Long-Term Effects of Group-Selection Timber Harvesting on Abundance of Forest Birds

Abstract:  Relatively few studies have examined the ecological effects of group-selection timber harvesting, and nearly all have been short-term and have lacked experimental manipulations that allow pre- and posttreatment comparisons. We have been documenting the effects of a group-selection timber harvest on bird abundance in a Maine forest for 24 years (preharvest, 1983–1987; postharvest, 1988–2006). Here we characterized the trends in bird abundance over the first 20 years of the study in the managed and control halves of the 40-ha study area. Species responses to the group-selection harvest were idiosyncratic, but in general the mature-forest bird community was retained and species dependent on early successional habitat temporarily (≤8 years) benefited. The Eastern Wood-Pewee ( Contopus virens ) , Winter Wren ( Troglodytes troglodytes ) , Pine Warbler ( Dendroica pinus ) , and White-throated Sparrow ( Zonotrichia albicollis ) increased in abundance in the managed half of the study area following timber harvest, whereas the Veery ( Catharus fuscescens ) decreased. The Black-and-White Warbler ( Mniotilta varia ) , Nashville Warbler ( Vermivora ruficapilla ) , and Common Yellowthroat ( Geothlypis trichas ) responded positively to harvesting, as indicated by decreases in abundance in the control area and more protracted declines or stable abundances in the managed area. This study constitutes the longest experimental investigation to date of the effects of a group-selection harvest on birds and thus provides important information on the strength, direction, and duration of temporal changes in bird populations following forest management.     

Forest structure and light regimes following moderate wind storms: implications for multi-cohort management.

Moderate-severity disturbances appear to be common throughout much of North America, but they have received relatively little detailed study compared to catastrophic disturbances and small gap dynamics. In this study, we examined the immediate impact of moderate-intensity wind storms on stand structure, opening sizes, and light regimes in three hemlock-hardwood forests of northeastern Wisconsin. These were compared to three stands managed by single-tree and group selection, the predominant forest management system for northern hardwoods in the region. Wind storms removed an average of 41% of the stand basal area, compared to 27% removed by uneven-aged harvests, but both disturbances removed trees from a wide range of size classes. The removal of nearly half of the large trees by wind in two old-growth stands caused partial retrogression to mature forest structure, which has been hypothesized to be a major disturbance pathway in the region. Wind storms resulted in residual stand conditions that were much more heterogeneous than in managed stands. Gap sizes ranged from less than 10 m2 up to 5000 m2 in wind-disturbed stands, whereas the largest opening observed in managed stands was only 200 m2. Wind-disturbed stands had, on average, double the available solar radiation at the forest floor compared to managed stands. Solar radiation levels were also more heterogeneous in wind-disturbed stands, with six times more variability at small scales (0.1225 ha) and 15 times more variability at the whole-stand level. Modification of uneven-aged management regimes to include occasional harvests of variable intensity and spatial pattern may help avoid the decline in species diversity that tends to occur after many decades of conventional uneven-aged management. At the same time, a multi-cohort system with these properties would retain a high degree of average crown cover, promote structural heterogeneity typical of old-growth forests, and maintain dominance by late-successional species.     

Habitat-tree protection concepts over 200 years

The protection and sustainable management of habitat trees is an integral part of modern forest nature conservation concepts such as retention forestry. Bats, cavity-nesting birds, arboreal marsupials, and many different saproxylic species depend on habitat trees and their great variety of microhabitats and old-growth characteristics. With a focus on insights from temperate forests, we traced the development of habitat-tree protection over 200 years. The idea was first conceptualized by foresters and natural scientists in the early 19th century. At that time, utilitarian conservation aimed to protect cavity trees that provided roosts and nesting holes for insectivorous bats and birds. By the second half of the 19th century, habitat-tree protection was well known to foresters and was occasionally implemented. Knowledge of the protection of large old trees, a special kind of habitat tree, for sociocultural and aesthetic reasons developed similarly. But, many foresters of that time and in the following decades fundamentally rejected protection of habitat trees for economic reasons. Beginning in the 1970s, forest conservation and integrative forest management became increasingly important issues worldwide. Since then, the protection of habitat trees has been implemented on a large scale. Long-term views on the development of conservation concepts are important to inform the implementation of conservation today. In particular, historical analyses of conservation concepts allow the testing of long-term conservation outcomes and make it possible to study the resilience of conservation approaches to changing social or ecological conditions. We encourage all conservation ecologists to assess the practical and conceptual impact of the initial ideas that led to modern conservation concepts in terms of long-term biodiversity conservation.     

