas maximus) to the dispersal of 3 large‐fruited species in a disturbed tropical moist forest and predicted the extent to which alternative dispersers could compensate for elephants in their absence. We created an empirical probability model with data on frugivory and seed dispersal from Buxa Tiger Reserve, India. These data were used to estimate the proportion of seeds consumed by elephants and other frugivores that survive handling and density‐dependent processes (Janzen‐Connell effects and conspecific intradung competition) and germinate. Without compensation, the number of seeds dispersed and surviving density‐dependent effects decreased 26% (Artocarpus chaplasha), 42% (Careya arborea), and 72% (Dillenia indica) when elephants were absent from the ecosystem. Compensatory fruit removal by other animals substantially ameliorated these losses. For instance, reductions in successful dispersal of D. indica were as low as 23% when gaur (Bos gaurus) persisted, but median dispersal distance still declined from 30% (C. arborea) to 90% (A. chaplasha) without elephants. Our results support the theory that the largest animal species in an ecosystem have nonredundant ecological functionality and that their extirpation is likely to lead to the deterioration of ecosystem processes such as seed dispersal. This effect is likely accentuated by the overall defaunation of many tropical systems.     

Habitat fragmentation affects movement behavior of migrating juvenile common toads

Displacement characteristics in animals are crucial drivers of successful movement decisions in resources acquisition, migration, and dispersal. As landscape structure is modified by human activity, mobility patterns are likely to evolve in response to habitat fragmentation. In species with complex life cycles that involve obligatory migrations between different habitats, one can predict that movement propensity would be promoted by fragmentation as long as it allows to reaching a habitat patch. Here, we compare the movement characteristics of naive toadlets sampled in populations distributed over a fragmentation gradient to test the hypothesis of a positive correlation between fragmentation and mobility levels. We studied toadlet movement in experimental arenas providing small patches of suitable conditions. We recorded the use of these patches (patch behavior) or the absence of their use (overtaking behavior). The more fragmented the original landscape, the more prone the toadlets were to combine these two behaviors, thus showing a higher motivation to explore. Moreover, the more fragmented the original landscape, the less the toadlets exhibited patch behavior. As the toadlets were reared in a common environment, the behavioral differences detected, relating to the level of fragmentation, resulted from inheritance. Our results thus illustrate that fragmentation is likely to create cross-generational transmittable variations in movement characteristics.     

Directional changes in the species composition of a tropical forest

Long-term studies have revealed that the structure and dynamics of many tropical forests are changing, but the causes and consequences of these changes remain debated. To learn more about the forces driving changes within tropical forests, we investigated shifts in tree species composition over the past 25 years within the 50-ha Forest Dynamics Plot on Barro Colorado Island (BCI), Panama, and examined how observed patterns relate to predictions of (1) random population fluctuations, (2) carbon fertilization, (3) succession from past disturbance, (4) recovery from an extreme El Ni?o drought at the start of the study period, and (5) long-term climate change. We found that there have been consistent and directional changes in the tree species composition. These shifts have led to increased relative representations of drought-tolerant species as determined by the species' occurrence both across a gradient of soil moisture within BCI and across a wider precipitation gradient from a dry forest near the Pacific coast of Panama to a wet forest near its Caribbean coast. These nonrandom changes cannot be explained by stochastic fluctuations or carbon fertilization. They may be the legacy of the El Ni?o drought, or alternatively, potentially reflect increased aridity due to long-term climate change. By investigating compositional changes, we increased not only our understanding of the ecology of tropical forests and their responses to large-scale disturbances, but also our ability to predict how future global change will impact some of the critical services provided by these important ecosystems.     

Independent effects of fragmentation on forest songbirds: an organism-based approach.

