Acceleration of Exotic Plant Invasion in a Forested Ecosystem by a Generalist Herbivore

Abstract: The successful invasion of exotic plants is often attributed to the absence of coevolved enemies in the introduced range (i.e., the enemy release hypothesis). Nevertheless, several components of this hypothesis, including the role of generalist herbivores, remain relatively unexplored. We used repeated censuses of exclosures and paired controls to investigate the role of a generalist herbivore, white‐tailed deer (Odocoileus virginianus), in the invasion of 3 exotic plant species (Microstegium vimineum, Alliaria petiolata, and Berberis thunbergii) in eastern hemlock (Tsuga canadensis) forests in New Jersey and Pennsylvania (U.S.A.). This work was conducted in 10 eastern hemlock (T. canadensis) forests that spanned gradients in deer density and in the severity of canopy disturbance caused by an introduced insect pest, the hemlock woolly adelgid (Adelges tsugae). We used maximum likelihood estimation and information theoretics to quantify the strength of evidence for alternative models of the influence of deer density and its interaction with the severity of canopy disturbance on exotic plant abundance. Our results were consistent with the enemy release hypothesis in that exotic plants gained a competitive advantage in the presence of generalist herbivores in the introduced range. The abundance of all 3 exotic plants increased significantly more in the control plots than in the paired exclosures. For all species, the inclusion of canopy disturbance parameters resulted in models with substantially greater support than the deer density only models. Our results suggest that white‐tailed deer herbivory can accelerate the invasion of exotic plants and that canopy disturbance can interact with herbivory to magnify the impact. In addition, our results provide compelling evidence of nonlinear relationships between deer density and the impact of herbivory on exotic species abundance. These findings highlight the important role of herbivore density in determining impacts on plant abundance and provide evidence of the operation of multiple mechanisms in exotic plant invasion.     

Exploitative competition between invasive herbivores benefits a native host plant

Although biological invasions are of considerable concern to ecologists, relatively little attention has been paid to the potential for and consequences of indirect interactions between invasive species. Such interactions are generally thought to enhance invasives' spread and impact (i.e., the "invasional meltdown" hypothesis); however, exotic species might also act indirectly to slow the spread or blunt the impact of other invasives. On the east coast of the United States, the invasive hemlock woolly adelgid (Adelges tsugae, HWA) and elongate hemlock scale (Fiorinia externa, EHS) both feed on eastern hemlock (Tsuga canadensis). Of the two insects, HWA is considered far more damaging and disproportionately responsible for hemlock mortality. We describe research assessing the interaction between HWA and EHS, and the consequences of this interaction for eastern hemlock. We conducted an experiment in which uninfested hemlock branches were experimentally infested with herbivores in a 2 x 2 factorial design (either, both, or neither herbivore species). Over the 2.5-year course of the experiment, each herbivore's density was approximately 30% lower in mixed- vs. single-species treatments. Intriguingly, however, interspecific competition weakened rather than enhanced plant damage: growth was lower in the HWA-only treatment than in the HWA + EHS, EHS-only, or control treatments. Our results suggest that, for HWA-infested hemlocks, the benefit of co-occurring EHS infestations (reduced HWA density) may outweigh the cost (increased resource depletion).     

Tsuga canadensis (L.) Carr. mortality will impact hydrologic processes in southern Appalachian forest ecosystems.

Eastern hemlock (Tsuga canadensis (L.) Carr.) is one of the principal riparian and cove canopy species in the southern Appalachian Mountains. Throughout its range, eastern hemlock is facing potential widespread mortality from the hemlock woolly adelgid (HWA). If HWA-induced eastern hemlock mortality alters hydrologic function, land managers will be challenged to develop management strategies that restore function or mitigate impacts. To estimate the impact that the loss of this forest species will have on the hydrologic budget, we quantified and modeled transpiration over a range of tree sizes and environmental conditions. We used heat dissipation probes, leaf-level gas-exchange measurements, allometric scaling, and time series modeling techniques to quantify whole-tree and leaf-level transpiration (E(L)) of eastern hemlock. We monitored trees ranging from 9.5 to 67.5 cm in diameter along a riparian corridor in western North Carolina, USA during 2004 and 2005. Maximum rates of daily tree water use varied by diameter and height, with large trees transpiring a maximum of 178-186 kg H2O x tree(-1) x d(-1). Values of E(L) could be predicted from current and lagged environmental variables. We forecasted eastern hemlock E(L) for inventoried stands and estimated a mean annual transpiration rate of 63.3 mm/yr for the hemlock component, with 50% being transpired in the winter and spring. In typical southern Appalachian stands, eastern hemlock mortality would thus reduce annual stand-level transpiration by approximately 10% and reduce winter and spring stand-level transpiration by approximately 30%. Eastern hemlock in the southern Appalachians has two distinct ecohydrological roles: an evergreen tree that maintains year-round transpiration rates and a riparian tree that has high transpiration rates in the spring. No other native evergreen in the southern Appalachians will likely fill the ecohydrological role of eastern hemlock if widespread mortality occurs. With the loss of this species, we predict persistent increases in discharge, decreases in the diurnal amplitude of streamflow, and increases in the width of the variable source area.     