Limited salvage logging effects on forest regeneration after moderate-severity windthrow.

Recent conceptual advances address forest response to multiple disturbances within a brief time period, providing an ideal framework for examining the consequences of natural disturbances followed by anthropogenic management activities. The combination of two or more disturbances in a short period may produce "ecological surprises," and models predict a threshold of cumulative disturbance severity above which forest composition will be drastically altered and regeneration may be impaired. Salvage logging (the harvesting of timber after natural disturbances; also called "salvaging" or "sanitary logging") is common, but there have been no tests of the manner in which salvaging after natural wind disturbance affects woody plant regeneration. Here we present findings from three years after a moderate-severity wind disturbance in west-central Tennessee, USA. We compare two unsalvaged sites and two sites that had intermediate-intensity salvaging. Our approach demonstrates the calculation of cumulative severity measures, which combine natural windthrow severity and anthropogenic tree cutting and removal, on a plot-by-plot basis. Seedling/sapling density and species richness were not influenced by cumulative disturbance severity, but species diversity showed a marginal increase with increasing cumulative severity. The amount of compositional change (from predisturbance trees to post-disturbance seedlings/saplings) increased significantly with cumulative severity of disturbance but showed no evidence of thresholds within the severity range examined. Overall, few deleterious changes were evident in these sites. Moderate-severity natural disturbances followed by moderate-intensity salvaging may have little detrimental effect on forest regeneration and diversity in these systems; the ecological surprises and threshold compositional change are more likely after combinations of natural and anthropogenic disturbances that have a much greater cumulative severity.     

Effects of timber harvesting on pond-breeding amphibian persistence: testing the evacuation hypothesis.

Numerous studies have documented the decline of amphibians following timber harvest. However, direct evidence concerning the mechanisms of population decline is lacking and hinders attempts to develop conservation or recovery plans and solutions for forest species. We summarized the mechanisms by which abundance of amphibians may initially decline following timber harvest into three testable hypotheses: (1) mortality, (2) retreat, and (3) evacuation. Here, we tested the evacuation hypothesis within a large-scale, replicated experiment. We used drift fences with pitfall traps to capture pond-breeding amphibians moving out of experimental clearcut quadrants and into control quadrants at four replicate arrays located within the Daniel Boone Conservation Area on the upper Ozark Plateau in Warren County, Missouri, USA. During the preharvest year of 2004, only 51.6% of the 312 individuals captured were moving out of pre-clearcut quadrants, and movement did not differ from random. In contrast, during both postharvest years of 2005 and 2006, the number of captures along the quadrant edge increased, and a higher proportion of individuals (59.9% and 56.6%, respectively, by year) were moving out of clearcut quadrants than entering. Salamanders moved out of clearcuts in large percentages (Ambystoma annulatum, 78.2% in 2005, 78.2% in 2006; A. maculatum, 64.0% in 2005, 57.1% in 2006). Frogs and toads also moved out of clearcut quadrants, but in lower percentages (Bufo americanus, 59.6% in 2005, 53.3% in 2006; Rana clamitans, 52.7% in 2006). Salamanders moved out of clearcuts with low-wood treatments more than out of clearcuts with high-wood treatments. Movement of salamanders out of clearcuts was independent of sex. Estimated movement out of clearcuts represented between 8.7% and 35.0% of the total breeding adults captured for two species of salamanders. Although we recognize that some portion of the amphibian population may retreat underground for short periods and others may not survive the effects of timber harvest, these data are the first direct evidence showing that individuals are capable of leaving clearcuts and shifting habitat use.     

