Effects of Pioneer Tree Species Hyperabundance on Forest Fragments in Northeastern Brazil

Abstract: Despite many studies on fragmentation of tropical forests, the extent to which plant and animal communities are altered in small, isolated forest fragments remains obscure if not controversial. We examined the hypothesis that fragmentation alters the relative abundance of tree species with different vegetative and reproductive traits. In a fragmented landscape (670 km²) of the Atlantic Forest of northeastern Brazil, we categorized 4056 trees of 182 species by leafing pattern, reproductive phenology, and morphology of seeds and fruit. We calculated relative abundance of traits in 50 1‐ha plots in three types of forest configurations: forest edges, small forest fragments (3.4–83.6 ha), and interior of the largest forest fragment (3500 ha, old growth). Although evergreen species were the most abundant across all configurations, forest edges and small fragments had more deciduous and semideciduous species than interior forest. Edges lacked supra‐annual flowering and fruiting species and had more species and stems with drupes and small seeds than small forest fragments and forest interior areas. In an ordination of species similarity and life‐history traits, the three types of configurations formed clearly segregated clusters. Furthermore, the differences in the taxonomic and functional (i.e., trait‐based) composition of tree assemblages we documented were driven primarily by the higher abundance of pioneer species in the forest edge and small forest fragments. Our work provides strong evidence that long‐term transitions in phenology and seed and fruit morphology of tree functional groups are occurring in fragmented tropical forests. Our results also suggest that edge‐induced shifts in tree assemblages of tropical forests can be larger than previously documented.     

Bryophyte responses to microclimatic edge effects across riparian buffers.

Although riparian buffers are an important aspect of forest management in the boreal forest of Canada, little is known about the habitat conditions within buffers, due in part to complex edge effects in response to both the upland clearcut and the stream. We investigated microclimatic conditions and bryophyte growth and vitality in seven locations between the stream edge and 60 m into the upland undisturbed conifer forests and at the clearcut sites with riparian buffer 30 km northwest of Thunder Bay, Ontario, Canada. We hypothesized that the growth and vitality of a pleurocarpous moss, Hylocomium splendens, and an acrocarpous moss, Polytrichum commune, would be directly related to the microclimatic gradients detected. We further hypothesized that sensitivity of the bryophytes to environmental factors will vary depending on their life form type, i.e., pleurocarpous moss will respond differently than the acrocarpous moss. Both bryophyte species were transplanted in pots and placed at 10-m intervals along 60-m transects perpendicular to the stream across the buffer and undisturbed sites. Bryophyte growth, cover, and vitality, as well as microclimatic parameters and plant cover, were measured over the summer in 2003. The riparian buffers were simultaneously affected by microclimatic gradients extending from both the clearcut edge and the riparian-upland ecotonal edge. Both bryophyte species responded to changes in the microclimatic conditions. However, vapor pressure deficit (VPD) was the most important factor influencing the growth of H. splendens, whereas for P. commune growth soil moisture was most important. Our study confirms earlier findings that interior forest bryophytes such as H. splendens can be used as indicators to monitor edge effects and biodiversity recovery following forest harvesting. We demonstrate that growth and vitality of these bryophytes reflect the prevailing near-ground microclimatic conditions at the forest edges. Abundance estimates of such bryophytes can be used to determine the depth of edge effects across both ecotonal edges (e.g., riparian-upland forest edge) and anthropogenically created edges (e.g., clearcut edge). Forest management practices must consider depth of edge in determining the appropriate width of riparian buffers that would be necessary to sustain biodiversity and associated values at the land/water interface.     

