Effects of Habitat Fragmentation on Bird Species in a Relict Temperate Forest in Semiarid Chile

Abstract: We analyzed the structure and composition of a bird assemblage in a fragmented relict temperate forest located in northcentral Chile ( Fray Jorge National Park). In terms of species composition, the bird assemblage we found in Fray Jorge was more similar to southern temperate forest sites, located more than 1200 km south of Fray Jorge, than to localities found in nearby scrub habitats. The relict character and long-term isolation of the Fray Jorge forest provides a natural experiment with which to establish the potential long-term effects of fragmentation and isolation on southern Chilean temperate forests. Between May 1996 and March 1997, we conducted seasonal surveys of birds in six forest fragments, ranging in size from 0.5 to 22.5 ha, at Fray Jorge. The number of bird species at each forest fragment was positively correlated with fragment area during all seasons. The relict forest system had a steeper species-area slope than that reported for similar temperate-forest bird assemblages in forest fragments within Chiloé Island and for islands across the Chiloé Archipelago in southern Chile. In this regard, this bird fauna resembled a depauperate oceanic archipelago. This difference in area effects is likely a consequence of the minimization of rescue effects because of the absence of large source forest areas nearby and the long-term isolation of the system. In addition, the distribution of species among forest fragments in Fray Jorge was not random, showing a nested subset pattern. Thus, some species occur across all fragments, regardless of their area, and therefore are less affected by habitat fragmentation and less prone to local extinction. These results suggest that, for south-temperate forest birds, large fragments (or reserves) should afford better protection against extinction than small forest patches.     

Effects of Residential Development on Forest-Dwelling Neotropical Migrant Songbirds

Many bird species are in local or regional decline because of habitat loss or degradation. We attempted to disentangle the effects of residential development from the effects of forest size on forest-bird communities, with particular emphasis on Neotropical migrant species. Two variables were examined for their influence on avian diversity and abundance: forest size and the number of houses outside a forest within 100 m of the edge. We found that Neotropical migrants consistently increased in number and abundance as forest size increased. Of greater interest, we found that the number of houses surrounding a forest severely undermined its suitability for Neotropical migrants. Neotropical migrants consistently decreased in diversity and abundance as the level of adjacent development increased, regardless of forest size. The effects of development were striking: 4-ha woodlots without any nearby houses had on average a richer, more abundant Neotropical community than did 25-ha urban woodlots. No predictable pattern of change concerning development or forest size was observed for short-distance migrants or permanent residents. Current planning regulations generally permit housing right up to forest edges. This practice may prevent protection of ecological features within the forest. Threshold distances for housing developments around forests need to be determined to prevent or minimize adverse effects on features and functions within the forests.     

Nested Faunas and Extinction in Fragmented Habitats

Abstract: The faunas of habitat islands, such as those produced by fragmentation of formerly continuous habitats, are commonly made up of nonrandom subsets of the total available species pool. Faunas within an archipelago may form a nested series, with depauperate faunas made up of subsets of more species-rich faunas. The pattern is seldom perfect: widespread species may be absent from otherwise rich faunas (holes), and uncommon species may occur in depauperate faunas (outliers). The "nestedness" of an assemblage of faunas (its fit to the "nested subset model"; Patterson & Atmar 1986) can be measured by summing the holes and outliers.
The distributions of boreal mammals and birds among mountain ranges in the Great Basin of western North America were analyzed by this method Despite differences in their derivation (mammalian faunas are thought to be relicts, isolated since the Pleistocene; bird faunas have probably experienced recurrent colonization throughout their history] both groups show an approximately equal (and highly significant) fit to the nested subset model. They differ, however, in the relative numbers of holes and outliers in their patterns. The mammalian pattern is hole-rich; the bird pattern is outlier-rich.
The present-day composition of the mammalian faunas is the result of selective extinction of species of originally richer faunas. The existence of nested subsets in these faunas suggests that extinction is a highly deterministic process: extinctions occurred in approximately the same sequence throughout the region, despite wide variation in extinction rates Extinction sequence has, in fact, been less variable than extinction rates.     