Impacts of Alternative Timber Harvest Practices on Leaf-Chewing Herbivores of Oak

Abstract:  Studies of the effects of logging on Lepidoptera rarely address landscape-level effects or effects on larval, leaf-feeding stages. We examined the impacts of uneven-aged and even-aged logging on the abundance, richness, and community structure of leaf-chewing insects of white ( Quercus alba L.) and black ( Q. velutina L.) oak trees remaining in unharvested areas by sampling 3 years before and 7 years after harvest. After harvest, white oaks in uneven-aged sites had 32% fewer species of leaf-chewing insects than control sites. This reduction in species richness may have resulted from changes in microclimate (reducing plant quality and/or changing leaf phenology) that affected a much larger total area of each site than did even-aged cuts. For black oak after harvest, species richness in uneven- and even-aged sites increased relative to levels before harvest. Harvesting did not alter total insect density or community structure in the unlogged habitat for either oak species with one exception: insect density on black oak increased in the oldest forest block. Community structure of herbivores of black and white oaks in clearcut gaps differed from that of oaks in intact areas of even-aged sites. Furthermore, both richness and total insect density of black oaks were reduced in clearcut gaps. We suggest that low-level harvests alter herbivore species richness at the landscape level. Treatment effects were subtle because we sampled untreated areas of logged landscapes, only one harvest had occurred, and large temporal and spatial variation in abundance and richness existed. Although the effects of logging were greater in uneven-aged sites, the effects of even-aged management are likely to increase as harvesting continues.     

Recovery of forest structure and spectral properties after selective logging in lowland Bolivia.

Effective monitoring of selective logging from remotely sensed data requires an understanding of the spatial and temporal thresholds that constrain the utility of those data, as well as the structural and ecological characteristics of forest disturbances that are responsible for those constraints. Here we assess those thresholds and characteristics within the context of selective logging in the Bolivian Amazon. Our study combined field measurements of the spatial and temporal dynamics of felling gaps and skid trails ranging from <1 to 19 months following reduced-impact logging in a forest in lowland Bolivia with remote-sensing measurements from simultaneous monthly ASTER satellite overpasses. A probabilistic spectral mixture model (AutoMCU) was used to derive per-pixel fractional cover estimates of photosynthetic vegetation (PV), non-photosynthetic vegetation (NPV), and soil. Results were compared with the normalized difference in vegetation index (NDVI). The forest studied had considerably lower basal area and harvest volumes than logged sites in the Brazilian Amazon where similar remote-sensing analyses have been performed. Nonetheless, individual felling-gap area was positively correlated with canopy openness, percentage liana coverage, rates of vegetation regrowth, and height of remnant NPV. Both liana growth and NPV occurred primarily in the crown zone of the felling gap, whereas exposed soil was limited to the trunk zone of the gap. In felling gaps >400 m2, NDVI, and the PV and NPV fractions, were distinguishable from unlogged forest values for up to six months after logging; felling gaps <400 m2 were distinguishable for up to three months after harvest, but we were entirely unable to distinguish skid trails from our analysis of the spectral data.     

Toward Meaningful Snag-Management Guidelines for Postfire Salvage Logging in North American Conifer Forests

Abstract:  The bird species in western North America that are most restricted to, and therefore most dependent on, severely burned conifer forests during the first years following a fire event depend heavily on the abundant standing snags for perch sites, nest sites, and food resources. Thus, it is critical to develop and apply appropriate snag-management guidelines to implement postfire timber harvest operations in the same locations. Unfortunately, existing guidelines designed for green-tree forests cannot be applied to postfire salvage sales because the snag needs of snag-dependent species in burned forests are not at all similar to the snag needs of snag-dependent species in green-tree forests. Birds in burned forests have very different snag-retention needs from those cavity-nesting bird species that have served as the focus for the development of existing snag-management guidelines. Specifically, many postfire specialists use standing dead trees not only for nesting purposes but for feeding purposes as well. Woodpeckers, in particular, specialize on wood-boring beetle larvae that are superabundant in fire-killed trees for several years following severe fire. Species such as the Black-backed Woodpecker ( Picoides arcticus ) are nearly restricted in their habitat distribution to severely burned forests. Moreover, existing postfire salvage-logging studies reveal that most postfire specialist species are completely absent from burned forests that have been (even partially) salvage logged. I call for the long-overdue development and use of more meaningful snag-retention guidelines for postfire specialists, and I note that the biology of the most fire-dependent bird species suggests that even a cursory attempt to meet their snag needs would preclude postfire salvage logging in those severely burned conifer forests wherein the maintenance of biological diversity is deemed important.     