Response of Red-Backed Voles to Recent Patch Cutting in Subalpine Forest

We examined the response of southern red-backed voles ( Clethrionomys gapperi ) to patch cutting in a forested landscape in which 23% of the forest cover had been removed by timber harvest. We live trapped voles in and around 18 patchcuts in one watershed of southern Wyoming. Although we found a significant difference in capture rates between patchcut interior and forest habitats in 1 of 2 years, voles did not strongly avoid the interior of patchcuts. This result contrasts with results from most studies of voles in clearcut ecosystems, which report that red-backed voles are generally rare or absent from clearcuts. Capture rates were highest on both sides of the patchcut edge, which also contrasts with studies of voles at the edges of forest remnants. The use of patchcut interior and edge habitats could not be explained as a consequence of juvenile voles dispersing to those habitats or males moving through the habitat in search of mates. We suggest that, despite similar physiognomy in patchcut and clearcut sites, the differences in landscape structure in perforated versus fragmented landscapes lead to very different patterns of vole movement. Understanding the scales at which voles perceive landscapes as coarse- or fine-grained will be key to developing predictive models to aid managers in designing timber sales that maintain high vole populations. Our results emphasize the importance of the spatial pattern and scale of disturbance in determining the response of vertebrates to landscape change and the need for more refined investigations of the consequences of deforestation.     

Vegetation Responses along Edge-to-Interior Gradients in the Mixed Hardwood Forests of the Roanoke River Basin, North Carolina

Compared with forest interiors, forest edges typically have a different plant species composition and community structure, a phenomenon known as "edge effect." Edge effects make the functional interior area of a forest smaller than its actual area. The objective of this study was to estimate how far the effects of agriculturally maintained edges penetrate the mixed hardwood forests of the Roanoke River Basin, North Carolina. I determined percentage cover for all vascular plant species in 10-by-100-meter belt transects on north-facing or south-facing edges of four relatively undisturbed forests. Changes in the percentage cover of individual species, the relative cover of exotic species, and species richness all indicated that edge effects penetrate deeper on south-facing edges (to 60 meters) than on north-facing edges (to 20 meters). Analyses of species responses to the edge showed a number of species to be edge oriented, but no species was found to be interior oriented. The results of multivariate analyses (ordination and cluster analysis) suggested that edge effects could be detected to 50 meters on south-facing edges and 10–30 meters on north-facing edges. These results allow us to better understand the difference between a forest's actual area and its functional interior area.     

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.     

Edge Effects and Isolation: Red-Backed Voles on Forest Remnants

Negative effects of habitat edge have been advanced as an important proximate cause of extinction, and a growing literature calls attention to the matrix surrounding habitat remnants as a critical factor determining population persistence. I examined spatial distribution of California red-backed voles ( Clethrionomys californicus ) on 13 forest remnants and five control sites in southwestern Oregon. The species was virtually isolated on remnants, making little use of the regenerating clearcuts surrounding the remnants. The effects of the clearcut also impinged on the remnants as edge effects: six times more voles were captured per trap in the interior of remnants than on the edge. Consequently, the density of voles per unit area on remnants increased with remnant size, despite the potential buildup of population density in small isolates due to limited emigration. I explored potential mechanisms of the negative edge effect on voles and found that the biomass of coarse woody debris, per se, did not explain the vole distribution because both number and volume of logs increased from the interior to the edge of remnants. However, the distribution of the vole's primary food item, hypogeous sporocarps of mycorrhizal fungi, did correspond to the vole edge effect     

Edge Effects on Lizards and Frogs in Tropical Forest Fragments

Abstract: We investigated whether forest-pasture edges affect the distribution of an assemblage of small vertebrate ectotherms in a consistent and predictable manner. We describe the abundance and distribution of two species of anoline lizards (   Norops ) and five species of leaf-litter frogs (   Eleutherodactylus ) along the edges and in the interiors of nine forest fragments near Las Cruces, Costa Rica. Over 4 months, we surveyed 44 pairs of plots by visual encounter. In each pair of plots, one was immediately adjacent to the pasture and the second was within the forest "interior." Both plots of a pair were searched simultaneously. This block design controlled for the effects of weather, topography, and searcher ability. The distribution of all species was highly variable with respect to edges. Only two species of frogs, Eleutherodactylus podiciferus and E. cruentus , were significantly more abundant in interior plots than in edge plots, although not consistently so. Both species of Norops lizards were more abundant along forest edges during the dry season. Both Norops species and several Eleutherodactylus species, however, appeared to become more abundant in the forest interior after the onset of the wet season, suggesting a seasonal edge effect. In Norops polylepis , the most abundant anole, rates of ectoparasitism were lower along edges than in forest interiors. The magnitude of the edge effect on any one species was not influenced by the size of fragments or by the distance of the interior plot from the nearest edge. We believe that edge effects should not be defined by the distance to which they are detected. Rather, they should be viewed as highly dynamic in space and time; taxa appear to respond to different components of edge effects according to their particular biological requirements.     