Forests Too Deer: Edge Effects in Northern Wisconsin

Abstract: Browsing by white-tailed deer (Odocoileus virginianus) can profoundly affect the abundance and population structure of several woody and herbaceous plant species. Enclosure studies and population surveys reveal that past and current deer densities as low as 4 deer/km² may prevent regeneration of the once common woody species, Canada yew (Taxus canadensis), eastern hemlock (Tsuja canadensis), and white cedar Puja occidentalis), as well as several herbaceous species. Prior to European settlement, forests in northern Wisconsin contained relatively sparse deer populations (<4/km²), but extensive timber cutting in the late nineteenth century boosted deer populations. Continued habitat fragmentation resulting from scattered timber harvests and the creation of "wildlife openings" to improve deer forage maintain these high densities throughout much of the Northeast.
Because deer wander widely, the effects of high deer densities penetrate deeply into remaining stands of old and mature forest, greatly modifying their composition Thus, abundant early successional and "edge" habitat, and the high deer densities they engender, represent significant external threats to these plant communities. We hypothesize that establishing large (200–400 km²) continuous areas of maturing forest, especially in conjunction with increased hunting, could reduce local deer densities and so provide a simple and inexpensive method for retaining species sensitive to the deleterious effects of browsing.     

The ecology of energy and nutrient fluxes in hemlock forests invaded by hemlock woolly adelgid

The hemlock woolly adelgid (HWA, Adelges tsugae Annand) is currently causing a severe decline in vitality and survival of eastern hemlock in North American forests. We analyzed the effects of light HWA infestation on vertical energy and nutrient fluxes from the canopy to the forest floor. Canopy throughfall, litter lysimeters, and laboratory litter microcosms were used to examine the effects of HWA-affected and unaffected throughfall on litter type, leachate, and litter chemistry. Early in the season adelgid infestation caused higher dissolved organic carbon (DOC; +24.6%), dissolved organic nitrogen (DON; +28.5%), and K (+39.3%) fluxes and lower inorganic nitrogen fluxes (-39.8%) in throughfall and in adjacent litter solutions collected beneath infested compared to uninfested trees. Needle litter collected beneath uninfested hemlock had significantly lower N concentrations compared to needles collected beneath infested trees, while no difference in N concentrations was found in birch litter. Bacteria were significantly more abundant on hemlock and birch litter beneath infested trees, while yeasts and filamentous fungi showed no consistent response to HWA throughfall. Litter microcosms showed that less DOC was leaching from birch than from hemlock needles when exposed to HWA throughfall. Overall, NH4-N and DON leachate concentrations were higher from birch than from hemlock litter. Thus, HWA-affected throughfall leads to qualitative and quantitative differences in nitrogen export from the litter layer. The N concentration of hemlock litter did not change with time, but the N concentration in birch litter increased significantly during the course of the experiment, especially when HWA-affected throughfall was applied. We suggest a nonlinear conceptual model for the temporal and vertical transition of energy and nutrient fluxes relative to progressing HWA infestation from a pure hemlock to a birch/maple-dominated forest. Progressive needle loss and changes in needle chemistry are likely to produce a humped-shaped DOC curve, while N fluxes initially decrease as infestation continues but rise eventually with hemlock decline and immigration of hardwood species. These findings suggest that it is necessary to understand the biology and specific physiological/trophic effects of exotic pests on their hosts and associated ecosystem processes in order to decipher the temporal dynamics, direction of change, and functional consequences.     