The degree to which spatial patterns influence the dynamics and distribution of populations is a central question in ecology. This question is even more pressing in the context of rapid habitat loss and fragmentation, which threaten global biodiversity. However, the relative influence of habitat loss and landscape fragmentation, the spatial patterning of remaining habitat, remains unclear. If landscape pattern affects population size, managers may be able to design landscapes that mitigate habitat loss. We present the results of a mensurative experiment designed to test four habitat loss vs. fragmentation hypotheses. Unlike previous studies, we measured landscape structure using quantitative, spatially explicit habitat distribution models previously developed for two species: Blackburnian Warbler (Dendroica fusca) and Ovenbird (Seiurus aurocapilla). We used a stratified sampling design that reduced the confounding of habitat amount and fragmentation variables. Occurrence and reoccurrence of both species were strongly influenced by characteristics at scales greater than the individual territory, indicating little support for the random-sample hypothesis. However, the type and spatial extent of landscape influence differed. Both occurrence and reoccurrence of Blackburnian Warblers were influenced by the amount of poor-quality matrix at 300- and 2000-m spatial extents. The occurrence and reoccurrence of Ovenbirds depended on a landscape pattern variable, patch size, but only in cases when patches were isolated. These results support the hypothesis that landscape pattern is important for some species only when the amount of suitable habitat is low. Although theoretical models have predicted such an interaction between landscape fragmentation and composition, to our knowledge this is the first study to report empirical evidence of such nonlinear fragmentation effects. Defining landscapes quantitatively from an organism-based perspective may increase power to detect fragmentation effects, particularly in forest mosaics where boundaries between patches and matrix are ambiguous. Our results indicate that manipulating landscape pattern may reduce negative impacts of habitat loss for Ovenbird, but not Blackburnian Warbler. We emphasize that most variance in the occurrence of both species was explained by local scale or landscape composition variables rather than variables reflecting landscape pattern.     

Effects of forest fragmentation on avian breeding activity

Biodiversity declines and ecosystem decay follow forest fragmentation; initially, abundant species may become rare or be extirpated. Underlying mechanisms behind delayed extirpation of certain species following forest fragmentation are unknown. Species declines may be attributed to an inadequate number of breeding adults required to replace the population or decreased juvenile survival rate due to reduced recruitment or increased nest predation pressures. We used 10 years of avian banding data, 5 years before and 4 years after fragment isolation, from the Biological Dynamics of Forest Fragments Project, carried out near Manaus, Brazil, to investigate the breeding activity hypothesis that there is less breeding activity and fewer young after relative to before fragment isolation. We compared the capture rates of active breeding and young birds in 3 forest types (primary forest, fragment before isolation, and fragment after isolation) and the proportion of active breeding and young birds with all birds in each unique fragment type before and after isolation. We grouped all bird species by diet (insectivore or frugivore) and nesting strategy (open cup, cavity, or enclosed) to allow further comparisons among forest types. We found support for the breeding activity hypothesis in insectivorous and frugivorous birds (effect sizes 0.45 and 0.53, respectively) and in birds with open-cup and enclosed nesting strategies (effect sizes 0.56 and 0.44, respectively) such that on average there were more breeding birds in fragments before isolation relative to after isolation. A larger proportion of birds in the community were actively breeding before fragment isolation (72%) than after fragment isolation (11%). Unexpectedly, there was no significant decrease in the number of young birds after fragment isolation, although sample sizes for young were small (n = 43). This may have been due to sustained immigration of young birds to fragments after isolation. Together, our results provide some of the strongest evidence to date that avian breeding activity decreases in response to fragment isolation, which could be a fundamental mechanism contributing to ecosystem decay.     

Leaf herbivory and decomposability in a Malaysian tropical rain forest

There is accumulating evidence that similar suites of plant traits may affect leaf palatability and leaf litter decomposability. However, the possible association between leaf herbivory and litter decomposition rates across species in species-diverse natural ecosystems such as tropical rain forests remains unexplored, despite its importance in estimating the herbivory effects on carbon and nutrient cycling of ecosystems. We found no strong association between leaf herbivory and litter decomposition rates across 40 tree species in a Malaysian tropical rain forest, even though the leaf and litter traits were tightly correlated. This is because the leaf and litter traits related to herbivory and decomposition rates in the field were inconsistent. Leaf toughness accounted for only a small part of the variation in the herbivory rate, whereas a number of litter traits (the leaf mass per area, lignin to nitrogen ratio, and condensed tannin concentration) accurately predicted the decomposition rate across species. These results suggest that herbivory rate across species may not be strongly related to single leaf traits, probably because plant-herbivore interactions in tropical rain forests are highly diverse; on the other hand, plant-decomposer interactions are less specific and can be governed by litter chemicals. We also investigated two factors, phylogeny and tree functional types, that could affect the relationship between herbivory and decomposition across species. Phylogenetic relatedness among the species did not affect the relationship between herbivory and decomposition. In contrast, when the plants were segregated according to their leaf emergence pattern, we found a significant positive relationship between herbivory and decomposition rates for continuous-leafing species. In these species, the condensed tannin to N ratios in leaves and litter were related to herbivory and decomposition rates, respectively. However, we did not observe a similar trend for synchronous-leafing species. These results suggest that the relationship between herbivory and decomposition may be more greatly affected by functional types than by phylogenetic relatedness among species. In conclusion, our results suggest that well-defended leaves are not necessarily less decomposable litter in a tropical rain forest community, implying that herbivory may not generate positive feedback for carbon and nutrient cycling in this type of ecosystem.     

