Summary1. Metacommunity theory predicts that increasing patch size and patch connectivity can alter local species diversity by affecting either colonization rates, extinction rates or both. Although species' dispersal abilities or 'dispersal mode' (e.g. gravity-, wind-or animal-dispersed seeds) can mediate the effects of patch size and connectivity on diversity, these important factors are frequently overlooked in empirical metacommunity work. 2. We use a natural metacommunity of aspen stands within a grassland matrix to determine whether dispersal mode alters the influence of stand size and connectivity on understorey plant diversity. We sampled the same area in each patch, controlled for the presence of matrix species in aspen stands, and tested for the effects of size, connectivity and dispersal mode on metacommunity richness. Because dispersal groups responded differently to patch size and connectivity, we created a null model and assessed ungulate activity to explore whether competitive dynamics or herbivory were driving diversity patterns. 3. Animal-dispersed species and species with no dispersal aid had higher diversity per unit area in larger stands, likely because large stands can both support larger populations that are less prone to extinction and may also attract seed-dispersing animals such as birds and small mammals that are sensitive to edge effects. Consistent with other empirical work, we found a positive relationship between diversity and connectivity for wind-dispersed species. However, we detected a negative effect of stand connectivity on the diversity of species with no dispersal aid, possibly due to the presence of other highly competitive species groups dominating well-connected patches, as our null model results suggest. We found no evidence for higher ungulate activity in highly connected patches, suggesting that herbivory may not be driving the decline in diversity of plants with no dispersal aid. 4. Synthesis. Overall, we see a positive effect of stand area on diversity for most groups despite sampling equal area in all stands, which is a prediction of metacommunity theory that is normally overlooked. Our results demonstrate the importance of considering variation in the dispersal modes of focal species for explaining the diversity patterns of natural metacommunities.
The species richness, community composition, and abundance of bryophytes within taxonomic and functional groups were examined in relation to habitat conditions in forest edge and interior habitats of nine old-growth temperate rain-forest patches remaining after logging in the Nimpkish River Valley of Vancouver Island, British Columbia. Bryophytes were sampled at a fine scale using 0.1 m × 0.3 m microplots to examine responses of species abundance on the forest floor, downed logs, and tree bases and at a coarser scale using 10 m × 2 m belt transects to determine changes in patterns of species richness and distribution. Edge habitats, sampled to a depth of 45 m into the forest fragments, were characterized by greater windthrow disturbance. Within the edge zone, increases in the richness of clearing-affiliated functional groups were associated primarily with the location of windthrown trees and tip-up mounds, rather than with distance from the edge per se. Interior habitats had both greater abundance of old-growth-associated functional groups and total bryophyte cover. The extension of the edge zone to at least 45 m into remnant patches carries implications for minimum patch size requirements in the context of variable-retention logging of coastal temperate rain forests.
The resilience (measured as changes in functional group representation and species composition) of bryophyte communities found in the younger-aged (“matrix”) forests surrounding old-growth remnants was examined in two different forest types, warmer, drier (Nimpkish) versus cooler, wetter (Sayward), on Vancouver Island, British Columbia. Bryophytes were sampled within 10 m × 2 m belt transects (species composition only) and using 10 cm × 30 cm microplots (composition and abundance) in two age classes of matrix forest, clearcuts (age 7–20 years), and second-growth (age 25–49 years) as well as in remnant old-growth forest stands (age >300 years). The cover of all bryophytes was diminished and more patchily distributed in younger-aged stands; however, the richness and frequency of bryophyte functional groups showed different responses in the two younger age class forests. Disturbance-associated species exhibited both higher richness and frequency in clearcut plots and higher richness in second-growth plots. In comparison, the richness of species associated with old-growth was largely unchanged in younger-aged forests compared with old-growth forests; however, the frequency of occurrence of species associated with old-growth was significantly reduced in younger-aged forests. The cooler, wetter forests exhibited greater resilience, as the difference in species composition between second-growth and old-growth stands was less than that between second-growth and old-growth stands in the warmer, drier forests. The greater difference in second-growth species composition in the warmer, drier forests was attributed mainly to the persistence of disturbance-associated species.
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