Effects of Trophy Hunting on Lion and Leopard Populations in Tanzania

Abstract: Tanzania holds most of the remaining large populations of African lions (Panthera leo) and has extensive areas of leopard habitat (Panthera pardus), and both species are subjected to sizable harvests by sport hunters. As a first step toward establishing sustainable management strategies, we analyzed harvest trends for lions and leopards across Tanzania's 300,000 km² of hunting blocks. We summarize lion population trends in protected areas where lion abundance has been directly measured and data on the frequency of lion attacks on humans in high‐conflict agricultural areas. We place these findings in context of the rapidly growing human population in rural Tanzania and the concomitant effects of habitat loss, human‐wildlife conflict, and cultural practices. Lion harvests declined by 50% across Tanzania between 1996 and 2008, and hunting areas with the highest initial harvests suffered the steepest declines. Although each part of the country is subject to some form of anthropogenic impact from local people, the intensity of trophy hunting was the only significant factor in a statistical analysis of lion harvest trends. Although leopard harvests were more stable, regions outside the Selous Game Reserve with the highest initial leopard harvests again showed the steepest declines. Our quantitative analyses suggest that annual hunting quotas be limited to 0.5 lions and 1.0 leopard/1000 km² of hunting area, except hunting blocks in the Selous Game Reserve, where harvests should be limited to 1.0 lion and 3.0 leopards/1000 km².     

Using forest inventory data to assess fisher resting habitat suitability in California.

The fisher (Martes pennanti) is a forest-dwelling carnivore whose current distribution and association with late-seral forest conditions make it vulnerable to stand-altering human activities or natural disturbances. Fishers select a variety of structures for daily resting bouts. These habitat elements, together with foraging and reproductive (denning) habitat, constitute the habitat requirements of fishers. We develop a model capable of predicting the suitability of fisher resting habitat using standard forest vegetation inventory data. The inventory data were derived from Forest Inventory and Analysis (FIA), a nationwide probability-based sample used to estimate forest characteristics. We developed the model by comparing vegetation and topographic data at 75 randomly selected fisher resting structures in the southern Sierra Nevada with 232 forest inventory plots. We collected vegetation data at fisher resting locations using the FIA vegetation sampling protocol and centering the 1-ha FIA plot on the resting structure. To distinguish used and available inventory plots, we used nonparametric logistic regression to evaluate a set of a priori biological models. The top model represented a dominant portion of the Akaike weights (0.87), explained 31.5% of the deviance, and included the following variables: average canopy closure, basal area of trees <51 cm diameter breast height (dbh), average hardwood dbh, maximum tree dbh, percentage slope, and the dbh of the largest conifer snag. Our use of routinely collected forest inventory data allows the assessment and monitoring of change in fisher resting habitat suitability over large regions with no additional sampling effort. Although models were constrained to include only variables available from the list of those measured using the FIA protocol, we did not find this to be a shortcoming. The model makes it possible to compare average resting habitat suitability values before and after forest management treatments, among administrative units, across regions and over time. Considering hundreds of plot estimates as a sample of habitat conditions over large spatial scales can bring a broad perspective, at high resolution, and efficiency to the assessment and monitoring of wildlife habitat.     

Influence of overstory removal on growth of epiphytic mosses and lichens in western Oregon.