Are shrubland birds edge specialists?

In studies of forest fragmentation, birds of scrubby, early-successional habitats are considered edge specialists. Because these birds are assumed to thrive in fragmented, edge-dominated areas, their landscape ecology has received little attention from ecologists. With populations of shrubland birds declining throughout the eastern United States, the question of whether or not these birds really prefer edge habitats has important conservation implications. We used a meta-analysis to test how edges affect the abundance of shrubland birds in early-successional habitats. We analyzed data for 17 species from seven studies that compared the abundances of birds in the interiors and edges of regenerating clearcuts surrounded by mature forest. The meta-analysis clearly showed that shrubland birds avoid edges. All 17 species tested had higher abundances in patch centers than along edges, and edge effects were significant for 8 of 17 species. The key implication of this result is that small or irregular patches, dominated by edge, are unlikely to provide suitable habitat for shrubland birds. Thus, management for these declining species should involve providing large patches and minimizing edges. These findings demonstrate the importance of testing widely accepted ecological classifications and the need to view landscape ecology from the perspective of non-forest wildlife.     

Nesting Success of a Neotropical Migrant in a Multiple-Use, Forested Landscape

Abstract: We studied the nesting success of an individually marked population of Kentucky Warblers ( Oporornis formosus ), a species that nests in disturbed and undisturbed forests, in a heterogeneous, managed forest site in the Shawnee National Forest in southern Illinois from 1992 to 1995. We examined the effects of forest stand type (clearcuts of various ages, tree plantations, and older forest) and distance from habitat edges on rates of nest predation and brood parasitism by Brown-headed Cowbirds (   Molothrus ater ). Brood parasitism levels gradually decreased from 60% to 3% ( nests) over a distance of 2 km from an agricultural edge proximal to a known cowbird foraging site (a pig feedlot), but they did not vary with distance from any other kinds of edges or with forest stand type. Rates of nest predation ( nests) did not vary with distance from any edges, but they were significantly lower in older forest than within even-aged clearcuts, a tree plantation, and in successional vegetation adjacent to a residential facility. These results suggest that, even in fragmented landscapes with high overall levels of parasitism and nest predation, management practices within and immediately adjacent to forest tracts can affect the nesting success of some species, but not necessarily as a simple function of distance from edge. For the Kentucky Warbler, our results suggest that a management strategy that avoids even-age silviculture and leaves core stands of older forest far from cowbird feeding areas can increase nesting success to levels similar to those measured in more forested landscapes.     

Effects of Clearcutting on Habitat Use and Reproductive Success of the Ovenbird in Forested Landscapes

We studied Ovenbirds ( Seiurus aurocapillus ) in northern New Hampshire during 1992 and 1993 to determine whether edge-related changes in habitat use and reproductive success reported in fragmented landscapes exist in predominantly forested landscapes. Six study plots were placed adjacent to four recent clearcuts (2.1–5 ha) and extended 400 m into the forest interior. Nests, territories, and territorial males obtaining mates were equally distributed in edge (0–200 m) and interior (201–400 m) areas. Nest survival was higher in the forest interior in 1992 and for 1992 and 1993 combined. The proportion of pairs fledging ≥ 1 young, fledgling weight, and fledgling wing-chord did not differ between edge and interior in either year. Number of young fledged per pair was slightly lower in edge areas, but these differences were not significant. We conclude that clearcutting in extensively forested landscapes can affect Ovenbird reproductive success. Nevertheless, the effect on Ovenbird populations is moderated by the abundance of mature forest cover in the region and by the tendency of Ovenbirds to renest after initial nest failure.     