Applying Principles of Landscape Design and Management to Integrate Old-Growth Forest Enhancement and Commodity Use

Using geographic information systems (GIS) and spatial analysis techniques, we developed a landscape design to maintain old-growth forest remnants and integrate commodity production in the surrounding second-growth matrix. The 4500-ha forest landscape in northern Wisconsin contains scattered patches of old-growth eastern hemlock ( Tsuga canadensis ) and northern hardwoods, predominately sugar maple ( Acer saccharum ). The design incorporates an old-growth restoration zone surrounding old-growth patches to buffer and enhance forest-interior habitat and link nearby old-growth remnants. This addition restores aspects of landscape patch size and structure and ecosystem juxtaposition that characterize a nearby, large, and contiguous natural old-growth landscape. A larger secondary zone is delineated for uneven-aged forest management. This zone provides a matrix structurally similar to the old-growth patches but also accommodates harvesting. A larger outer zone is retained primarily in even-aged forest of aspen ( Populus tremuloides ) and paper birch ( Betula papyrifera ), but traditional clearcutting practices are modified to partial cutting and mixed-species rotations. This design meets limited goals of biodiversity enhancement and integrated commodity production in a landscape that will remain largely harvested. The landscape design is therefore improved not only by buffers and corridors provided to old-growth ecosystems, but by modifying the management of the majority commodity lands matrix as well.     

Indirect Effects of Pandemic Deer Overabundance Inferred from Caterpillar‐Host Relations

Externally feeding phytophagous insect larvae (i.e., caterpillars, here, larval Lepidoptera and sawflies, Hymenoptera: Symphyta) are important canopy herbivores and prey resources in temperate deciduous forests. However, composition of forest trees has changed dramatically in the eastern United States since 1900. In particular, browsing by high densities of white‐tailed deer (Odocoileus virginianus) has resulted in forests dominated by browse‐tolerant species, such as black cherry (Prunus serotina), and greatly reduced relative abundance of other tree species, notably pin cherry (Prunus pensylvanica) and birches (Betula spp.). To quantify effects of these changes on caterpillars, we sampled caterpillars from 960 branch tips of the 8 tree species that comprise 95% of trees in Allegheny hardwood forests: red maple (Acer rubrum), striped maple (Acer pensylvanicum), sugar maple (Acer saccharum), sweet birch (Betula lenta), yellow birch (Betula allegheniensis), American beech (Fagus grandifolia), black cherry, and pin cherry. We collected 547 caterpillar specimens that belonged to 66 Lepidoptera and 10 Hymenoptera species. Caterpillar density, species richness, and community composition differed significantly among tree species sampled. Pin cherry, nearly eliminated at high deer density, had the highest density and diversity of caterpillars. Pin cherry shared a common caterpillar community with black cherry, which was distinct from those of other tree hosts. As high deer density continues to replace diverse forests of cherries, maples, birches, and beech with monodominant stands of black cherry, up to 66% of caterpillar species may be eliminated. Hence, deer‐induced changes in forest vegetation are likely to ricochet back up forest food webs and therefore negatively affect species that depend on caterpillars and moths for food and pollination. Efectos Indirectos de la Sobreabundancia de Venados Pandémicos Inferida de Relaciones Orugas‐Huéspedes     

Modelling forest succession in two southeastern Canadian mixedwood ecosystem types using the ZELIG model

The gap model ZELIG was validated for red spruce–balsam fir–yellow birch and yellow birch–sugar maple–balsam fir forest types in southern Quebec, Canada. Long-term historical data originating from the Lake Edward Experimental Forest, La Mauricie National Park, were used. The effect of the variation in plot size, representing the space within which trees uptake site resources, was also examined. Several species were included in both forest types: red spruce (Picea rubens Sarg.), balsam fir (Abies balsamea (L.) Mill.), yellow birch (Betula alleghaniensis Britton), white birch (Betula papyrifera Marsh.), red maple (Acer rubrum L.), sugar maple (Acer saccharum Marsh.), American beech (Fagus grandifolia Ehrh.), eastern hemlock (Tsuga canadensis (L.) Carr.) and northern white cedar (Thuja occidentalis L.). The pattern of change in basal area growth varied among species, ranging from a steady increase to a more or less rapid decline. There was a good agreement between observations and predictions for yellow birch, red spruce, red maple, sugar maple, balsam fir and northern white cedar. Plot size had a significant impact on the dynamics of the different species. Depending on the species, the decline was accelerated, the amplitude of the fluctuations varied, or the maximum basal area reached changed. Predicted regeneration varied among species and the number of seedlings generally increased with increase in plot size. The pattern of development for most species was related to their life characteristics. The results highlighted the fact that there is a critical lack of knowledge and data on the dynamics of regeneration from the seedling to the sapling stages for the two forest types studied, which resulted in poor predictions for some species. As the life characteristics varied among species, the use of only one plot size for all species may not be realistic.     

