Seedling establishment patterns on the Pumice Plain,Mount St. Helens,Washington

Abstract. We examined the factors that control seedling establishment on barren substrates on the pyroclastic flows from Mount St. Helens. From June to September in 1993, we monitored seedling and microhabitat changes in 240 20 cm × 20 cm quadrats on the Pumice Plain. Seedlings emerged in 104 quadrats (43.3 %). The most abundant species were Anaphalis margaritacea, Hypochaeris radicata, Lupinus lepidus and Epilobium angustifolium. Measured site characteristics included topography, particle size distribution, ground surface movements, soil water content, organic matter, pH, and presence or absence of dead lupines. Quadrats with seedlings had higher cover of dead lupines, higher amount of rock and gravel substrate, and a greater cover of rills. More seedlings emerged where eroded material accumulated. Compared to coarse-textured surfaces, silt surfaces had higher organic matter, held more water, and showed higher pH. However, seedlings became established more frequently on coarse-textured surfaces. In greenhouse experiments, a higher percentage of Hypochaeris seeds germinated on silt than on sand or gravel. The germination of Anaphalis and Epilobium did not differ with soil texture, but was higher at higher moisture levels. Seedling colonization is more dependent on ground surface microtopography, particle size, and ground movement than on the chemical status of these volcanic deposits.     

The role of refugia and dispersal in primary succession on Mount St. Helens,Washington

Abstract. An intense lateral blast devastated Mount St. Helens in 1980, but forest understory species survived in some north‐slope ‘refugia’. We explored the effects of refugia on colonization of barren pumice in 1997 and 1998, 18 yr after the eruption. The seed rain of 23 colonizers came mostly from populations that had previously established in refugia. Parachutists had small, vagile seeds, parasailors had winged seeds, and tumblers were blown along the ground. The latter two groups are heavier and dispersed more slowly, but are more likely to survive. The proportion of the vegetation represented by wind‐dispersed species increased with distance from refugia. Parachutist's density declined with time and proximity to refugia. As vegetation adjacent to refugia developed, populations of parasailors and tumblers expanded, foreshadowing their dominance in more remote pumice. Refugia played a critical role in determining the rate and course of succession by providing fertile islands that permitted pioneers and dry meadow species to establish near barren pumice. Species that survived in refugia played a negligible role in colonization. This study showed that when refugia contrast sharply with new substrates, they accelerate recovery by facilitating the invasion of pioneer species.     

Early succession on lahars spawned by Mount St. Helens

The effects of isolation on primary succession are poorly documented. I monitored vegetation recovery on two Mount St. Helens lahars (mud flows) with different degrees of isolation using contiguous plots. Seventeen years after the eruption, species richness was stable, but cover continued to increase. That isolation affects community structure was confirmed in several ways. The dominance hierarchies of the lahars differed sharply. Detrended correspondence analysis on Lahar I showed a trend related to distance from an adjacent woodland, whereas vegetation on Lahar II was relatively homogeneous. Spectra of growth forms and dispersal types also differed. Lahar I was dominated by species with modest dispersal ability, while Lahar II was dominated by species with better dispersal. Variation between plots should decline through time, a prediction confirmed on Lahar II. Lahar I remained heterogeneous despite having developed significantly higher cover. Here, the increasing distance from the forest has prevented plots from becoming more homogeneous. At this stage of early primary succession, neither lahar is converging towards the species composition of adjacent vegetation. This study shows that isolation and differential dispersal ability combine to determine initial vegetation structure. Stochastic effects resulting from dispersal limitations may resist the more deterministic effects of competition that could lead to floristic convergence.     

Early primary succession on Mount St. Helens,Washington, USA

Abstract. The north slope of Mount St. Helens was sampled with 141 circular 100-m² plots to describe vegetation and environmental patterns 13 yr after the 1980 eruption. At least 114 vascular plant taxa were encountered. We recognized four habitat types: Refugia, Pumice barrens, Pyroclastic surfaces and Drainages. A fifth category, Lupine patches, includes samples on primary surfaces that were rapidly colonized. Refugia provided small enclaves where underground portions of several species survived the eruption. They retained an inconsistent array of forest understory species and contained 86 species (mean = 20.8 per plot). Refugia are dominated by woody species such as Penstemon cardwellii, Rubus spp., Ribes spp. and Alnus sinuata, with herbs such as Agrostis diegoensis, Luzula parviflora and Anaphalis margaritacea. Anaphalis represents a suite of species that invaded Refugia after the eruption. Diversity (N₂ and H′) is significantly greater in Refugia than in any other habitat. No plants survived on primary sites, which remain sparsely vegetated and dominated by readily dispersed taxa. Total richness ranges from 36 species (9.9 per plot) on pyroclastic surfaces, through 42 species (11.2 per plot) in drainages, to 66 (11.7 per plot) on Pumice barrens. H′ and N₂ of the three habitats do not differ significantly. Lupine-dominated vegetation occurs sporadically in Pyroclastic and drainage habitats. Lupine patches are characterized by high Lupinus cover and a suite of invaders. These sites have high cover and 52 species (12.6 per plot). H′ and N₂ scores were significantly lower than any other habitat due to strong lupine dominance. Canonical Correspondence Analysis showed that site history and slope contributed most to species composition. Geographic effects accounted for 10 25 % of the explained species-environment relationship. Forest understory species have migrated only short distances and have made negligible contributions to vegetation development. A few species common in Refugia, including Agrostis diegoensis and Carex mertensii, have invaded barren surfaces, but most have not. Refugia also have been invaded by open site species abundant on the Pumice Plain. The heterogeneity of plots within habitat types and small statistical linkage of vegetation to environmental and spatial factors suggests that stochastic events have played a leading role in early primary succession.     