Evaluating Effects of Habitat Loss and Land-Use Continuity on Ant Species Richness in Seminatural Grassland Remnants

Abstract:  Seminatural grasslands in Europe are susceptible to habitat destruction and fragmentation that result in negative effects on biodiversity because of increased isolation and area effects on extinction rate. However, even small habitat patches of seminatural grasslands might be of value for conservation and restoration of species richness in a landscape with a long history of management, which has been argued to lead to high species richness. We tested whether ant communities have been negatively affected by habitat loss and increased isolation of seminatural grasslands during the twentieth century. We examined species richness and community composition in seminatural grasslands of different size in a mosaic landscape in Central Sweden. Grasslands managed continuously over centuries harbored species-rich and ecologically diverse ant communities. Grassland remnant size had no effect on ant species richness. Small grassland remnants did not harbor a nested subset of the ant species of larger habitats. Community composition of ants was mainly affected by habitat conditions. Our results suggest that the abandonment of traditional land use and the encroachment of trees, rather than the effects of fragmentation, are important for species composition in seminatural grasslands. Our results highlight the importance of considering land-use continuity and dispersal ability of the focal organisms when examining the effects of habitat loss and fragmentation on biodiversity. Landscape history should be considered in conservation programs focusing on effects of land-use change.     

Inf luence of Within-Plantation Heterogeneity and Surrounding Landscape Composition on Avian Communities in Hybrid Poplar Plantations

We conducted breeding bird surveys in Minnesota, Wisconsin, and South Dakota in 12 hybrid poplar plantations and surrounding landscapes from 1992 to 1994. Plantations varied in age, shape, composition of surrounding landscape, and internal vegetative heterogeneity. Numbers of breeding bird individuals and species in plantations were lower than in surrounding forest/shrub habitat, but higher than in row crops. Numbers of individuals observed within several bird groups based on migratory status and habitat preference also differed among plantations and surrounding land-use types. Most differences were between numbers in plantations and row crops. Year-to-year changes in bird species composition in plantations were more likely in plantations between ages 2 and 4 years than in younger or older plantations. Correlative evidence from canonical correspondence analysis illustrated that plantation bird communities were related to habitat in surrounding landscapes, plantation age, size, latitude, and longitude. Additionally, more heterogeneous plantations had more species, individuals, and numbers of long-distance migrants. Plantations will likely not support bird communities that are comparable to natural forests in either species composition or species diversity. A goal would be to position them in the landscape to minimize impacts on regional biodiversity. This could be accomplished by maintaining structural diversity of plantations by creating a broad range of successional stages (ages) throughout plantations within a region. Size and connectivity of existing forest fragments may be increased by plantations, but fragmentation of natural open areas should be avoided.     

Mediation of area and edge effects in forest fragments by adjacent land use

Habitat loss, fragmentation, and degradation have pervasive detrimental effects on tropical forest biodiversity, but the role of the surrounding land use (i.e., matrix) in determining the severity of these impacts remains poorly understood. We surveyed bird species across an interior-edge-matrix gradient to assess the effects of matrix type on biodiversity at 49 different sites with varying levels of landscape fragmentation in the Brazilian Atlantic Forest—a highly threatened biodiversity hotspot. Both area and edge effects were more pronounced in forest patches bordering pasture matrix, whereas patches bordering Eucalyptus plantation maintained compositionally similar bird communities between the edge and the interior and exhibited reduced effects of patch size. These results suggest the type of matrix in which forest fragments are situated can explain a substantial amount of the widely reported variability in biodiversity responses to forest loss and fragmentation.     

