Bryan L. Foster

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.

SPECIES RICHNESS IN A SUCCESSIONAL GRASSLAND: EFFECTS OF NITROGEN ENRICHMENT AND PLANT LITTER

We conducted a field experiment in a successional grassland to investigate the short-term effects of nitrogen enrichment and plant litter on plant species richness and on the establishment of a native grass (Andropogon gerardi) that was experimentally introduced to the study site as seed. Additions of nitrogen fertilizer to experimental plots over two growing seasons increased plant production as indicated by increases in both living plant biomass and litter biomass. Increased productivity reduced species richness by effectively preventing the seedling establishment of the subordinate forb species and reduced the recruitment of Andropogon gerardi by inhibiting both germination and survival. Litter removals carried out at each of two levels of nitrogen enrichment (no nitrogen, nitrogen added) showed that litter significantly reduced species richness by the same amount in fertilized and unfertilized plots, suggesting that living biomass and litter were purely additive, rather than interactive in their effects. In contrast, the results of litter additions suggested that the declines in richness associated with fertilization could largely be due to the suppressive effects of increased litter biomass alone. As a whole, the results indicate that litter and living biomass are largely substitutable in their inhibitory effects on species richness in highly productive successional grasslands due to their independent and equivalent capacities to attenuate light to very low levels. This study highlights the combined roles of competition and plant litter in influencing the diversity of grasslands through effects on seedling establishment.

Worldwide evidence of a unimodal relationship between productivity and plant species richness

Grassland diversity and ecosystem productivity The relationship between plant species diversity and ecosystem productivity is controversial. The debate concerns whether diversity peaks at intermediate levels of productivity—the so-called humped-back model—or whether there is no clear predictable relationship. Fraser et al. used a large, standardized, and geographically diverse sample of grasslands from six continents to confirm the validity and generality of the humped-back model. Their findings pave the way for a more mechanistic understanding of the factors controlling species diversity. Science, this issue p. 302 The humped-back model of plant species diversity is confirmed by a global grassland survey. The search for predictions of species diversity across environmental gradients has challenged ecologists for decades. The humped-back model (HBM) suggests that plant diversity peaks at intermediate productivity; at low productivity few species can tolerate the environmental stresses, and at high productivity a few highly competitive species dominate. Over time the HBM has become increasingly controversial, and recent studies claim to have refuted it. Here, by using data from coordinated surveys conducted throughout grasslands worldwide and comprising a wide range of site productivities, we provide evidence in support of the HBM pattern at both global and regional extents. The relationships described here provide a foundation for further research into the local, landscape, and historical factors that maintain biodiversity.

SECONDARY SUCCESSION IN AN EXPERIMENTALLY FRAGMENTED LANDSCAPE: COMMUNITY PATTERNS ACROSS SPACE AND TIME

Secondary succession reflects, at least in part, community assembly—the sequences of colonizations and extinctions. These processes in turn are expected to be sensitive to the size of the site undergoing assembly and its location relative to source pools. In this paper we describe patterns of succession over 18 years in an experimentally fragmented landscape created in eastern Kansas, USA, in 1984. The design of the experiment permits one to assess the influence of patch size and landscape position on successional dynamics. The general trajectory of succession follows that typical of succession in much of the eastern United States. In the initial years of the study, there was relatively little effect of patch size or distance to sources. Here we show that spatial effects in this system have become increasingly evident with time, as gauged both by repeated-measures ANOVA and ordination techniques. Woody plants have colonized more rapidly (per unit area) on large and nearby patches. Species richness at a local (within-quadrat) scale in general has increased, with slightly greater richness in large than in small patches later in the study. Temporal stability in community composition has generally been greater in large patches. Spatial heterogeneity in community composition has increased during succession, but with different patterns in large and small patches. This long-term experiment suggests that landscape structure influences many aspects of community structure and dynamics during succession, and that such effects become more pronounced with the passage of time.

Coexistence through spatio-temporal heterogeneity and species sorting in grassland plant communities

The effect of spatial heterogeneity on species coexistence relies on the degree of niche heterogeneity in the habitat and the ability of species to exploit the available niche opportunities. We studied species coexistence in a perennial grassland, and tested whether small-scale disturbances create environmental heterogeneity that affects coexistence and whether the functional diversity of species in the species pool affects the ability of community composition to reflect heterogeneity through species sorting. We manipulated the spatio-temporal heterogeneity of disturbance and the functional diversity of species added as seed and measured their impact on the spatial turnover of species composition. Disturbance increased environmental heterogeneity and spatial turnover, and the effect of heterogeneity on turnover was greatest in the presence of a functionally diverse species pool, showing the importance of trait variation among species for exploiting environmental heterogeneity, and suggesting that coexistence occurred due to species sorting among heterogeneous niches.

GRASSLAND DIVERSITY AND PRODUCTIVITY: THE INTERPLAY OF RESOURCE AVAILABILITY AND PROPAGULE POOLS

Processes operating at multiple spatial scales govern the structure and functioning of ecological communities. We conducted a resource manipulation and propagule addition experiment in grassland to evaluate the interaction of local resource availability and propagule pools in governing local-scale plant colonization, biodiversity, and aboveground productivity. The availabilities of establishment microsites and water were manipulated in field plots for two years through the application of experimental soil disturbances and irrigation, respectively. Resource manipulations led to increased invasibility of the community, as predicted by the theory of fluctuating resources. Rates of colonization, enhanced by the sowing of 32 grassland species, increased plant diversity and aboveground productivity, but to a greater extent under conditions of resource enrichment. Although resource enrichment generally increased diversity and productivity, these responses were contingent upon species availability and tended to be more pronounced in the presence of an expanded propagule pool. These findings suggest that biodiversity at the level of the available propagule pool and fluctuations in resources interact to regulate local resident diversity and productivity by determining opportunities for species sorting, by mediating community assembly, and by governing the potential for functional compensation in the community.

