Nick M. Haddad

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.

Corridors affect plants, animals, and their interactions in fragmented landscapes

Among the most popular strategies for maintaining populations of both plants and animals in fragmented landscapes is to connect isolated patches with thin strips of habitat, called corridors. Corridors are thought to increase the exchange of individuals between habitat patches, promoting genetic exchange and reducing population fluctuations. Empirical studies addressing the effects of corridors have either been small in scale or have ignored confounding effects of increased habitat area created by the presence of a corridor. These methodological difficulties, coupled with a paucity of studies examining the effects of corridors on plants and plant–animal interactions, have sparked debate over the purported value of corridors in conservation planning. We report results of a large-scale experiment that directly address this debate. In eight large-scale experimental landscapes that control for patch area and test alternative mechanisms of corridor function, we demonstrate that corridors not only increase the exchange of animals between patches, but also facilitate two key plant–animal interactions: pollination and seed dispersal. Our results show that the beneficial effects of corridors extend beyond the area they add, and suggest that increased plant and animal movement through corridors will have positive impacts on plant populations and community interactions in fragmented landscapes.

Contrasting Effects of Plant Richness and Composition on Insect Communities: A Field Experiment

We experimentally separated the effects of two components of plant diversity—plant species richness and plant functional group richness—on insect communities. Plant species richness and plant functional group richness had contrasting effects on insect abundances, a result we attributed to three factors. First, lower insect abundances at higher plant functional group richness were explained by a sampling effect, which was caused by the increasing likelihood that one low‐quality group, C4 grasses, would be present and reduce average insect abundances by 25%. Second, plant biomass, which was positively related to plant functional group richness, had a strong, positive effect on insect abundances. Third, a positive effect of plant species richness on insect abundances may have been caused by greater availability of alternate plant resources or greater vegetational structure. In addition, a greater diversity of insect species, whose individual abundances were often unaffected by changes in plant species richness, may have generated higher total community abundances. After controlling for the strong, positive influence of insect abundance on insect diversity through rarefaction, insect species richness increased as plant species richness and plant functional group richness increased. Although these variables did not explain a high proportion of variation individually, plant species richness and plant functional group richness had similar effects on insect diversity and opposing effects on insect abundances, and both factors may explain how the loss of plant diversity influences higher trophic levels.

CORRIDOR USE BY DIVERSE TAXA

One of the most popular approaches for maintaining populations and con- serving biodiversity in fragmented landscapes is to retain or create corridors that connect otherwise isolated habitat patches. Working in large-scale, experimental landscapes in which open-habitat patches and corridors were created by harvesting pine forest, we showed that corridors direct movements of different types of species, including butterflies, small mam- mals, and bird-dispersed plants, causing higher movement between connected than between unconnected patches. Corridors directed the movement of all 10 species studied, with all corridor effect sizes .68%. However, this corridor effect was significant for five species, not significant for one species, and inconclusive for four species because of small sample sizes. Although we found no evidence that corridors increase emigration from a patch, our results show that movements of disparate taxa with broadly different life histories and functional roles are directed by corridors.

Plant species loss decreases arthropod diversity and shifts trophic structure

Plant diversity is predicted to be positively linked to the diversity of herbivores and predators in a foodweb. Yet, the relationship between plant and animal diversity is explained by a variety of competing hypotheses, with mixed empirical results for each hypothesis. We sampled arthropods for over a decade in an experiment that manipulated the number of grassland plant species. We found that herbivore and predator species richness were strongly, positively related to plant species richness, and that these relationships were caused by different mechanisms at herbivore and predator trophic levels. Even more dramatic was the threefold increase, from low- to high-plant species richness, in abundances of predatory and parasitoid arthropods relative to their herbivorous prey. Our results demonstrate that, over the long term, the loss of plant species propagates through food webs, greatly decreasing arthropod species richness, shifting a predator-dominated trophic structure to being herbivore dominated, and likely impacting ecosystem functioning and services.

Corridor Use Predicted from Behaviors at Habitat Boundaries

Through empirical studies and simulation, I demonstrate how simple behaviors can be used in lieu of detailed dispersal studies to predict the effects of corridors on interpatch movements. Movement paths of three butterfly species were measured in large (1.64 ha) experimental patches of open habitat, some of which were connected by corridors. Butterflies that “reflected” off boundaries between open patches and the surrounding forest also emigrated from patches through corridors at rates higher than expected from random movement. This was observed for two open‐habitat species, Eurema nicippe and Phoebis sennae; however, edges and corridors had no effect on a habitat generalist, Papilio troilus. Behaviorally based simulation models, which departed from correlated random walks only at habitat boundaries, predicted that corridors increase interpatch movement rates of both open‐habitat species. Models also predicted that corridors have proportionately greater effects as corridor width increases, that movement rates increase before leveling off as corridor width increases, and that corridor effects decrease as patch size increases. This study suggests that corridors direct movements of habitat‐restricted species and that local behaviors may be used to predict the conservation potential of corridors in fragmented landscapes.