Effects of Patch Size on Birds in Old-Growth Montane Forests

Following habitat alteration or fragmentation, competition, parasitism, and predation from species that live in the new habitats may reduce the survival and reproductive success of species living in the original habitats. Negative influences from species that live outside the remnant patches are expected to be greater in small rather than in large remnant patches because more "external" species are expected to move through the centers of small remnant patches. We surveyed birds within remnant patches of old-growth montane forests on Vancouver Island, Canada, (1) to evaluate whether the richness and abundance of non-old-growth bird species were larger at the center of small rather than large patches and (2) to evaluate whether the opposite was true of old-growth bird species. More non-old-growth bird species were present at the center of small remnant patches of old growth than in large old-growth patches. We found no relationship, however, between patch size and richness or abundance of old-growth bird species at the center of remnant patches of old growth. This was true for old-growth species with open, cup-shaped nests and cavity nests. Old-growth birds may have been affected less in our study area than in other areas because they evolved within heterogeneous montane forests and interacted with non-old-growth species throughout their evolutionary histories or because the contrast between old-growth forests and logged areas was less than that between the forests and agricultural/urban areas that were surveyed in other studies.     

Effects of Restoring Oak Savannas on Bird Communities and Populations

Abstract:  Efforts to restore and maintain oak savannas in North America, with emphasis on the use of prescribed fire, have become common. Little is known, however, about how restoration affects animal populations, especially those of birds. I compared the breeding densities, community structure, and reproductive success of birds in oak savannas maintained by prescribed fire (12 sites) with those in closed-canopy forests (13 sites). All sampling was conducted in Illinois (U.S.A.). Of the 31 bird species analyzed, 12 were more common in savannas, 14 were not affected by habitat structure, and 5 were more common in forest habitat. The species favored by disturbance and restoration included Northern Bobwhites ( Colinus virginianus ), Mourning Doves (  Zenaida macroura ), Red-headed Woodpeckers (  Melanerpes erythrocephalus ), Indigo Buntings (  Passerina cyanea ), and Baltimore Orioles ( Icterus galbula ). Those more common in closed-canopy forest included Ovenbirds ( Seiurus aurocapilla ) and Wood Thrushes (  Hylocichla mustelina ). Few species were unique to one type of habitat, but overall avian community structure in oak savannas and closed-canopy forests was generally distinctive. Estimates of nesting success (derived from 785 nests) revealed that 6 of the 13 species considered experienced greater productivity in the savanna habitat. Rates of brood parasitism were unaffected by restoration and habitat structure. Within savannas, tract size had little effect on breeding abundances and reproductive success. My results illustrate that restoration techniques can significantly affect the ecology of constituent animal populations and communities and have key implications regarding avian conservation and the management of forest habitat in fragmented landscapes. Small patches of forest habitat that regularly function as population sinks may offer far better prospects for birds if they are subjected to disturbance and ecosystem restoration.     

Effect of Slash-and-Burn Shifting Cultivation on Rainforest Birds in Mizoram, Northeast India

Abstract: Slash-and-burn shifting cultivation, or jhum , the predominant form of agriculture in the hill tracts of northeast India, is believed to have caused considerable loss of forest cover in the species-rich tropical rainforests of the region. In this study I sought to understand how rainforest bird communities are affected by shifting cultivation in Mizoram State. I studied bird occurrence and abundance patterns in secondary successional and mature tropical rainforests in a shifting-cultivation habitat mosaic in Dampa Tiger Reserve. To compare replicate sites in fallows aged 1, 5, 10, 25, and 100 years with undisturbed primary forest, I used systematic strip-transect sampling over the winter and early summer ( breeding) seasons during 1994–1995. Many forest bird species, especially those with ranges restricted to northeast India, declined in abundance or disappeared in successional fallows that had regenerated for ≤10 years. Birds that colonized or increased in abundance in regrowth habitats were mainly common and widespread species of open-country and secondary-forest habitats, and of low importance for conservation. Primary forest was the main habitat for specialized forest birds, intrinsically rare species, and elevational migrants. Although protection and conservation of relatively undisturbed mature forests is imperative in the core area of the reserve, management in the buffer zone for long-rotation shifting cultivation (>10-year cycles) instead of plantation of monocultures may be important until alternate means of livelihood are available to cultivators.     