Increasing deterministic control of primary succession on Mount St. Helens,Washington

Question: Does the relative importance of stochastic and deterministic factors change during primary succession? Location: Small depressions (potholes) located on Mount St. Helens, Washington (46°13′51″N, 122°09′10″W, 1290 m). Methods: Pothole vegetation was described in 1993, 1997 and from 2001 to 2008. Explanatory variables included location and elevation (spatial factors), soil factors and Lupinus lepidus cover from prior years (a fertility surrogate). RDA assessed species‐variable relationships. DCA calculated β diversity and within‐year heterogeneity. Flexible sorting classified the vegetation. Species composition, richness, cover, H′ and evenness were also calculated. Results: Vegetation cover increased through 2001, and then fluctuated due to changes in L. lepidus cover. Richness peaked in 2005, after which pioneer species began to decline as persistent evergreen species increased. The six CTs recognized in 2008 were more scattered than were the six different CTs from 2001. DCA demonstrated that woody and rhizomatous species increased as pothole vegetation became less variable. RDA revealed weak spatial relationships in 1993, 1997 and 2001; thereafter, environmental and biological factors became important. The species‐explanatory data relationship increased during this study from 10.2% to 36.0%, leaving 64.0% of the variation unexplained. Conclusions: This is the first temporal study to demonstrate that deterministic control of vegetation development increases during succession. Pothole vegetation has converged somewhat due to deterministic factors, but the initial effects of chance, local disturbances and history remain large and may prevent strong convergence.     

Long-term Effects of <Emphasis Type="Italic">Lupinus lepidus</Emphasis> on Vegetation Dynamics at Mount St. Helens

The nitrogen-fixing legume Lupinus lepidus is the most abundant herb on new volcanic surfaces at Mount St. Helen. We compared vegetation structure in 30 Lupinus colonies in three age classes (old, mature, or young based on known years of their establishment) to adjacent sites that were sparsely populated by Lupinus. Our goals were to determine if the age of colonies affected either species composition or vegetation structure and if colonies altered vegetation structure of a site. Species richness increased with site age and colonies had significantly greater richness. Mature colonies had lower cover and frequency of other species, while old colonies had significantly higher cover of other species, particularly mosses. Frequencies in colonies were higher than in adjacent sites. Diversity measures were highest in old colonies and least in mature ones. ANOVA showed that density effects of Lupinus were stronger than age effects on richness, cover, frequency, dominance, and diversity. The floristic similarity between colonies and adjacent sites declined with colony age, and sites without Lupinus became heterogeneous with age. Detrended correspondence analysis demonstrated floristic change as colonies aged and that the differences between colonies and sparse sites increased with age. Regressions of Lupinus frequency with measures of structure showed weak second order relationships that suggested that low densities of Lupinus inhibited other species, while higher densities promoted abundance and diversity. Together these results suggest that Lupinus is a key element that alters floristic successional trajectories and accelerates succession. However, its effects are complicated by the wide ranges of Lupinus density, the sequence of invasion, and by herbivores.     

Species attributes in early primary succession on volcanoes

Abstract. We predicted that plants that can establish on volcanic soils with similar disturbance histories will have similar growth characteristics. We tested this prediction by a multivariate analysis of 27 traits of 84 species found six years after an eruption on Mount St. Helens, Washington State, USA, and Mount Usu, Hokkaido, Japan. These traits include vegetative, life-history, phenological and seed-biology characteristics. Cluster analysis revealed five species groups: annual herbs, perennial forbs, graminoids, shrubs and trees. Each group has distinct vegetative, life-history, and seed-biology traits. Except for shrubs, which were lacking on Mount Usu, both floras were well represented in each group. On intensely disturbed sites on both volcanoes, perennial forbs, whose development is dependent primarily on well-developed below-ground organs and wind-dispersal, expanded their cover more rapidly than did graminoids. These graminoids generally produce gravity-dispersed seeds and have close-set rhizomes and/or shoots. These results suggest that species that can establish during the early stages of succession on each volcano have similar vegetative, life-history, and seed-biology traits.     