Nonredundancy in the dispersal network of a generalist tropical forest tree

Plant species with generalized dispersal mutualisms are considered to be robust to local frugivore extinctions because of redundancy between dispersal agents. However, real redundancy can only occur if frugivores have similar foraging and ranging patterns and if fruit is a limiting resource. We evaluated the quantitative and qualitative contributions of seed dispersers for an endochorus mast-fruiting species, Prunus javanica (Rosaceae) in Khao Yai National Park, Thailand, to evaluate the potential redundancy of dispersers. Data were collected from tree watches, seed/fruit traps, and seed transects under and away from fruiting trees, feeding and seed deposition by gibbons (Hylobates lar), and evaluations of seed and first-year seedling survival. We identified three clusters of dispersers within the network. Most (>80%) frugivore species observed were small birds and squirrels that were not functional dispersers, dropping most seeds under or very near the tree crown, where seedling survival was ultimately nil. Monkeys (Macaca leonina) were low-quality, short-range dispersers, but they dispersed large numbers of seeds and were responsible for 67% of surviving first-year seedlings. Gibbons and Oriental Pied Hornbills (Anthracoceros albirostris) handled few fruits, but they provided the highest quality service by carrying most seeds away from the canopy to medium and long distances, respectively. Although there was overlap in the deposition patterns of the functional dispersers, they displayed complementary, rather than redundant, roles in seed dispersal. Satiation of all functional dispersers further limited their capacity to "replace" one another. Redundancy must be evaluated at the community level because each type of disperser may shift to different species in the non-masting years of P. javanica. Our results underscore the need for research on broader spatial and temporal scales, which combines studies of dispersal and plant recruitment, to better understand mechanisms that maintain network stability.     

Nutrient regulation of organic matter decomposition in a tropical rain forest

Terrestrial biosphere-atmosphere CO2 exchange is dominated by tropical forests, so understanding how nutrient availability affects carbon (C) decomposition in these ecosystems is central to predicting the global C cycle's response to environmental change. In tropical rain forests, phosphorus (P) limitation of primary production and decomposition is believed to be widespread, but direct evidence is rare. We assessed the effects of nitrogen (N) and P fertilization on litter-layer organic matter decomposition in two neighboring tropical rain forests in southwest Costa Rica that are similar in most ways, but that differ in soil P availability. The sites contain 100-200 tree species per hectare and between species foliar nutrient content is variable. To control for this heterogeneity, we decomposed leaves collected from a widespread neotropical species, Brosimum utile. Mass loss during decomposition was rapid in both forests, with B. utile leaves losing >80% of their initial mass in <300 days. High organic matter solubility throughout decomposition combined with high rainfall support a model of litter-layer decomposition in these rain forests in which rapid mass loss in the litter layer is dominated by leaching of dissolved organic matter (DOM) rather than direct CO2 mineralization. While P fertilization did not significantly affect mass loss in the litter layer, it did stimulate P immobilization in decomposing material, leading to increased P content and a lower C:P ratio in soluble DOM. In turn, increased P content of leached DOM stimulated significant increases in microbial mineralization of DOM in P-fertilized soil. These results show that, while nutrients may not affect mass loss during decomposition in nutrient-poor, wet ecosystems, they may ultimately regulate CO2 losses (and hence C storage) by limiting microbial mineralization of DOM leached from the litter layer to soil.     

Group size and ectoparasitism affect daily survival probability in a colonial bird