Defining and Evaluating the Umbrella Species Concept for Conserving and Restoring Landscape Connectivity

Conserving or restoring landscape connectivity between patches of breeding habitat is a common strategy to protect threatened species from habitat fragmentation. By managing connectivity for some species, usually charismatic vertebrates, it is often assumed that these species will serve as conservation umbrellas for other species. We tested this assumption by developing a quantitative method to measure overlap in dispersal habitat of 3 threatened species—a bird (the umbrella), a butterfly, and a frog—inhabiting the same fragmented landscape. Dispersal habitat was determined with Circuitscape, which was parameterized with movement data collected for each species. Despite differences in natural history and breeding habitat, we found substantial overlap in the spatial distributions of areas important for dispersal of this suite of taxa. However, the intuitive umbrella species (the bird) did not have the highest overlap with other species in terms of the areas that supported connectivity. Nevertheless, we contend that when there are no irreconcilable differences between the dispersal habitats of species that cohabitate on the landscape, managing for umbrella species can help conserve or restore connectivity simultaneously for multiple threatened species with different habitat requirements. Definición y Evaluación del Concepto de Especie Paraguas para Conservar y Restaurar la Conectividad de Paisajes     

Importance of Reserve Size and Landscape Context to Urban Bird Conservation

Abstract:  We tested whether reserve size, landscape surrounding the reserve, and their interaction affect forest songbirds in the metropolitan area of Seattle, Washington (U.S.A.), by studying 29 reserves of varying size (small, medium, large) and surrounding urbanization intensity (urban, suburban, exurban). Larger reserves contained richer and less even bird communities than smaller reserves. These size effects disappeared when we removed the positive correlation of shrub diversity with reserve size, suggesting that greater habitat diversity in large reserves supported additional species, some of which were rare. Standardizing the number of individuals detected among all reserve size classes reversed the effect of size on richness in exurban landscapes and reduced the magnitude of the effect in suburban or urban landscapes. The latter change suggested that richness increased with reserve size in most landscapes because larger areas also supported larger samples from the regional bird species pool. Most bird species associated with native forest habitat (native forest species) and with human activity (synanthropic species) were present in reserves larger than 42 ha and surrounded by >40% urban land cover, respectively. Thus, we recommend these thresholds as means for conserving the composition of native bird communities in this mostly forested region. Native forest species were least abundant and synanthropic species most abundant in urban landscapes, where exotic ground and shrub vegetation was most common. Therefore, control of exotic vegetation may benefit native songbird populations. Bird nests in shrubs were most dense in medium (suburban) and large reserves (urban) and tended to be most successful in medium (suburban) and large reserves (exurban), potentially supplying another mechanism by which reserve size increased retention of native forest species.     

Edge Effects in the Great Tit: Analyses of Long-term Data with GIS Techniques

Abstract:  In contemporary fragmented landscapes, edges are commonplace, and understanding the effects of edge environments is thus essential for the conservation of forest communities. The reproductive output of forest passerines is often reduced close to forest edges. Possible explanations include overcrowding by conspecifics, elevated rates of predation, and the occurrence of lower-quality habitat and/or individuals at forest edges. We attempted to separate these processes by examining edge effects in the absence of nest predation and by effectively controlling for differences in breeding density and the quality of habitats and individuals. We used an edge distance index (EDI), which accounts for the number and distribution of edges in close proximity to a breeding location, to help explain variation in breeding density, nesting success, and reproductive traits of 8308 pairs of Great Tits ( Parus major ) breeding between 1965 and 2005, in Wytham, near Oxford, United Kingdom. Results from linear mixed modeling confirmed higher breeding density and a higher proportion of immigrant individuals at forest edges. Nevertheless, independently of these effects, we also found that birds laying later, with smaller clutches but larger eggs, were typical of edge environments. The number of offspring recruited to the breeding offspring per breeding attempt was also reduced at edges, both directly and mediated through changes in clutch size and laying date. Edge effects on life histories were detectable within individual females and up to 500 m from the woodland edge. Woodland edges are increasingly common in contemporary fragmented landscapes. Therefore these results, which suggest a pervasive effect of edges on reproduction, are of considerable importance to the management and conservation of forest communities.     