Will old-growth-associated epiphytes survive if the forest canopy is opened around them by thinning or partial harvest? If old-growth association is due to a species' environmental tolerances, it may not survive in the relatively open stands that result from such treatments. If, however, old-growth association is due to dispersal limitations rather than environmental tolerances, retention of host trees as refugia and sources of inoculum might carry populations of old-growth-associated epiphytes into young stands. We studied growth rates of lichen and moss transplants in a Douglas-fir (Pseudotsuga menziesii) forest (tree ages approximately 55 yr) in western Oregon for nine months before and 27 months after moderate thinning, creation of 0.4-ha patch cuts, and in control areas. We also assessed moss sporophyte production. We contrasted responses of one moss species, Isothecium myosuroides sensu lato, which is ubiquitous in forests of varying ages, with those of another moss, Antitrichia curtipendula, and a lichen, Lobaria oregana, which are both associated with old-growth forests. Both old-growth associates grew faster in thinned areas and patch cuts than in controls, while Isothecuim grew most slowly and produced fewest sporophytes in patch cuts. These species are likely to survive in remnants, assuming they can remain attached, and may be successful in young stands if they can disperse and establish there. Our results suggest that logging with green-tree retention and other silvicultural practices that preserve trees or shrubs hosting the species studied here are likely to encourage these species' development in managed forests.     

The effects of forest harvest intensity in combination with wind disturbance on carbon dynamics in Lake States Mesic Forests

Total forest carbon (C) storage is determined by succession, disturbances, climate, and the edaphic properties of a site or region. Forest harvesting substantially affects C dynamics; these effects may be amplified if forest harvesting is intensified to provide biofuel feedstock. We tested the effects of harvest intensity on landscape C using a simulation modeling approach that included C dynamics, multiple disturbances, and successional changes in composition. We developed a new extension for the LANDIS-II forest landscape disturbance and succession model that incorporates belowground soil C dynamics derived from the CENTURY soil model. The extension was parameterized and calibrated using data from an experimental forest in northeastern Wisconsin, USA. We simulated a 9800 ha forested landscape over 400 years with wind disturbance combined with no harvesting, harvesting with residual slash left on site (‘standard harvest’), and whole-tree harvesting. We also simulated landscapes without wind disturbance and without eastern hemlock (Tsuga canadensis) to examine the effects of detrital quantity and quality on C dynamics. We estimated changes in live C, detrital C, soil organic C, total C, and forest composition. Overall, the simulations without harvesting had substantially greater total C and continued to sequester C. Standard harvest simulations had more C than the whole tree harvest simulations. Under both harvest regimes, C accrual was not evident after 150 years. Without hemlock, SOC was reduced due to a decline in detritus and a shift in detrital chemistry. In conclusion, if the intensity of harvesting increases we can expect a corresponding reduction in potential C storage. Compositional changes due to historic circumstances (loss of hemlock) may also affect forest C although to a lesser degree than harvesting. The modeling approach presented enabled us to consider multiple, interacting drivers of landscape change and the subsequent changes in forest C.     

Bat Use of Remnant Old-Growth Redwood Stands

Most of the old-growth redwood ( Sequoia sempervirens ) in California has been cut; regenerating forests will probably never resemble those that were harvested, and what old growth remains on private land occurs in small, isolated remnant patches. The landscapes in which these stands occur differ so markedly from their original condition that their value as habitat to many species of wildlife, including bats, is unknown. Previous research in unfragmented redwood forests demonstrated that bats use basal hollows in old-growth redwoods as roosts. We sought to determine whether bats use similar old-growth trees as roosts when they occur in small, remnant patches of isolated old growth on commercial forest land. We compared bat occurrence in remnant and contiguous stands by collecting guano in traps suspended in hollows and by monitoring flight activity with ultrasonic bat detectors. Hollows in trees within the remnant stands had significantly more guano deposited per tree than the trees within the contiguous forest. The mean numbers of bat passes per night were statistically indistinguishable between the two treatments, although mean flight activity in the remnant stands was greater than in the contiguous forest. Bats frequently used basal hollows in small (<5 ha) stands of remnant old growth, which may be due to the closer proximity of remnants to stream courses, to their greater interface with edge where foraging success may be greater, or to the fact that the lower density of hollow-bearing trees in remnants than in contiguous forest favored greater use per tree. Significant use of small, residual old-growth redwood provides reason to maintain these remnants in managed landscapes as potentially important habitat for forest bats.     

