Lars A. Brudvig

Habitat fragmentation and its lasting impact on Earth's ecosystems

Urgent need for conservation and restoration measures to improve landscape connectivity. We conducted an analysis of global forest cover to reveal that 70% of remaining forest is within 1 km of the forest’s edge, subject to the degrading effects of fragmentation. A synthesis of fragmentation experiments spanning multiple biomes and scales, five continents, and 35 years demonstrates that habitat fragmentation reduces biodiversity by 13 to 75% and impairs key ecosystem functions by decreasing biomass and altering nutrient cycles. Effects are greatest in the smallest and most isolated fragments, and they magnify with the passage of time. These findings indicate an urgent need for conservation and restoration measures to improve landscape connectivity, which will reduce extinction rates and help maintain ecosystem services.

The restoration of biodiversity: Where has research been and where does it need to go?

The practice of ecological restoration is a primary option for increasing levels of biodiversity by modifying human-altered ecosystems. The scientific discipline of restoration ecology provides conceptual guidance and tests of restoration strategies, with the ultimate goal of predictive landscape restoration. I construct a conceptual model for restoration of biodiversity, based on site-level (e.g., biotic and abiotic) conditions, landscape (e.g, interpatch connectivity and patch geometry), and historical factors (e.g., species arrival order and land-use legacies). I then ask how well restoration ecology has addressed the various components of this model. During the past decade, restoration research has focused largely on how the restoration of site-level factors promotes species diversity-primarily of plants. Relatively little attention has been paid to how landscape or historical factors interplay with restoration, how restoration influences functional and genetic components of biodiversity, or how a suite of less-studied taxa might be restored. I suggest that the high level of variation seen in restoration outcomes might be explained, at least in part, by the contingencies placed on site-level restoration by landscape and historical factors and then present a number of avenues for future research to address these often ignored linkages in the biodiversity restoration model. Such work will require carefully conducted restoration experiments set across multiple sites and many years. It is my hope that by considering how space and time influence restoration, we might move restoration ecology in a direction of stronger prediction, conducted across landscapes, thus providing feasible restoration strategies that work at scales over which biodiversity conservation occurs.

The movement ecology and dynamics of plant communities in fragmented landscapes

A conceptual model of movement ecology has recently been advanced to explain all movement by considering the interaction of four elements: internal state, motion capacity, navigation capacities, and external factors. We modified this framework to generate predictions for species richness dynamics of fragmented plant communities and tested them in experimental landscapes across a 7-year time series. We found that two external factors, dispersal vectors and habitat features, affected species colonization and recolonization in habitat fragments and their effects varied and depended on motion capacity. Bird-dispersed species richness showed connectivity effects that reached an asymptote over time, but no edge effects, whereas wind-dispersed species richness showed steadily accumulating edge and connectivity effects, with no indication of an asymptote. Unassisted species also showed increasing differences caused by connectivity over time, whereas edges had no effect. Our limited use of proxies for movement ecology (e.g., dispersal mode as a proxy for motion capacity) resulted in moderate predictive power for communities and, in some cases, highlighted the importance of a more complete understanding of movement ecology for predicting how landscape conservation actions affect plant community dynamics.

How fragmentation and corridors affect wind dynamics and seed dispersal in open habitats

Significance Understanding how widespread human-induced global changes are affecting the movement and dispersal of organisms is critical for maintaining species diversity and making sound land management decisions. In contrast with animal-dispersed species, little is known about how wind-dispersed species are affected by conservation strategies such as corridors. We use a combination of mechanistic models and field data to show that habitat corridors alter wind dynamics in a way that promotes seed dispersal and appears to increase plant diversity. Wind direction also interacts with landscape orientation to determine when corridors can provide connectivity. Determining how widespread human-induced changes such as habitat loss, landscape fragmentation, and climate instability affect populations, communities, and ecosystems is one of the most pressing environmental challenges. Critical to this challenge is understanding how these changes are affecting the movement abilities and dispersal trajectories of organisms and what role conservation planning can play in promoting movement among remaining fragments of suitable habitat. Whereas evidence is mounting for how conservation strategies such as corridors impact animal movement, virtually nothing is known for species dispersed by wind, which are often mistakenly assumed to not be limited by dispersal. Here, we combine mechanistic dispersal models, wind measurements, and seed releases in a large-scale experimental landscape to show that habitat corridors affect wind dynamics and seed dispersal by redirecting and bellowing airflow and by increasing the likelihood of seed uplift. Wind direction interacts with landscape orientation to determine when corridors provide connectivity. Our results predict positive impacts of connectivity and patch shape on species richness of wind-dispersed plants, which we empirically illustrate using 12 y of data from our experimental landscapes. We conclude that habitat fragmentation and corridors strongly impact the movement of wind-dispersed species, which has community-level consequences.