Effect of Edge Structure on the Flux of Species into Forest Interiors

Abstract: A key goal of conservation biology is to prevent the spread of exotic species. Previous work on exotic invasion has two limitations: (1) the lack of a spatially explicit approach and (2) a primary focus on the net effect of invasion by examining invasive species already present in the community. We address these limitations by focusing on the arrival of a potential invader into a community and use a spatially explicit approach to quantify the flow of seeds from the surrounding landscape into the interior of a forest. We hypothesize that the structure of forest-edge vegetation influences how the edge mediates seed flux. To test our hypothesis, we experimentally altered vegetation structure within 20 m of the edge to create two edge treatments: thinned and intact. We quantified the flux of seeds moving into the forest interior across the two treatments. We used seed traps randomly arrayed on transects from 5 to 50 m into the forest. More seeds crossed the thinned treatment than crossed the intact treatment to reach the forest interior. In addition, seeds that crossed the thinned treatment dispersed farther into the forest than those that crossed the intact treatment. These results were consistent throughout the period of maximum autumn dispersal, including periods before and after leaf drop. Our results show that the structure of vegetation on the edge interacts with the flux of wind-dispersed seed across the edge. We demonstrated that an edge with intact vegetation can function as a physical barrier to seed dispersal. Therefore, the structure of vegetation on edges can influence the function of edges as barriers to seed flux into the forest interior.     

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.     

Edge Effects on the Understory Bird Community in a Logged Forest in Uganda

Abstract: Understanding how the fauna of logged tropical rainforests responds to fragmentation and the creation of edges is vital to ensure conservation of biodiversity. We studied the composition of the understory bird community from the edge of a 15-ha clearing toward the interior of the forest in a part of Budongo Forest Reserve, Uganda, that was selectively logged about 45 years ago. Mist netting was conducted along five transects from the edge and 500 m into the interior. The total number of individuals captured did not change with distance from the edge, but there was a significant increase in the number of species. We sampled fewer, but more common species near the edge, whereas the interior of the forest had more, and less common species. Guild composition also changed with distance from the edge. Frugivore-insectivores and nectarivores were most common close to the edge. Among insectivores, ground foragers, bark-gleaners, and leaf-gleaners were most common in the interior of the forest, whereas sallying insectivores favored the edge. Graminivores were unaffected by the edge. Analysis of common species showed that Ispidina picta , Andropadus curvirostris , A. latirostris , Camaroptera brachyura , Terpsiphone rufiventer , and Nectarinia olivacea were associated with the edge, but no species showed significant avoidance of the edge. This finding may be explained by the generally low sample sizes of interior species. Our results show that even bird communities in logged forests respond to edges. Estimates of edge effects suggested that changes in bird densities may have occurred several hundred meters from the edge. In conclusion, logged forests provide habitat for bird species avoiding forest edges, and this should be considered in the management of such forests for conservation.     

Disappearance and Return of Plethodontid Salamanders to Clearcut Plots in the Southern Blue Ridge Mountains

The disappearance and return of plethodontid salamanders on clearcuts has been monitored since 1979 in the southern Blue Ridge Mountains at three sites near Highlands, North Carolina. Salamander abundance on 225 m² plots located on clearcuts and in nearby forest was determined by nightly non-destructive searches. Abundances on clearcut and forest plots at a given site were compared for each year in which sampling occurred. Numbers of salamanders on clearcut plots decreased to 30–50% of forested plots in the first year after logging and were almost zero by the second year. Decreases in standing crop and moisture content of leaf litter seem responsible for salamander disappearance. Salamanders returned to clearcuts 4–6 years after cutting, and their numbers increased rapidly. Linear regressions estimate that salamander numbers on clearcut plots will equal or exceed numbers on forested plots by 20–24 years after cutting. The pattern of salamander return to clearcuts appears closely correlated with the timing of litter layer reformation. All sex and age classes of the most common species, Plethodon jordani, disappear from clearcuts at equal rates, whereas the earliest colonizers are predominantly large adults. Plethodon oconaluftee, a desiccation-resistant species, exists on regenerating clearcuts in disproportionately large numbers. Large adults of all species, including Plethodon oconaluftee, may be better able to withstand the drier, sparse litter cover of young, regenerating stands. Adults might move to clearcuts to avoid competition from smaller and immature salamanders restricted to mature forests with abundant, moist litter.     