Examining the effects of alternative management strategies on landscape-scale forest patterns in northeastern Minnesota using LANDIS

Spatial modeling of forest patterns can provide information on the potential impact of various management strategies on large landscapes over long time frames. We used LANDIS, a stochastic, spatially-explicit, ecological landscape model to simulate 120 years of forest change on the Nashwauk Uplands, a 328,000 ha landscape in northeastern Minnesota that lies in the transition between boreal and temperate forests. We ran several forest management scenarios including current harvesting practices, no harvests, varied rotation ages, varied clearcut sizes, clustered clearcuts, and landowner coordination. We examined the effects of each scenario on spatial patterns of forests by covertype, age class, and mean and distribution of patch sizes. All scenarios reveal an increase in the spruce-fir (Picea-Abies) covertype relative to the economically paramount aspen-birch (Populus-Betula) covertype. Our results also show that most covertypes occur in mostly small patches <5 ha in size and the ability of management to affect patch size is limited by the highly varied physiography and landuse patterns on the landscape. However, coordination among landowners, larger clearcuts, and clustered clearcuts were all predicted to increase habitat diversity by creating some larger patches and older forest patches. These three scenarios along with the no harvest scenario also create more old forest than current harvesting practices, by concentrating harvesting on some portion of the landscape. The no harvest scenario retained large, fire-regenerated aspen-birch patches. Harvests fragment large aspen-birch patches by changing the age structure and releasing the shade-tolerant understory species. More sapling forest, and larger sapling patches resulted from the shortened rotation scenario.     

Effects of Fragmentation of Araucarian Vine Forest on Small Mammal Communities

Abstract: Habitat loss and fragmentation are major threats to the survival of forest-dependent fauna. We examined the abundance of small mammal species in forests, corridors, remnants of araucarian vine forest, and Araucaria cunninghamii plantations and pastures. None of the forest mammal species persisted following conversion of forest to pasture. Plantations supported lowered abundances of a subset of forest species that were mainly habitat generalists with respect to their occurrence in different floristic types of undisturbed native forest. Within plantations, an increased subcanopy cover was associated with a more forest-like small mammal assemblage. Species' responses to habitat fragmentation varied. The floristic habitat generalists were largely tolerant of habitat fragmentation, their abundance being similar in forests, corridors, and remnants, and were capable of persisting in remnants a few hectares in area. Floristic habitat specialists were vulnerable to habitat fragmentation and thus were abundant in continuous forest, were less abundant in corridors, and were generally absent from remnants. Species that avoid the corridor matrix and are therefore constrained to the corridor may be disadvantaged by the linearity of the habitat, consistent with the predictions of central-place foraging theory. Although small remnants and corridors provide habitat for some species, those that are more specialized in their use of undisturbed habitat types require the retention or reestablishment of large intact areas.     

The influence of land use and climate change on forest biomass and composition in Massachusetts, USA

Land use and climate change have complex and interacting effects on naturally dynamic forest landscapes. To anticipate and adapt to these changes, it is necessary to understand their individual and aggregate impacts on forest growth and composition. We conducted a simulation experiment to evaluate regional forest change in Massachusetts, USA over the next 50 years (2010-2060). Our objective was to estimate, assuming a linear continuation of recent trends, the relative and interactive influence of continued growth and succession, climate change, forest conversion to developed uses, and timber harvest on live aboveground biomass (AGB) and tree species composition. We examined 20 years of land use records in relation to social and biophysical explanatory variables and used regression trees to create "probability-of-conversion" and "probability-of-harvest" zones. We incorporated this information into a spatially interactive forest landscape simulator to examine forest dynamics as they were affected by land use and climate change. We conducted simulations in a full-factorial design and found that continued forest growth and succession had the largest effect on AGB, increasing stores from 181.83 Tg to 309.56 Tg over 50 years. The increase varied from 49% to 112% depending on the ecoregion within the state. Compared to simulations with no climate or land use, forest conversion reduced gains in AGB by 23.18 Tg (or 18%) over 50 years. Timber harvests reduced gains in AGB by 5.23 Tg (4%). Climate change (temperature and precipitation) increased gains in AGB by 17.3 Tg (13.5%). Pinus strobus and Acer rubrum were ranked first and second, respectively, in terms of total AGB throughout all simulations. Climate change reinforced the dominance of those two species. Timber harvest reduced Quercus rubra from 10.8% to 9.4% of total AGB, but otherwise had little effect on composition. Forest conversion was generally indiscriminate in terms of species removal. Under the naive assumption that future land use patterns will resemble the recent past, we conclude that continued forest growth and recovery will be the dominant mechanism driving forest dynamics over the next 50 years, and that while climate change may enhance growth rates, this will be more than offset by land use, primarily forest conversion to developed uses.     