PARTITIONING THE EFFECTS OF PLANT BIOMASS AND LITTER ON ANDROPOGON GERARDI IN OLD‐FIELD VEGETATION

We examined the effects of living plant neighbors and litter on the perfor- mance of a native C4 grass, Andropogon gerardi, at five old-field sites that differ in com- munity biomass and soil fertility. We used plant removal experiments in which both living neighbors and litter were manipulated in a factorial arrangement of treatments over one growing season. Andropogon was added to treatment plots as seeds and as established transplants to examine the effects of the surrounding plant community on the recruitment and established phases of its life history. Neighbors negatively affected Andropogon performance at all sites, indicating that resource exploitation by living plants was an important constraint to seedling recruitment and growth across the range of community biomass examined. Plant litter negatively affected recruitment at sites with the greatest community biomass, but had no effect on the growth of established transplants at any of the sites. The total effect of the surrounding plant community on recruitment was positively correlated with community biomass due to an increasing impact of plant litter. However, the total effect of the surrounding community on the growth of established transplants was unrelated to community biomass. The results suggest that it may be during the recruitment phase of the life history, when seeds and seedlings are especially susceptible to the effects of both litter and living neigh- bors, that Andropogon is most sensitive to variation in community biomass. Variation in the effects of litter on recruitment may be important in regulating plant species diversity and the distribution of native C4 grasses along old-field biomass gradients in southwest Michigan.

Fertilization decreases plant biodiversity even when light is not limiting

Many researchers hypothesize that plant richness declines at high soil fertility (and high productivity) due to light limitation. We tested this hypothesis in an old-field by independently manipulating fertilization and light levels via shade cloth (decreased light), vegetation tie-backs (increased light) and vegetation clipping (increased light). Droughts occurred during two of the four years of the study, and we found that higher light levels were generally associated with decreased plant richness in drought years but increased plant richness in wet years. Most importantly, fertilization decreased richness whether light availability limited richness (wet years) or did not limit richness (drought years), and the effects of fertilization and light manipulation treatments were additive. These results suggest that effects of fertilization on plant richness are at least partly independent of light levels and that competition for resources other than light plays a substantial role in the decline of plant richness after fertilization.

Community-level consequences of mycorrhizae depend on phosphorus availability

In grasslands, arbuscular mycorrhizal fungi (AMF) mediate plant diversity; whether AMF increase or decrease diversity depends on the relative mycotrophy in dominant vs. subordinate plants. In this study we investigated whether soil nutrient levels also influence the ability of AMF to mediate plant species coexistence. First, we developed a conceptual model that predicts the influence of AMF on diversity along a soil nutrient gradient for plant communities dominated by mycotrophic and non-mycotrophic species. To test these predictions, we manipulated phosphorus to create a soil nutrient gradient for mesocosm communities composed of native prairie grasses and then compared community properties for mesocosms with and without AMF. We found that, where P was limiting, AMF increased plant diversity and productivity, and also altered community structure; however, at high P, AMF had little influence on aboveground communities. Compositional differences among treatments were due largely to a trade-off in the relative abundance of C3 vs. C4 spes. Our study emphasizes how environmental constraints on mutualisms may govern community- and ecosystem-level properties.

Effects of hay management and native species sowing on grassland community structure, biomass, and restoration.

Prairie hay meadows are important reservoirs of grassland biodiversity in the tallgrass prairie regions of the central United States and are the object of increasing attention for conservation and restoration. In addition, there is growing interest in the potential use of such low-input, high-diversity (LIHD) native grasslands for biofuel production. The uplands of eastern Kansas, USA, which prior to European settlement were dominated by tallgrass prairie, are currently utilized for intensive agriculture or exist in a state of abandonment from agriculture. The dominant grasslands in the region are currently high-input, low-diversity (HILD) hay fields seeded to introduced C3 hay grasses. We present results from a long-term experiment conducted in a recently abandoned HILD hay field in eastern Kansas to evaluate effects of fertilization, haying, and native species sowing on community dynamics, biomass, and potential for restoration to native LIHD hay meadow. Fertilized plots maintained dominance by introduced grasses, maintained low diversity, and were largely resistant to colonization throughout the study. Non-fertilized plots exhibited rapid successional turnover, increased diversity, and increased abundance of C4 grasses over time. Haying led to modest changes in species composition and lessened the negative impact of fertilization on diversity. In non-fertilized plots, sowing increased representation by native species and increased diversity, successional turnover, and biomass production. Our results support the shifting limitations hypothesis of community organization and highlight the importance of species pools and seed limitations in constraining successional turnover, community structure, and ecosystem productivity under conditions of low fertility. Our findings also indicate that several biological and functional aspects of LIHD hay meadows can be restored from abandoned HILD hay fields by ceasing fertilization and reintroducing native species through sowing. Declines in primary production and hay yield that result from the cessation of fertilization may be at least partially compensated for by restoration.