Managing the Abundance and Diversity of Breeding Bird Populations through Manipulation of Deer Populations

Abstract: Deer densities in forests of eastern North America are thought to have significant effects on the abundance and diversity of forest birds through the role deer play in structuring forest understories. We tested the ability of deer to affect forest bird populations by monitoring the density and diversity of vegetation and birds for 9 years at eight 4-ha sites in northern Virginia, four of which were fenced to exclude deer. Both the density and diversity of understory woody plants increased following deer exclosure. The numerical response of the shrubs to deer exclosure was significantly predicted by the soil quality (ratio of organic carbon to nitrogen) at the sites. Bird populations as a whole increased following exclosure of deer, particularly for ground and intermediate canopy species. The diversity of birds did not increase significantly following exclosure of deer, however, primarily because of replacement of species as understory vegetation proceeded through successional processes. Changes in understory vegetation accounted for most of the variability seen in the abundance and diversity of bird populations. Populations of deer in protected areas are capable of causing significant shifts in the composition and abundance of bird communities. These shifts can be reversed by increasing the density and diversity of understory vegetation, which can be brought about by reducing deer density.     

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.     

A climatic driver for abrupt mid-Holocene vegetation dynamics and the hemlock decline in New England

The mid-Holocene decline of eastern hemlock is widely viewed as the sole prehistorical example of an insect- or pathogen-mediated collapse of a North American tree species and has been extensively studied for insights into pest-host dynamics and the consequences to terrestrial and aquatic ecosystems of dominant-species removal. We report paleoecological evidence implicating climate as a major driver of this episode. Data drawn from sites across a gradient in hemlock abundance from dominant to absent demonstrate: a synchronous, dramatic decline in a contrasting taxon (oak); changes in lake sediments and aquatic taxa indicating low water levels; and one or more intervals of intense drought at regional to continental scales. These results, which accord well with emerging climate reconstructions, challenge the interpretation of a biotically driven hemlock decline and highlight the potential for climate change to generate major, abrupt dynamics in forest ecosystems.     

Interactive Effects of Harvest and Deer Herbivory on the Population Dynamics of American Ginseng

Abstract:  Few demographic models for any species consider the role of multiple, interacting ecological threats. Many forest herbs are heavily browsed by white-tailed deer ( Odocoileus virginianus ) and a number of these are also harvested for the medicinal, floral, or horticultural trades. Previous studies of the viability of American ginseng ( Panax quinquefolius ) have separately examined the effects of harvesting and deer herbivory. We followed individually marked ginseng plants in 6 populations for 8 years and documented deer browse levels, conducted helicopter surveys to estimate the deer herd size, and documented 2 ginseng harvests. We used this long-term data set to develop a stochastic demographic model that quantified the separate and interactive role of these threats to ginseng viability. Although harvesting and deer herbivory negatively affected ginseng population growth, their effects were not additive. Deer herbivory negatively affected population growth in the absence but not in the presence of harvesting. Life table response experiments revealed that in the presence of harvesting, deer herbivory had some positive effects on vital rates because browsed plants were less apparent to harvesters. Ginseng populations that were harvested responsibly (i.e., planting seeds from harvested individuals) had higher growth rates than those that were harvested irresponsibly. We concluded that both deer populations and harvesting must be managed to ensure sustainable populations of American ginseng. Our findings underscore the importance of long-term monitoring to assess threats to viability and the need for a broad ecological understanding of the complexity of ecosystem management.     