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.     

Quantifying relationships between bird and butterfly community shifts and environmental change.

Quantifying the manner in which ecological communities respond during a time of decreasing precipitation is a first step in understanding how they will respond to longer-term climate change. Here we coupled analysis of interannual variability in remotely sensed data with analyses of bird and butterfly community changes in montane meadow communities of the Greater Yellowstone Ecosystem. Landsat satellite imagery was used to classify these meadows into six types along a hydrological gradient. The northern portion of the ecosystem, or Gallatin region, has smaller mean patch sizes separated by ridges of mountains, whereas the southern portion of the ecosystem, or Teton region, has much larger patches within the Jackson Hole valley. Both support a similar suite of butterfly and bird species. The Gallatin region showed more overall among-year variation in the normalized difference vegetation index (NDVI) when meadow types were pooled within regions, perhaps because the patch sizes are smaller on average. Bird and butterfly communities showed significant relationships relative to meadow type and NDVI. We identified several key species that are tightly associated with specific meadow types along the hydrological gradient. Comparing taxonomic groups, fewer birds showed specific habitat affinities than butterflies, perhaps because birds are responding to differences in habitat structure among meadow types and using the landscape at a coarser scale than the butterflies. Comparing regions, the Teton region showed higher predictability of community assemblages as compared to the Gallatin region. The Gallatin region exhibited more significant temporal trends with respect to butterflies. Butterfly communities in wet meadows showed a distinctive shift along the hydrological gradient during a drought period (1997-2000). These results imply that the larger Teton meadows will show more predictable (i.e., static) species-habitat associations over the long term, but that the smaller Gallatin meadows may be an area that will exhibit the effects of global climate change faster.     

How well do sunken vessels approximate fish assemblages on coral reefs? Conservation implications of vessel-reef deployments

The amount of artificial habitat (termed ??artificial reef??, AR) in marine systems is rapidly increasing, yet the effect of most types of AR on reef communities remains unknown. We examined the role of well-established vessel-reefs in structuring coral reef fish assemblages by comparing assemblages on 7 World War II wrecks (>65?years old) to those on interspersed coral patch reefs of comparable size in a tropical lagoon. Fish abundance, species richness, diversity and feeding guild structure on wrecks were similar to natural reefs; however, species composition differed between the two reef types (R?=?0.189?C0.341, average dissimilarity: 67.3?C68.8?%). Despite being more species-rich and diverse, fish assemblages on larger wrecks were less similar to assemblages on their adjacent natural reefs than smaller wrecks. Wrecks may also have affected fish abundance on adjacent natural reefs, with reefs adjacent to larger wrecks supporting higher abundances than reefs adjacent to smaller wrecks. Our results indicate that increases in vessel-reef habitat may not greatly affect reef fish assemblage parameters, but may affect the relative abundances of particular species.     

Linking edge effects and patch size effects: Importance of matrix nest predators

The edge effect is usually considered to be the proximate cause of area sensitivity in forest birds. We tested if birds nesting in large patches are less vulnerable to the edge effect using a simple model that assumes an increase in patch size reduces the probability of a matrix predator moving to the core areas of forest and that larger perimeter/area ratios result in a higher number of matrix predators per unit of area. The probability of a nest being successful decreased asymptotically with an increase in either the patch penetration distance of predators or predator density, but those effects were reduced when patch size was increased. Large patches have a lower probability of being affected by an Allee effect and they can function as sink habitats only if penetration distance and predator density are largely increased. However, the transition from an Allee effect to a sink condition occurs with a small increase in penetration distance and predator density. Since birds nesting in large patches are less vulnerable to an increase in matrix predator populations, persistence of bird populations may be possible by increasing the size of habitat patches that can act as source populations.     