EDITOR'S CHOICE: Confronting contingency in restoration: management and site history determine outcomes of assembling prairies, but site characteristics and landscape context have little effect

Summary 1. The outcomes of ecological restoration are notoriously unpredictable, but we have no general predictive understanding of this contingency. Management decisions can have strong effects on restoration outcomes, but in other cases may be overwhelmed by site characteristics (e.g. soil conditions), landscape context (e.g. abundance of similar habitat) or historical factors (e.g. priority effects). However, we generally cannot predict which of these four classes of drivers will affect restoration outcomes. Disparate aspects of restoration outcomes (e.g. species richness, beta diversity and community composition) and their unique responses further complicate our understanding. Finally, these four classes of drivers might differentially affect subsets of the restored community, where, for example, management might shape the abundance and distribution of species of the target community, while other species are more contingent on site, landscape or historical factors. 2. Here, we used variation partitioning to compare the relative importance of management, site, landscape and historical factors for determining the plant community outcomes of 27 prairie restorations in south-west Michigan. 3. We found that management, especially the composition, diversity and density of seed mixes applied, and history, especially site age, were the most important drivers of prairie restoration species richness, beta diversity and composition. Site and landscape factors were only rarely important for restoration outcomes. 4. Finally, we found that comparing the unique responses of sown and non-sown species typically increased our understanding of the dynamics contributing to community-wide restoration outcomes. 5. Synthesis and applications. This is, to our knowledge, the first quantitative comparison of how four major classes of drivers determine the outcome of restoration. Historical legacies and management decisions, but generally not landscape context or local site conditions, shaped plant communities at restored sites. These findings represent an important step towards developing a more predictive framework for understanding contingency in restoration outcomes.

Experimental evidence does not support the Habitat Amount Hypothesis

&NA; For a half century, habitat configuration – the arrangement of habitat patches within a landscape – has been central to theories of landscape ecology, population dynamics, and community assembly, in addition to conservation strategies. A recent hypothesis advanced by Fahrig (2013) would, if supported, greatly diminish the relevance of habitat configuration as a predictor of diversity. The Habitat Amount Hypothesis posits that the sample area effect overrides patch size and patch isolation effects of habitat fragmentation on species richness. It predicts that the amount of habitat in a local landscape, regardless of configuration, is the main landscape‐level determinant of species richness. If habitat amount is indeed the major, landscape‐level driver of species richness, the slopes of the species–area relationship (SAR) for otherwise similar fragmented and unfragmented landscapes should be indistinguishable. We tested the Habitat Amount Hypothesis with data from two replicated and controlled habitat fragmentation experiments that disentangle the effects of habitat amount and configuration. One experiment provided time‐series data on plant species richness and the other on micro‐arthropod species richness. We found that, relative to less fragmented habitats, the SARs for fragmented habitats have significantly higher slopes and that the magnitude of the difference in slopes increased over time. Relatively more species were lost in smaller areas when fragments were more isolated. In both experiments, the proportion of species lost due to increased habitat fragmentation was nearly identical to the proportion lost due to reduced habitat amount. Our results provide a direct and experimentally derived refutation of the Habitat Amount Hypothesis, supporting the long‐held view that in addition to area, patch isolation and configuration are important determinants of species richness. Differences in species richness between fragmented and non‐fragmented habitats increase over time, demonstrating that long‐term studies are needed to understand the effects of fragmentation, above and beyond the amount of habitat lost.