Modeling range dynamics in heterogeneous landscapes: invasion of the hemlock woolly adelgid in eastern North America

Range expansion by native and exotic species will continue to be a major component of global change. Anticipating the potential effects of changes in species distributions requires models capable of forecasting population spread across realistic, heterogeneous landscapes and subject to spatiotemporal variability in habitat suitability. Several decades of theory and model development, as well as increased computing power and availability of fine-resolution GIS data, now make such models possible. Still unanswered, however, is the question of how well this new generation of dynamic models will anticipate range expansion. Here we develop a spatially explicit stochastic model that combines dynamic dispersal and population processes with fine-resolution maps characterizing spatiotemporal heterogeneity in climate and habitat to model range expansion of the hemlock woolly adelgid (HWA; Adelges tsugae). We parameterize this model using multiyear data sets describing population and dispersal dynamics of HWA and apply it to eastern North America over a 57-year period (1951-2008). To evaluate the model, the observed pattern of spread of HWA during this same period was compared to model predictions. Our model predicts considerable heterogeneity in the risk of HWA invasion across space and through time, and it suggests that spatiotemporal variation in winter temperature, rather than hemlock abundance, exerts a primary control on the spread of HWA. Although the simulations generally matched the observed current extent of the invasion of HWA and patterns of anisotropic spread, it did not correctly predict when HWA was observed to arrive in different geographic regions. We attribute differences between the modeled and observed dynamics to an inability to capture the timing and direction of long-distance dispersal events that substantially affected the ensuing pattern of spread.     

Decomposing the mid-Holocene Tsuga decline in eastern North America

The mid-Holocene decline of Tsuga canadensis (hereafter Tsuga) populations across eastern North America is widely perceived as a synchronous event, driven by pests/pathogens, rapid climate change, or both. Pattern identification and causal attribution are hampered by low stratigraphic density of pollen-sampling and radiometric dates at most sites, and by absence of highly resolved, paired pollen and paleoclimate records from single sediment cores, where chronological order of climatic and vegetational changes can be assessed. We present an intensely sampled (contiguous 1-cm intervals) record of pollen and water table depth (inferred from testate amoebae) from a single core spanning the Tsuga decline at Irwin Smith Bog in Lower Michigan, with high-precision chronology. We also present an intensively sampled pollen record from Tower Lake in Upper Michigan. Both sites show high-magnitude fluctuations in Tsuga pollen percentages during the pre-decline maximum. The terminal decline is dated at both sites ca. 5000 cal yr BP, some 400 years later than estimates from other sites and data compilations. The terminal Tsuga decline was evidently heterochronous across its range. A transient decline ca. 5350 cal yr BP at both sites may correspond to the terminal decline at other sites in eastern North America. At Irwin Smith Bog, the terminal Tsuga decline preceded an abrupt and persistent decline in water table depths by approximately 200 years, suggesting the decline was not directly driven by abrupt climate change. The Tsuga decline may best be viewed as comprising at least three phases: a long-duration pre-decline maximum with high-magnitude and high-frequency fluctuations, followed by a terminal decline at individual sites, followed in turn by two millennia of persistently low Tsuga populations. These phases may not be causally linked, and may represent dynamics taking place at multiple temporal and spatial scales. Further progress toward understanding the phenomenon requires an expanded network of high-resolution pollen and paleoclimate chronologies.     

Nonadditive effects of leaf litter species diversity on breakdown dynamics in a detritus-based stream

Since species loss is predicted to be nonrandom, it is important to understand the manner in which those species that we anticipate losing interact with other species to affect ecosystem function. We tested whether litter species diversity, measured as richness and composition, affects breakdown dynamics in a detritus-based stream. Using full-factorial analyses of single- and mixed-species leaf packs (15 possible combinations of four dominant litter species; red maple [Acer rubrum], tulip poplar [Liriodendron tulipifera], chestnut oak [Quercus prinus], and rhododendron [Rhododendron maximum]), we tested for single-species presence/absence (additive) or species interaction (nonadditive) effects on leaf pack breakdown rates, changes in litter chemistry, and microbial and macroinvertebrate biomass. Overall, we found significant nonadditive effects of litter species diversity on leaf pack breakdown rates, which were explained both by richness and composition. Leaf packs containing higher litter species richness had faster breakdown rates, and antagonistic effects of litter species composition were observed when any two or three of the four litter species were mixed. Less-consistent results were obtained with respect to changes in litter chemistry and microbial and macroinvertebrate biomass. Our results suggest that loss of litter species diversity will decrease species interactions involved in regulating ecosystem function. To that end, loss of species such as eastern hemlock (Tsuga canadensis) accompanied by predicted changes in riparian tree species composition in the southeastern United States could have nonadditive effects on litter breakdown at the landscape scale.     