Little is known in general about how group size or ectoparasitism affect survival in colonial animals. We estimated daily within-season survival probabilities for nesting adult and recently fledged juvenile cliff swallows (Petrochelidon pyrrhonota) at 239 colonies from 1983 to 2003 in southwestern Nebraska, USA. Some colonies had been fumigated to remove ectoparasites. We conducted mark-recapture at each colony site to estimate daily survival. There were no systematic differences between males and females in daily survival. Adults and juveniles occupying parasite-free colonies had, on average, 4.4% and 62.2% greater daily survival, respectively, than their counterparts in naturally infested colonies. Daily survival of all birds increased with colony size for both parasite-free colonies and those under natural conditions, although the effect was stronger for adults at fumigated sites and for juveniles. Average daily survival probability for adults tended to increase during warmer and drier summers. Although daily survival varied at some sites over the course of the nesting cycle, there were no strongly consistent within-year temporal effects on survival. Even small differences in daily survival probability can translate into large effects on mean lifespan. The deleterious effects of ectoparasites on daily survival within the season represent a previously unknown cost of ectoparasitism. The increase in within-season survival with colony size reflects the net effects of many costs and benefits associated with colony size. Ectoparasitism is probably the most important cost that tends to partly balance the positive effects of large colonies. The greater survival of cliff swallows in the larger colonies is a previously unknown advantage of colonial nesting.Communicated by P. Heeb     

Rain forest fragmentation and the proliferation of successional trees

The effects of habitat fragmentation on diverse tropical tree communities are poorly understood. Over a 20-year period we monitored the density of 52 tree species in nine predominantly successional genera (Annona, Bellucia, Cecropia, Croton, Goupia, Jacaranda, Miconia, Pourouma, Vismia) in fragmented and continuous Amazonian forests. We also evaluated the relative importance of soil, topographic, forest dynamic, and landscape variables in explaining the abundance and species composition of successional trees. Data were collected within 66 permanent 1-ha plots within a large (approximately 1000 km2) experimental landscape, with forest fragments ranging from 1 to 100 ha in area. Prior to forest fragmentation, successional trees were uncommon, typically comprising 2-3% of all trees (> or =10 cm diameter at breast height [1.3 m above the ground surface]) in each plot. Following fragmentation, the density and basal area of successional trees increased rapidly. By 13-17 years after fragmentation, successional trees had tripled in abundance in fragment and edge plots and constituted more than a quarter of all trees in some plots. Fragment age had strong, positive effects on the density and basal area of successional trees, with no indication of a plateau in these variables, suggesting that successional species could become even more abundant in fragments over time. Nonetheless, the 52 species differed greatly in their responses to fragmentation and forest edges. Some disturbance-favoring pioneers (e.g., Cecropia sciadophylla, Vismia guianensis, V. amazonica, V. bemerguii, Miconia cf. crassinervia) increased by >1000% in density on edge plots, whereas over a third (19 of 52) of all species remained constant or declined in numbers. Species responses to fragmentation were effectively predicted by their median growth rate in nearby intact forest, suggesting that faster-growing species have a strong advantage in forest fragments. An ordination analysis revealed three main gradients in successional-species composition across our study area. Species gradients were most strongly influenced by the standlevel rate of tree mortality on each plot and by the number of nearby forest edges. Species-composition also varied significantly among different cattle ranches, which differed in their surrounding matrices and disturbance histories. These same variables were also the best predictors of total successional-tree abundance and species richness. Successional-tree assemblages in fragment interior plots (>150 m from edge), which are subjected to fragment area effects but not edge effects, did not differ significantly from those in intact forest, indicating that area effects per se had little influence on successional trees. Soils and topography also had little discernable effect on these species. Collectively, our results indicate that successional-tree species proliferate rapidly in fragmented Amazonian forests, largely as a result of chronically elevated tree mortality near forest edges and possibly an increased seed rain from successional plants growing in nearby degraded habitats. The proliferation of fast-growing successional trees and correlated decline of old-growth trees will have important effects on species composition, forest dynamics, carbon storage, and nutrient cycling in fragmented forests.     

Common ectomycorrhizal networks may maintain monodominance in a tropical rain forest

Most tropical rain forests contain diverse arrays of tree species that form arbuscular mycorrhizae. In contrast, the less common monodominant rain forests, in which one tree species comprises more than 50% of the canopy, frequently contain ectomycorrhizal (ECM) associates. In this study, I explored the potential for common ECM networks, created by aggregations of ECM trees, to enhance seedling survivorship near parent trees. I determined the benefit conferred by the common ECM network on seedling growth and survivorship of an ECM monodominant species in Guyana. Seedlings with access to an ECM network had greater growth (73% greater), leaf number (55% more), and survivorship (47% greater) than seedlings without such access, suggesting that the ECM network provides a survivorship advantage. A survey of wild seedlings showed positive distance-dependent distribution and survival with respect to conspecific adults. These experimental and survey results suggest that the negative distance-dependent mechanisms at the seedling stage thought to maintain tropical rain forest diversity are reversed for ECM seedlings, which experience positive feedbacks from the ECM network. These results may in part explain the local monodominance of an ECM tree species within the matrix of high-diversity, tropical rain forest.     