Understory plant communities and the functional distinction between savanna trees, forest trees, and pines

Although savanna trees and forest trees are thought to represent distinct functional groups with different effects on ecosystem processes, few empirical studies have examined these effects. In particular, it remains unclear if savanna and forest trees differ in their ability to coexist with understory plants, which comprise the majority of plant diversity in most savannas. We used structural equation modeling (SEM) and data from 157 sites across three locations in the southeastern United States to understand the effects of broadleaf savanna trees, broadleaf forest trees, and pine trees on savanna understory plant communities. After accounting for underlying gradients in fire frequency and soil moisture, abundances (i.e., basal area and stem density) of forest trees and pines, but not savanna trees, were negatively correlated with the cover and density (i.e., local-scale species richness) of C4 graminoid species, a defining savanna understory functional group that is linked to ecosystem flammability. In analyses of the full understory community, abundances of trees from all functional groups were negatively correlated with species density and cover. For both the C4 and full communities, fire frequency promoted understory plants directly, and indirectly by limiting forest tree abundance. There was little indirect influence of fire on the understory mediated through savanna trees and pines, which are morefire tolerant than forest trees. We conclude that tree functional identity is an important factor that influences overstory tree relationships with savanna understory plant communities. In particular, distinct relationships between trees and C4 graminoids have implications for grass-tree coexistence and vegetation-fire feedbacks that maintain savanna environments and their associated understory plant diversity.

Potential Negative Ecological Effects of Corridors

Despite many studies showing that landscape corridors increase dispersal and species richness for disparate taxa, concerns persist that corridors can have unintended negative effects. In particular, some of the same mechanisms that underlie positive effects of corridors on species of conservation interest may also increase the spread and impact of antagonistic species (e.g., predators and pathogens), foster negative effects of edges, increase invasion by exotic species, increase the spread of unwanted disturbances such as fire, or increase population synchrony and thus reduce persistence. We conducted a literature review and meta-analysis to evaluate the prevalence of each of these negative effects. We found no evidence that corridors increase unwanted disturbance or non-native species invasion; however, these have not been well-studied concerns (1 and 6 studies, respectively). Other effects of corridors were more often studied and yielded inconsistent results; mean effect sizes were indistinguishable from zero. The effect of edges on abundances of target species was as likely to be positive as negative. Corridors were as likely to have no effect on antagonists or population synchrony as they were to increase those negative effects. We found 3 deficiencies in the literature. First, despite studies on how corridors affect predators, there are few studies of related consequences for prey population size and persistence. Second, properly designed studies of negative corridor effects are needed in natural corridors at scales larger than those achievable in experimental systems. Third, studies are needed to test more targeted hypotheses about when corridor-mediated effects on invasive species or disturbance may be negative for species of management concern. Overall, we found no overarching support for concerns that construction and maintenance of habitat corridors may result in unintended negative consequences. Negative edge effects may be mitigated by widening corridors or softening edges between corridors and the matrix. Other negative effects are relatively small and manageable compared with the large positive effects of facilitating dispersal and increasing diversity of native species.

Landscape connectivity strengthens local–regional richness relationships in successional plant communities

Local species diversity is maintained over ecological time by a balance between dispersal and species interactions. Local-regional species richness relationships are often used to investigate the relative importance of these two processes and the scales at which they operate. For communities undergoing succession, theory predicts a temporal progression in local-regional species richness relationships: from no relationship to positive linear to saturating. However, observational tests have been mixed, and experiments have been rare. Using a replicated large-scale experiment, we evaluate the impact of two dispersal-governing processes at the regional scale, connectivity and shape of the region (i.e., patches), on the progression of local-regional species richness relationships for plant communities undergoing succession. Regional connectivity accelerates the transition from no relationship to a positive linear relationship, while the shape of the region has no consistent effect nine years post-disturbance. Our results experimentally demonstrate the importance of dispersal in structuring local-regional species richness relationships over time and suggest that conservation corridors among regions can increase local diversity through regional enrichment of plant communities undergoing reassembly.

Can dispersal mode predict corridor effects on plant parasites

Habitat corridors, a common management strategy for increasing connectivity in fragmented landscapes, have experimentally validated positive influences on species movement and diversity. However, long-standing concerns that corridors could negatively impact native species by spreading antagonists, such as disease, remain largely untested. Using a large-scale, replicated experiment, we evaluated whether corridors increase the incidence of plant parasites. We found that corridor impacts varied with parasite dispersal mode. Connectivity provided by corridors increased incidence of biotically dispersed parasites (galls on Solidago odora) but not of abiotically dispersed parasites (foliar fungi on S. odora and three Lespedeza spp.). Both biotically and abiotically dispersed parasites responded to edge effects, but the direction of responses varied across species. Although our results require additional tests for generality to other species and landscapes, they suggest that, when establishing conservation corridors, managers should focus on mitigating two potential negative effects: the indirect effects of narrow corridors in creating edges and direct effects of corridors in enhancing connectivity of biotically dispersed parasites.