Structural segregation and scales of spatial dependency in Abies amabilis forests

Question: Is a mosaic structure apparent in the spatial distribution of trees in old‐growth Abies amabilis forests? Location: Montane forests of the western Cascade Range, Washington, USA. Methods: Maps of tree locations were created for study areas located in two, 300‐year old stands and a single 600‐year old stand. Stand structure parameters were calculated using several subsample quadrats sizes (56.25 ‐ 306.25 m²), which were drawn randomly with replacement at a density of 250 quadrats per ha from the stem maps in the computing environment. Spatial cross‐covariance functions between different canopy strata were estimated using the spline cross‐correlogram. Results: Negative spatial correlation (segregation) between subcanopy tree density and areas of high overstorey occupancy was detected. Understorey and midstorey tree densities were positively spatially correlated. These general trends were apparent across the range of observational scales investigated. Significant spatial correlation between canopy strata was observed at spatial scales of 12 ‐ 44 m and extended to the largest scales in the 600‐year old stand. Conclusion: The observed spatial segregation between canopy strata supports the hypothesis that old A. amabilis forests form fine‐scale structural mosaics. Structural segregation at small scales may be due to competitive interactions as well as exogenous forcing of tree locations (e.g. by mortality due to pathogens or disturbance), however segregation at large scales in the 600‐year old stand is likely due to exogenous factors alone. This study reinforces the idea that horizontal heterogeneity is an emergent property of old‐growth forests.     

Early primary succession on a barren volcanic plain at Mount St. Helens,Washington

The invasion pattern on a barren plain on the eastern flank of Mount St. Helens, Washington, devastated by the 1980 volcanic eruption was monitored between 1988 and 1992. All vascular plants on a grid consisting of 400 contiguous 100-square-meter quadrats were recorded with a cover score. The substrate was initially homogeneous, but significant heterogeneity had developed by 1988. Vascular plant species richness increased from 24 in 1988 to 41 in 1992. Mean species richness per quadrat increased from 0.44 to 5.71, mean cover increased from 0.04% to 0.51%, and mean diversity index (H') increased from 0.08 to 1.56. A variance/mean test of species richness pattern showed that invasion occurred sporadically since plots tended to have either several or no species. By 1992, mean species richness was more evenly distributed. Most seedlings continue to result from long-distance dispersal, but reproductive colonies of species are developing. Seedling distributions are controlled by microsites. Eleven common species strongly and similarly preferred safe-sites created by small rocks, undulations, or rills. However, many apparent safe-sites are empty, suggesting that seeds are scarce and that even the most favorable microsites are marginal. The niches of these species seem to overlap broadly. The Plains of Abraham is in the earliest stage of primary succession. The detailed invasion pattern permitted us to distinguish species still dependent on immigration from those now locally established. Pronounced microsite preferences emphasize that physical amelioration (e.g., nutrient input and erosion) must occur before further succession can commence. We have observed the early stages of succession where an inhospitable site is gradually and heterogeneously changed into a habitat where safe-sites do not limit succession, but where stochastic factors remain important.     

Tree invasion into a mountain-top meadow in the Oregon Coast Range,USA

Abstract. The grassy meadow on the top of Marys Peak, in the Oregon Coast Range, is being invaded, primarily along the margins, by Abies procera. To examine vegetation and environmental changes across the forest-meadow transition and to evaluate factors affecting tree invasion, belt transects were established at 10 sites. Correlations of environmental variables with ordination axes from Detrended Correspondence Analysis suggest that the sites were distributed along a moisture gradient. Although the sites varied considerably in species composition and environment, tree invasion of the meadow was occurring at all sites. Reduction in abundance of the dense herbaceous vegetation of the meadow is required before the small seedlings of Abies procera can become established. Almost all tree invasion of the meadow occurs in the narrow forest-meadow ecocline because trees at the edge of the forest reduce the cover of herbaceous plants in the adjacent meadow. On some sites, above average snow accumulations further reduce vigor of meadow vegetation and increase tree establishment. On the driest sites wet summers facilitate tree establishment. Infrequent fires, which remove the fire sensitive Abies procera, are probably required to reverse the slow but persistent trend of forest expansion and ultimately maintain the meadow.     