Effects of Land‐Use Change on Community Composition of Tropical Amphibians and Reptiles in Sulawesi,Indonesia

Abstract: Little is known about the effects of anthropogenic land‐use change on the amphibians and reptiles of the biodiverse tropical forests of Southeast Asia. We studied a land‐use modification gradient stretching from primary forest, secondary forest, natural‐shade cacao agroforest, planted‐shade cacao agroforest to open areas in central Sulawesi, Indonesia. We determined species richness, abundance, turnover, and community composition in all habitat types and related these to environmental correlates, such as canopy heterogeneity and thickness of leaf litter. Amphibian species richness decreased systematically along the land‐use modification gradient, but reptile richness and abundance peaked in natural‐shade cacao agroforests. Species richness and abundance patterns across the disturbance gradient were best explained by canopy cover and leaf‐litter thickness in amphibians and by canopy heterogeneity and cover in reptiles. Amphibians were more severely affected by forest disturbance in Sulawesi than reptiles. Heterogeneous canopy cover and thick leaf litter should be maintained in cacao plantations to facilitate the conservation value for both groups. For long‐term and sustainable use of plantations, pruned shade trees should be permanently kept to allow rejuvenation of cacao and, thus, to prevent repeated forest encroachment.     

Effects of Conversion of Dry Tropical Forest to Agricultural Mosaic on Herpetofaunal Assemblages

Abstract: We explored the impact of forest conversion to agricultural mosaic on anuran, lizard, snake, and turtle assemblages of Neotropical dry forests. Over 2 years, we sampled 6 small watersheds on the west coast of Mexico, 3 conserved and 3 disturbed. The disturbed watersheds were characterized by a mosaic of pastures and cultivated fields (corn, beans, squash) intermingled with patches of different successional stages of dry forest. In each watershed, we conducted 11 diurnal and nocturnal time‐constrained searches in 10 randomly established plots. We considered vulnerability traits of species in relation to habitat modification. Eighteen anuran, 18 lizard, 23 snake, and 3 turtle species were recorded. Thirty‐six species (58%) occurred in both forest conditions, and 14 (22%) and 12 species (19%) occurred only in the conserved and disturbed sites, respectively. Assemblages responded differently to disturbance. Species richness, diversity, and abundance of lizards were higher in disturbed forests. Anuran diversity and species richness were lower in disturbed forest but abundance was similar in both forest conditions. Diversity, richness, and abundance of turtles were lower in disturbed forest. The structure and composition of snake assemblages did not differ between forest conditions. We considered species disturbance sensitive if their abundance was significantly less in disturbed areas. Four anuran (22%), 2 lizard (11%), and 3 turtle (100%) species were sensitive to disturbance. No snake species was sensitive. The decline in abundance of disturbance‐sensitive species was associated with the reduction of forest canopy cover, woody stem cover, roots, and litter‐layer ground cover. Anuran species with small body size and direct embryonic development were especially sensitive to forest disturbance. An important goal for the conservation of herpetofauna should be the determination of species traits associated with extinction or persistence in agricultural mosaics.     

Effects of Selective Logging on the Butterflies of a Bornean Rainforest

Abstract: Selective logging has been the main cause of disturbance to tropical forests in Southeast Asia, so the extent to which biodiversity is maintained in selectively logged forest is of prime conservation importance. We compared the butterfly assemblages of Bornean primary rainforest to those of rainforest selectively logged 6 years previously. We sampled by means of replicated transects stratified into riverine and ridge forests and we included roads in the logged forest. There was a three-fold variation in species richness and abundance over the 8-month sampling period. More species and individuals were observed in the logged forest, although between-replicate variability was high. Rarefied species richness was positively correlated with canopy openness within the range of disturbance levels encountered at our forest sites. Within families, there was no significant difference in the number of species between primary and logged forest. There was a significant difference in the relative abundance of species, but this was due largely to the abundance of one or two species. Community ordination separated the sites along a gradient of disturbance and revealed strong differences between riverine and ridge-forest butterfly assemblages in primary forest that were obscured in logged forest. There was no evidence that logging has resulted in a change in the composition of the butterfly assemblages from species with a local distribution to more widespread species. We conclude that at a logged forest site in close proximity to primary forest, low intensities of logging do not necessarily reduce the species richness or abundance of butterflies, although assemblage composition is changed.     