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.     

Effects of landscape transformation on bird colonization and extinction patterns in a large‐scale,long‐term natural experiment

Conversion of agricultural land to forest plantations is a major driver of global change. Studies on the impact of forest plantations on biodiversity in plantations and in the surrounding native vegetation have been inconclusive. Consequently, it is not known how to best manage the extensive areas of the planet currently covered by plantations. We used a novel, long‐term (16 years) and large‐scale (30,000 ha) landscape transformation natural experiment (the Nanangroe experiment, Australia) to test the effects of land conversion on population dynamics of 64 bird species associated with woodland and forest. A unique aspect of our study is that we focused on the effects of plantations on birds in habitat patches within plantations. Our study design included 56 treatment sites (Eucalyptus patches where the surrounding matrix was converted from grazed land to pine plantations), 55 control sites (Eucalyptus patches surrounded by grazed land), and 20 matrix sites (sites within the pine plantations and grazed land). Bird populations were studied through point counts, and colonization and extinction patterns were inferred through multiple season occupancy models. Large‐scale pine plantation establishment affected the colonization or extinction patterns of 89% of studied species and thus led to a comprehensive turnover in bird communities inhabiting Eucalyptus patches embedded within the maturing plantations. Smaller bodied species appeared to respond positively to plantations (i.e., colonization increased and extirpation of these species decreased in patches surrounded by plantations) because they were able to use the newly created surrounding matrix. We found that the effects of forest plantations affected the majority of the bird community, and we believe these effects could lead to the artificial selection of one group of species at the expense of another.     

Distribution of Carabid Beetles (Coleoptera, Carabidae) across a Boreal Forest–Clearcut Ecotone

Abstract: We studied the occurrence of carabid beetles (Coleoptera, Carabidae) in boreal forest fragments, their edges, and adjacent clearcuts in central Finland. Beetles were collected with pitfall traps along transects extending 60 m from the edge into clearcuts and 60 m into forest interior. Our main findings were that (1) species richness was significantly higher in the clearcut than in the forest fragments, (2) clearcuts hosted many open-habitat species, which increased overall species richness in these sites, (3) carabid assemblages in the edges were more similar to forest assemblages than to those found in the clearcuts, (4) no edge specialists were found, and (5) open-habitat species did not penetrate into the forest fragments from the clearcut. Because forest specialists occurred all the way to the edge on the forest side, it seems that edge effects per se do not adversely affect these species, at least in the short term. In the long term, however, habitat conditions in the edges may deteriorate for interior species because of trees falling over in strong winds, thereby reducing the size of the fragments and widening the edge zone.     

Diversity-invasibility across an experimental disturbance gradient in Appalachian Forests

Research examining the relationship between community diversity and invasions by nonnative species has raised new questions about the theory and management of biological invasions. Ecological theory predicts, and small-scale experiments confirm, lower levels of nonnative species invasion into species-rich compared to species-poor communities, but observational studies across a wider range of scales often report positive relationships between native and nonnative species richness. This paradox has been attributed to the scale dependency of diversity-invasibility relationships and to differences between experimental and observational studies. Disturbance is widely recognized as an important factor determining invasibility of communities, but few studies have investigated the relative and interactive roles of diversity and disturbance on nonnative species invasion. Here, we report how the relationship between native and nonnative plant species richness responded to an experimentally applied disturbance gradient (from no disturbance up to clearcut) in oak-dominated forests. We consider whether results are consistent with various explanations of diversity-invasibility relationships including biotic resistance, resource availability, and the potential effects of scale (1 m2 to 2 ha). We found no correlation between native and nonnative species richness before disturbance except at the largest spatial scale, but a positive relationship after disturbance across scales and levels of disturbance. Post-disturbance richness of both native and nonnative species was positively correlated with disturbance intensity and with variability of residual basal area of trees. These results suggest that more nonnative plants may invade species-rich communities compared to species-poor communities following disturbance.