Disjunct performance and distribution in the sedge Carex prasina.

Efforts to understand species distributions and predict responses to environmental changes depend on specifying how the abiotic environment determines distributions. At landscape scales, it is critical to distinguish effects of environmental factors from other mechanisms such as competition and dispersal limitation. We examined how environmental factors affect the distribution and performance of the sedge Carex prasina across a 10-km(2) old-growth forest in southern Québec. We isolated the effects of soil characteristics by conducting a greenhouse experiment that assessed the performance of C. prasina on soils from a range of wetland habitats where it could potentially occur. This allowed us to compare how the species' performance and its distribution across the landscape relate to the same soil characteristics. In the experiment, the biomass and leaf chlorophyll content of C. prasina increased with increasing soil organic matter (OM). Across the landscape, however, the species' probability of occurrence and abundance decreased with increasing soil OM. C. prasina had similar biomass on soils from sites where it did and did not occur, but it had higher leaf chlorophyll content on soils from sites where it did not occur. We found no evidence that differential performance across environments determines the distribution of this species, as C. prasina tended to occur on soils where it showed reduced performance. Rather, other processes such as competition or dispersal limitation likely override any direct effects of the soil environment on distribution. Our results caution against the common assumption that the environments where a species tends to occur or be most abundant are the environments where it performs best. C. prasina presents a clear example of a species whose performance, at least along edaphic gradients, cannot explain its distribution. This example highlights the importance of distinguishing the relative roles of biotic and abiotic factors that shape species distributions across landscapes.

Plant species diversity and composition of wetlands within an upland forest.

Though often overlooked, small wetlands in an upland matrix can support diverse plant communities that increase both local and regional species richness. Here we characterize the full range of wetland vegetation within an upland forest landscape and compare the diversity and composition of different wetland plant communities. In an old-growth forest reserve in southern Quebec, Canada, we sampled wet habitats including lakeshores, permanent and seasonal ponds, swamps, glades, and streamsides. We used clustering, indicator species analysis, and nonmetric multidimensional scaling ordination to identify and compare vegetation types. The wetlands contained 280 species of vascular plants, 45% of the reserve's flora, in only 1.1% of its area. Local diversity averaged 24 ± 0.7 species per 7 m(2), much higher than in the surrounding upland forests. Plant communities sorted into five types, whose strongest indicator species were Osmunda regalis, Glyceria striata, O. cinnamomea, Deparia acrostichoides, and Matteuccia struthiopteris, respectively. Both local species richness and compositional variation among sites differed among the vegetation types. By combining species representative of the region's major wetlands with species from the upland forest matrix, the plant assemblages of these wetlands make disproportionately important contributions to landscape-level diversity.

Agricultural legacies in forest environments: tree communities, soil properties, and light availability.

Temperate deciduous forests across much of Europe and eastern North America reflect legacies of past land use, particularly in the diversity and composition of plant communities. Intense disturbances, such as clearing forests for agriculture, may cause persistent environmental changes that continue to shape vegetation patterns as landscapes recover. We assessed the long-term consequences of agriculture for environmental conditions in central New York forests, including tree community structure and composition, soil physical and chemical properties, and light availability. To isolate the effects of agriculture, we compared 20 adjacent pairs of forests that were never cleared for agriculture (primary forests) and forests that established 85-100 years ago on plowed fields (secondary forests). Tree communities in primary and secondary forests had similar stem density, though secondary forests had 14% greater basal area. Species composition differed dramatically between the two forest types, with primary forests dominated by Acer saccharum and Fagus grandifolia and secondary forests by Acer rubrum and Pinus strobus. Primary and secondary forests showed no consistent differences in soil physical properties or in the principal gradient of soil fertility associated with soil pH. Within stands, however, soil water content and pH were more variable in primary forests. Secondary forest soils had 15% less organic matter, 16% less total carbon, and 29% less extractable phosphorus in the top 10 cm than adjacent primary stands, though the ranges of the forest types mostly overlapped. Understory light availability in primary and secondary forests was similar. These results suggest that, within 100 years, post-agricultural stands have recovered conditions comparable to less disturbed forests in many attributes, including tree size and number, soil physical properties, soil chemical properties associated with pH, and understory light availability. The principal legacies of agriculture that remain in these forests are the reduced levels of soil organic matter, carbon, and phosphorus; the spatial homogenization of soil properties; and the altered species composition of the vegetation.

Microsite-limited recruitment controls fern colonization of post-agricultural forests.

Assessing the relative roles of dispersal limitation and environmental effects in population dynamics and community assembly is fundamental to understanding patterns of species distribution and diversity. In forests growing on abandoned agricultural lands, both legacies of vegetation disturbance and changes in the abiotic environment shape the diversity and composition of recovering communities. Here I specify how interactions among historical, environmental, and biological factors influence species distributions, focusing on three fern species with contrasting distributions across forests of different history in central New York, USA: Dryopteris carthusiana, Dryopteris intermedia, and Polystichum acrostichoides. Using population surveys, spore-trap and spore-bank studies, and a three-year field experiment, I compare demographic rates among species and between forest types to determine which life history stages limit colonization and which traits explain species distributions. Adult plants of all three species were larger and more likely to produce spores in post-agricultural forests than in adjacent, uncleared stands. Though lower population densities led to fewer spores in post-agricultural soils, spore availability still exceeded recruitment by four to five orders of magnitude. Sowing additional spores had relatively little effect, while microhabitat conditions had the greatest impact on establishment rates. Given similar microsites, the two forest types had equal rates of establishment, but some forest-floor features preferentially occupied by juvenile plants were less frequent in post-agricultural stands. The availability of suitable sites for establishment, created by small-scale heterogeneity on forest floors, thus limits both the growth of fern populations and the colonization of new habitats. In fact, reduced microtopographic variation in post-agricultural forests may represent a greater hindrance to plant establishment than changes in mean environmental conditions. Among the three fern species, establishment rates differed as species distributions would predict, with the strongest colonizer consistently having the highest rates and the slowest colonizer the lowest. Rather than random or trait-mediated dispersal, the different distributions of these species reflect life history traits that determine establishment rates and thus colonization ability. This case study demonstrates that ecological interactions based on the unique life histories of individual species can override dispersal in determining species distributions.

Reproductive biology of three fern species may contribute to differential colonization success in post-agricultural forests.

Because selfing enables a single individual to reproduce in a new location, the ability to self-fertilize should enhance plants' capacity for colonization. This study examined whether selfing ability correlated with successful migration in three fern species, Dryopteris carthusiana, Dryopteris intermedia, and Polystichum acrostichoides, which vary in their ability to colonize forests on abandoned agricultural lands in central New York, USA. Polystichum acrostichoides is much more frequent in forests that were never cleared for agriculture, D. carthusiana is more frequent in forests that developed on former fields, and D. intermedia is equally frequent in the two forest types. To test the hypothesis that better-colonizing species and post-agricultural forest populations have greater selfing ability, I assessed the sporophyte production of gametophytes grown in isolation and in pairs of varying relatedness. Dryopteris carthusiana had the highest reproductive success and selfing ability and P. acrostichoides the lowest. These results support the hypothesis that selfing may facilitate colonization in these species. They also exemplify the general pattern that polyploid fern species have higher rates of self-fertilization than related diploids, as the allotetraploid D. carthusiana had greater selfing ability than both diploid species.