Corridors Increase Plant Species Richness at Large Scales

Habitat fragmentation is one of the largest threats to biodiversity. Landscape corridors, which are hypothesized to reduce the negative consequences of fragmentation, have become common features of ecological management plans worldwide. Despite their popularity, there is little evidence documenting the effectiveness of corridors in preserving biodiversity at large scales. Using a large-scale replicated experiment, we showed that habitat patches connected by corridors retain more native plant species than do isolated patches, that this difference increases over time, and that corridors do not promote invasion by exotic species. Our results support the use of corridors in biodiversity conservation.

CORRIDOR AND DISTANCE EFFECTS ON INTERPATCH MOVEMENTS: A LANDSCAPE EXPERIMENT WITH BUTTERFLIES

The hypothesis that corridors increase animal movement between habitat fragments, a central tenet of conservation biology, has been virtually untested. This study demonstrates that corridors increase interpatch movement rates of two butterfly species. The hypothesis was tested in a large-scale, replicated experiment, in which 27 equal-sized (1.64-ha) patches of early successional habitat were created within large areas of pine forest. Patches varied in whether or not they were connected to another patch by a corridor, and in their distance from other patches (64–384 m). The results of mark–release–recapture studies showed that two open-habitat butterfly species, Junonia coenia and Euptoieta claudia, moved more frequently between patches connected by corridors than between unconnected patches. Interpatch movement was significantly, negatively related to interpatch distance. Interpatch movement rates of J. coenia were significantly, positively related to the density of its host and nectar resource, Linaria canadensis. Corridor effects were stronger for males than for females and were most pronounced within three days after butterflies were marked. Pine forest was not a complete barrier to butterfly movement; both species moved between unconnected patches, even at the longest distances. However, the results of this study suggest that corridors will increase long-distance movements of habitat-restricted species.

AN EXPERIMENTAL TEST OF CORRIDOR EFFECTS ON BUTTERFLY DENSITIES

In a large-scale experiment, we found that three habitat-restricted butterfly species reached higher densities in patches connected by corridors than in similar, isolated patches. We conducted our study in 27 equal-sized (1.64-ha) patches that varied in whether or not they were connected to another patch by a corridor. Patches and corridors were open, early-successional habitat that contrasted strongly with the surrounding pine forest. Of four butterfly species studied, three open-habitat specialists (Junonia coenia, Euptoieta claudia, and Phoebis sennae) reached higher densities in patches connected by corridors than in isolated patches. A fourth species, Papilio troilus, showed no preference for open habitat or pine forest, and its density did not differ in connected or isolated patches. Although butterfly densities were often significantly, positively associated with densities of host plant or flower resources, plant densities did not confound corridor effects on butterfly densities. Higher densities i...

The effects of long-term nitrogen loading on grassland insect communities

Abstract Just as long-term nitrogen loading of grasslands decreases plant species richness and increases plant biomass, we have found that nitrogen loading decreases insect species richness and increases insect abundances. We sampled 54 plots that had been maintained at various rates of nitrogen addition for 14 years. Total insect species richness and effective insect diversity, as well as herbivore and predator species richness, were significantly, negatively related to the rate of nitrogen addition. However, there was variation in trophic responses to nitrogen. Detritivore species richness increased as nitrogen addition increased, and parasitoids showed no response. Insect abundances, measured as the number of insects and insect biovolume (an estimate of biomass), were significantly, positively related to the rate of nitrogen addition, as were the abundances of herbivores and detritivores. Parasitoid abundance was negatively related to the rate of nitrogen addition. Changes in the insect community were correlated with changes in the plant community. As rates of nitrogen addition increased, plant species richness decreased, plant productivity and plant tissue nitrogen increased, and plant composition shifted from C4 to C3 grass species. Along this gradient, total insect species richness and effective insect diversity were most strongly, positively correlated with plant species richness. Insect biovolume was negatively correlated with plant species richness. Responses of individual herbivores varied along the nitrogen gradient, but numbers of 13 of the 18 most abundant herbivores were positively correlated with their host plant biomass. Although insect communities did not respond as strongly as plant communities, insect species richness, abundance, and composition were impacted by nitrogen addition. This study demonstrates that long-term nitrogen loading affects the entire food chain, simplifying both plant and insect communities.