Conservation of Epiphyte Diversity in an Andean Landscape Transformed by Human Land Use

Abstract: Epiphytes are diverse and important elements of tropical forests, but as canopy‐dwelling organisms, they are highly vulnerable to deforestation. To assess the effect of deforestation on epiphyte diversity and the potential for epiphyte conservation in anthropogenically transformed habitats, we surveyed the epiphytic vegetation of an Ecuadorian cloud forest reserve and its surroundings. Our study was located on the western slopes of the Andes, a global center of biodiversity. We sampled vascular epiphytes of 110 study plots in a continuous primary forest; 14 primary forest fragments; isolated remnant trees in young, middle‐aged, and old pastures; and young and old secondary forests. It is the first study to include all relevant types of habitat transformation at a single study site and to compare epiphyte diversity at different temporal stages of fragmentation. Epiphyte diversity was highest in continuous primary forest, followed by forest fragments and isolated remnant trees, and lowest in young secondary forests. Spatial parameters of habitat transformation, such as fragment area, distance to the continuous primary forest, or distance to the forest edge from inside the forest, had no significant effect on epiphyte diversity. Hence, the influence of dispersal limitations appeared to be negligible or appeared to operate only over very short distances, whereas microclimatic edge effects acted only in the case of completely isolated trees, but not in larger forest fragments. Epiphyte diversity increased considerably with age of secondary forests, but species assemblages on isolated remnant trees were impoverished distinctly with time since isolation. Thus, isolated trees may serve for recolonization of secondary forests, but only for a relatively short time. We therefore suggest that the conservation of even small patches of primary forest within agricultural landscape matrices is essential for the long‐term maintenance of the high epiphyte diversity in tropical cloud forests.     

Mortality of large trees and lianas following experimental drought in an Amazon forest

Severe drought episodes such as those associated with El Ni?o Southern Oscillation (ENSO) events influence large areas of tropical forest and may become more frequent in the future. One of the most important forest responses to severe drought is tree mortality, which alters forest structure, composition, carbon content, and flammability, and which varies widely. This study tests the hypothesis that tree mortality increases abruptly during drought episodes when plant-available soil water (PAW) declines below a critical minimum threshold. It also examines the effect of tree size, plant life form (palm, liana, tree) and potential canopy position (understory, midcanopy, overstory) on drought-induced plant mortality. A severe, four-year drought episode was simulated by excluding 60% of incoming throughfall during each wet season using plastic panels installed in the understory of a 1-ha forest treatment plot, while a 1-ha control plot received normal rainfall. After 3.2 years, the treatment resulted in a 38% increase in mortality rates across all stems >2 cm dbh. Mortality rates increased 4.5-fold among large trees (>30 cm dbh) and twofold among medium trees (10-30 cm dbh) in response to the treatment, whereas the smallest stems were less responsive. Recruitment rates did not compensate for the elevated mortality of larger-diameter stems in the treatment plot. Overall, lianas proved more susceptible to drought-induced mortality than trees or palms, and potential overstory tree species were more vulnerable than midcanopy and understory species. Large stems contributed to 90% of the pretreatment live aboveground biomass in both plots. Large-tree mortality resulting from the treatment generated 3.4 times more dead biomass than the control plot. The dramatic mortality response suggests significant, adverse impacts on the global carbon cycle if climatic changes follow current trends.     

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.