Forest Edges as Nutrient and Pollutant Concentrators: Potential Synergisms between Fragmentation, Forest Canopies, and the Atmosphere

Abstract: Forest fragmentation leads to a dramatic increase in forest edge, and these edges may function as traps and concentrators for wind-borne nutrients and pollutants. We assessed the influence of forest edges on atmospheric deposition and subsequent inputs to the forest floor in deciduous-forest fragments in the eastern United States. To quantify these inputs, we collected throughfall—water that has passed through the forest canopy—from edge and interior zones of forests adjacent to open fields. During the 1995 growing season, atmospheric input (wet and dry deposition) of sulfur to forest edge zones was elevated compared with input to forest interiors. Throughfall fluxes of dissolved inorganic nitrogen and calcium were also greater at edges than interiors. The mean edge increases ranged from 17% to 56% for the nutrients and pollutants we measured. When we manipulated the structure of forest edges by removing all vegetation below half the canopy height, throughfall flux in the edge zone declined sharply and was less than that of the respective interior zone. Changing the vegetation structure of the edge also shifted the zone of highest throughfall flux farther into the interior of the forest. Our data suggest that forest edges can function both as significant traps for airborne nutrients and pollutants from adjoining agricultural or urban landscapes and effective concentrators of below-canopy chemical fluxes. These enhanced fluxes may have cascading effects on soil-nutrient cycling, microbial activity, seedling dominance, and other ecological processes near forest edges.     

Effects of Forest Management on Amphibians and Reptiles in Missouri Ozark Forests

Abstract:  As part of the Missouri Ozark Forest Ecosystem Project (MOFEP), we experimentally evaluated the impacts of forest management on the relative abundance of amphibians and reptiles in Missouri's Ozark forests (U.S.A.). Using large study sites (average size of 400 ha) as the experimental unit, we tested the effects of uneven-aged and even-aged forest management treatments compared with no-harvest management (i.e., control) on the relative abundance of 13 focal amphibian and reptile species. Within even-aged management sites, we also focused on the local-scale effects of clearcutting on these species by comparing relative abundance among plots located within clearcut stands, 50 m away from clearcut stands, and 200 m away from clearcut stands. Pretreatment sampling of species abundance occurred from 1992 through 1995, and post-treatment sampling occurred from 1997 through 2000. At the landscape scale, treatment significantly affected the abundance of Bufo americanus . This species declined less on even-aged management sites than on control sites, but the general decline on all sites suggests that other factors may have contributed to this result. Within even-aged management sites, most amphibian species declined and some reptile species increased relative to pretreatment abundances within clearcut stands. We found significant effects of distance from clearcut for two amphibian species, Ambystoma maculatum and Rana clamitans, and two reptile species, Scincella lateralis and Sceloporus undulatus . In general, we conclude that clearcuts within even-aged management sites locally affected amphibian and reptile species but, at a larger spatial scale, we did not detect significant effects of even-aged and uneven-aged forest management. These findings represent relatively short-term data but suggest that forest management and maintenance of biodiversity may be compatible when relatively small amounts of the landscape are disturbed.     

Edge Effects and Isolation: Red-Backed Voles Revisited

Abstract:  We examined demographic responses of California red-backed voles ( Clethrionomys californicus ) to forest fragmentation in southwestern Oregon at sites where this species has previously shown negative responses to fragmentation. Voles were captured in live traps and released. Voles were rarely caught in clearcuts surrounding 11 forest fragments, but relative vole density did not decrease from the forest-fragment interiors to edges. The first result agrees with previous findings at these sites 6 years earlier, but the latter result does not. There was no evidence that vole response to edge changes with fragment age. Two years of intensive mark-recapture efforts at two forest-fragment sites and two unfragmented (control) sites did not show negative effects of fragmentation on vole survival, an important demographic rate. Vole capture probabilities varied greatly across space and time on these four sites, which may explain the differences in vole responses to edge seen between this and the previous study. These results suggest that reliable appraisal of edge effects may be difficult for many species on small fragments because the data necessary to apply population estimators require great efforts to obtain and the use of indices leads to a confounding of detection probabilities with demographic change.     