Habitat selection increases the isolating effect of habitat fragmentation on the gopher tortoise

Habitat selection requires choice, which differentiates it from habitat use, and choice, in turn, is dependent upon the responses of organisms to the environmental, social, and other cues that they perceive. Habitat selection by the gopher tortoise (Gopherus polyphemus) was investigated by translocating tortoises and monitoring their movements within two sites in central Florida. The first site supported a stable preponderance of high-quality habitat, and tortoises avoided areas with a dense tree canopy cover caused by fire exclusion. The second site was badly invaded by an introduced weed, and tortoises avoided areas where the weed had formed a dense monoculture. At both sites, individuals appeared to be responding to visual cues to avoid areas that were relatively dark. In landscapes with relatively large amounts of high-quality habitat, this avoidance behavior serves the gopher tortoise well by keeping individuals within the dominant habitat type. In degraded areas, high-quality habitat often becomes increasingly uncommon, and the avoidance behavior exhibited by the tortoises will result in individuals becoming confined to small patches, causing a significant reduction in fitness and hence questioning their long-term survival in such areas. The results from our study show that in order to maintain viable tortoise populations in areas increasingly subjected to human fragmentation and degradation, it is crucial not only to suppress tree canopy cover continually and prevent invasion by exotic weeds, but also to be mindful that the avoidance behavior of the gopher tortoise could prevent individuals from fully occupying a high-quality habitat in response to restoration and management efforts.     

Associations of forest bird species richness with housing and landscape patterns across the USA.

In the United States, housing density has substantially increased in and adjacent to forests. Our goal in this study was to identify how housing density and human populations are associated with avian diversity. We compared these associations to those between landscape pattern and avian diversity, and we examined how these associations vary across the conterminous forested United States. Using data from the North American Breeding Bird Survey, the U.S. Census, and the National Land Cover Database, we focused on forest and woodland bird communities and conducted our analysis at multiple levels of model specificity, first using a coarse-thematic resolution (basic models), then using a larger number of fine-thematic resolution variables (refined models). We found that housing development was associated with forest bird species richness in all forested ecoregions of the conterminous United States. However, there were important differences among ecoregions. In the basic models, housing density accounted for < 5% of variance in avian species richness. In refined models, 85% of models included housing density and/or residential land cover as significant variables. The strongest guild response was demonstrated in the Adirondack-New England ecoregion, where 29% of variation in richness of the permanent resident guild was associated with housing density. Model improvements due to regional stratification were most pronounced for cavity nesters and short-distance migrants, suggesting that these guilds may be especially sensitive to regional processes. The varying patterns of association between avian richness and attributes associated with landscape structure suggested that landscape context was an important mediating factor affecting how biodiversity responds to landscape changes. Our analysis suggested that simple, broadly applicable, land use recommendations cannot be derived from our results. Rather, anticipating future avian response to land use intensification (or reversion to native vegetation) has to be conditioned on the current landscape context and the species group of interest. Our results show that housing density and residential land cover were significant predictors of forest bird species richness, and their prediction strengths are likely to increase as development continues.     

Extinction debt of forest plants persists for more than a century following habitat fragmentation

Following habitat fragmentation individual habitat patches may lose species over time as they pay off their "extinction debt." Species with relatively low rates of population extinction and colonization ("slow" species) may maintain extinction debts for particularly prolonged periods, but few data are available to test this prediction. We analyzed two unusually detailed data sets on forest plant distributions and land-use history from Lincolnshire, United Kingdom, and Vlaams-Brabant, Belgium, to test for an extinction debt in relation to species-specific extinction and colonization rates. Logistic regression models predicting the presence-absence of 36 plant species were first parameterized using data from Lincolnshire, where forest cover has been relatively low (approximately 5-8%) for the past 1000 years. Consistent with extinction debt theory, for relatively slow species (but not fast species) these models systematically underpredicted levels of patch occupancy in Vlaams-Brabant, where forest cover was reduced from approximately 25% to <10% between 1775 and 1900 (it is presently 6.5%). As a consequence, the ability of the Lincolnshire models to predict patch occupancy in Vlaams-Brabant was worse for slow than for fast species. Thus, more than a century after forest fragmentation reached its current level an extinction debt persists for species with low rates of population turnover.