Riparian forest composition affects stream litter decomposition despite similar microbial and invertebrate communities

Cross-boundary flows of energy and nutrients link biodiversity and functioning in adjacent ecosystems. The composition of forest tree species can affect the structure and functioning of stream ecosystems due to physical and chemical attributes, as well as changes in terrestrial resource subsidies. We examined how variation in riparian canopy composition (coniferous, deciduous, mixed) affects adjacent trophic levels (invertebrate and microbial consumers) and decomposition of organic matter in small, coastal rainforest streams in southwestern British Columbia. Breakdown rates of higher-quality red alder (Alnus rubra) litter were faster in streams with a greater percentage of deciduous than coniferous riparian canopy, whereas breakdown rates of lower-quality western hemlock (Tsuga heterophylla) litter were independent of riparian forest composition. When invertebrates were excluded using fine mesh, breakdown rates of both litter species were an order of magnitude less and were not significantly affected by riparian forest composition. Stream invertebrate and microbial communities were similar among riparian forest composition, with most variation attributed to leaf litter species. Invertebrate taxa richness and shredder biomass were higher in A. rubra litter; however, taxa evenness was greatest for T. heterophylla litter and both litter species in coniferous streams. Microbial community diversity (determined from terminal restriction fragment length polymorphisms) was unaffected by riparian forest or litter species. Fungal allele richness was higher than bacterial allele richness, and microbial communities associated with lower-quality T. heterophylla litter had higher diversity (allele uniqueness and richness) than those associated with higher-quality A. rubra litter. Percent variation in breakdown rates was mostly attributed to riparian forest composition in the presence of invertebrates and microbes; however, stream consumer biodiversity at adjacent trophic levels did not explain these patterns. Riparian and stream ecosystems and their biotic communities are linked through exchange and decomposition of detrital resources, and we provide evidence that riparian forest composition affects stream ecosystem catabolism despite similarities in microbial and invertebrate communities.     

The influence of successional processes and disturbance on the structure of Tsuga canadensis forests

Old-growth forests are valuable sources of ecological, conservation, and management information, yet these ecosystems have received little study in New England, due in large part to their regional scarcity. To increase our understanding of the structures and processes common in these rare forests, we studied the abundance of downed coarse woody debris (CWD) and snags and live-tree size-class distributions in 16 old-growth hemlock forests in western Massachusetts. Old-growth stands were compared with eight adjacent second-growth hemlock forests to gain a better understanding of the structural differences between these two classes of forests resulting from contrasting histories. In addition, we used stand-level dendroecological reconstructions to investigate the linkages between disturbance history and old-growth forest structure using an information-theoretic model selection framework. Old-growth stands exhibit a much higher degree of structural complexity than second-growth forests. In particular, old-growth stands had larger overstory trees and greater volumes of downed coarse woody debris (135.2 vs. 33.2 m3/ha) and snags (21.2 vs. 10.7 m3/ha). Second-growth stands were characterized by either skewed unimodal or reverse-J shaped diameter distributions, while old-growth forests contained bell-shaped, skewed unimodal, rotated sigmoid, and reverse J-shaped distributions. The variation in structural attributes among old-growth stands, particularly the abundance of downed CWD, was closely related to disturbance history. In particular, old-growth stands experiencing moderate levels of canopy disturbance during the last century (1930s and 1980s) had greater accumulations of CWD, highlighting the importance of gap-scale disturbances in shaping the long-term development and structural characteristics of old-growth forests. These findings are important for the development of natural disturbance-based silvicultural systems that may be used to restore important forest characteristics lacking in New England second-growth stands by integrating structural legacies of disturbance (e.g., downed CWD) and resultant tree-size distribution patterns. This silvicultural approach would emulate the often episodic nature of CWD recruitment within old-growth forests.