Invertebrates and Boreal Forest Management

Logging has ecological effects on invertebrates in Fennoscandian boreal forest. Especially affected are species associated with micro-habitats of natural old-growth forest, such as coarse woody debris, large deciduous trees, and patches of wet swamp-forest. Furthermore, the effective control of forest fires has adversely affected many species that require burned substrates. Although old-growth specialists tend to disappear from clear-cuts, local invertebrate species richness (α-diversity) may increase as forest generalists persist and numerous open-habitat species appear. This seems true at least for taxa (e.g., ground-beetles and spiders) that are not directly associated with the characteristic micro-habitats of old growth mentioned above. However, at the landscape or biogeographical scale intensive logging tends to homogenize forest habitats and lead to declines of sensitive species. Based on these findings three complementary approaches appear useful for the maintenance of boreal biodiversity while harvesting timber. (1) Undisturbed old-growth forests must be set aside to sustain specialist species and to serve as sources for recolonization. (2) Because these reserves are not, and will probably not be, large and regionally representative enough, ecologically sound silvicultural practices must be developed using natural disturbance regimes as guidelines. (3) Restoration of habitats and the process of natural regeneration through fire is imperative for the recovery of associated species assemblages. However, the process of recovery presupposes that colonists are available, which may not always be the case.     

Contribution of Roads to Forest Fragmentation in the Rocky Mountains

The contribution of roads to forest fragmentation has not been adequately analyzed. We quantified fragmentation due to roads in a 30,213-ha section of the Medicine Bow-Routt National Forest in sout heastern Wyoming with several indices of landscape structure using a geographic information system. The number of patches, mean patch area, mean interior area, mean area of edge influence, mean patch perimeter, total perimeter, and mean patch shape identified patch- and edge-related landscape changes. Shannon-Wiener diversity, dominance, contagion, contrast, and angular second moment indicated effects on landscape diversity and texture. Roads added to forest fragmentation more than clearcuts by dissecting large patches into smaller pieces and by converting forest interior habitat into edge habitat. Edge habitat created by roads was 1.54–1.98 times the edge habitat created by clearcuts. The total landscape area affected by clearcuts and roads was 2.5–3.5 times the actual area occupied by these disturbances. Fragmentation due to roads could be minimized if road construction is minimized or rerouted so that its fragmentation effects are reduced. Geographic information system technology can be used to quantify the potential fragmentation effects of individual roads and the cumulative effects of a road network on landscape structure.     

Frog, Bat, and Dung Beetle Diversity in the Cloud Forest and Coffee Agroecosystems of Veracruz, Mexico

Abstract:  We compared the species diversity of copronecrophagous beetles (Scarabaeinae) , bats, and frogs in tropical montane cloud forest (original vegetation) and shaded coffee plantations (an agroecosystem common to the region) for a landscape in central Veracruz, Mexico. We sampled in three tropical montane cloud forest fragments and in three coffee plantations with traditional polyculture shade between 1998 and 2001. The three indicator groups responded differently to the transformation of tropical montane cloud forest into shaded coffee plantations. The species richness of frogs was one-fifth less in coffee plantations than in forest fragments, and only one-third of the frog species occurred in both forest fragments and coffee plantations. The number of beetle species and their abundance was significantly greater in coffee plantations than in the forest fragments, whereas species richness and species composition of bats were virtually the same in both habitats. The majority of the abundant species remained as such in both communities, but species that were less abundant were not scarce in both habitats. We attributed differences in the species assemblages to the differing degrees of penetrability of the borders of the two habitat types (especially for the coffee plantations) and to the differences in life-history traits among species. Shaded coffee plantations form a matrix that envelops the remaining fragments of cloud forest. Together they connect the forest fragments with the other habitats of the landscape and represent a highly functional resource for the preservation of biodiversity that serves as a complement to but not a substitute for cloud forest in this notably modified landscape.