Dynamics of herbaceous vegetation recovery on Mount St. Helens,Washington, USA,after a volcanic eruption

Recovery of herbaceous vegetation on Mount St. Helens was studied annually after the massive lateral eruption of May 18, 1980. Measures such as species richness, cover, and diversity were combined with detrended correspondence analysis to describe vegetation recovery rates under different combinations of initial impact intensity and degree of isolation from recolonization sources. A major key to recovery is whether any plants survived the devastation. Survival of even a few individuals markedly accelerated recovery. Where no plants survived, the degree of isolation becomes paramount. New, barren substrates, a few meters from undisturbed sites, have begun to develop some vegetation, while more isolated sites have scarcely any subalpine plants present. On any site, plant-mediated processes that improve conditions for growth and the invasion of other species predominate in the early stages, but as vegetation develops, biotic inhibition and establishment of seedlings from adults already in the habitat gain importance. The rate at which this conversion occurs is a function of the size and intensity of the initial impact.Abbreviations DCA Detrended correspondence analysis     

Primary succession trajectories on a barren plain,Mount St. Helens,Washington

Questions: Have predictable relationships between environmental variables and vegetation developed in primary succession following a volcanic eruption? Has the rate of succession changed? Have vegetation trajectories converged or diverged? Location: The Abraham Plain of Mount St. Helens, Washington, USA (46°12′42″N, 122°08′27″W, elevation 1360 m), was sterilized in 1980 by a blast, scoured by lahars and buried by pumice. Method: We monitored 400 100 m² contiguous permanent plots annually (1988–2008), and classified each plot from every year into ten community types (CTs). We characterized the terrain by topography and surface features. Redundancy analysis assessed relationships between vegetation and possible explanatory variables, which included sample location. We used detrended correspondence analysis (DCA) to assess successional rates and trends. Results: Relationships between species composition and explanatory variables were only significant after 1996, when position and presence of rills became significant. By 2006, explained variation remained low (13%) but significant. Species accumulated slowly, restricted by stress and isolation. Changes in mean DCA position slowed. Composition shifted from pioneer to persistent species and vegetation became more stable with time. Species accumulated for two decades and then stabilized, while cover has continued to increase. Diversity increased and then declined slightly as dominance developed and pioneer species became less common. Conclusions: We demonstrate weak but increasingly predictable trends in species composition using environmental variables. The rate of succession slowed and trajectories formed a reticulate network of transitions dominated by divergence. Convergence was not evident because vegetation responded distinctively to minor topographic features that allowed alternative stable communities to develop.     

N-P co-limitation of primary production and response of arthropods to N and P in early primary succession on Mount St. Helens volcano

Background

The effect of low nutrient availability on plant-consumer interactions during early succession is poorly understood. The low productivity and complexity of primary successional communities are expected to limit diversity and abundance of arthropods, but few studies have examined arthropod responses to enhanced nutrient supply in this context. We investigated the effects of nitrogen (N) and phosphorus (P) addition on plant productivity and arthropod abundance on 24-yr-old soils at Mount St. Helens volcano.

Methodology/Principal Findings

We measured the relative abundance of eight arthropod orders and five families in plots that received N, P, or no nutrients for 3–5 years. We also measured plant % cover, leaf %N, and plant diversity. Vegetation responded rapidly to N addition but showed a lagged response to P that, combined with evidence of increased N fixation, suggested P-limitation to N availability. After 3 yrs of fertilization, orthopterans (primarily Anabrus simplex (Tettigoniidae) and Melanoplus spp (Acrididae)) showed a striking attraction to P addition plots, while no other taxa responded to fertilization. After 5 yrs of fertilization, orthopteran density in the same plots increased 80%–130% with P addition and 40% with N. Using structural equation modeling, we show that in year 3 orthopteran abundance was associated with a P-mediated increase in plant cover (or correlated increases in resource quality), whereas in year 5 orthopteran density was not related to cover, diversity or plant %N, but rather to unmeasured effects of P, such as its influence on other aspects of resource quality.

Conclusions/Significance

The marked surprising response to P by orthopterans, combined with a previous observation of P-limitation in lepidopteran herbivores at these sites, suggests that P-mediated effects of food quantity or quality are critical to insect herbivores in this N-P co-limited primary successional system. Our results also support a previous suggestion that the availability of N in these soils is P-limited.     

Forest dynamics,differential mortality and variable recruitment probabilities

Abstract. Both spatial and temporal variability in recruitment probabilities can lead to coexistence in gap-phase regenerating forests which would otherwise tend to be dominated by fewer species. Using modified Markov models, the potential roles were examined of temporal variability and differential mortality rates among species in the dynamics of a forest for which spatial variability has been rejected as a strong factor leading to coexistence. Differential longevity modifies results obtained from a simple Markov model: it exerts a strong influence on the equilibrium species composition, on the rate of community change and on the time a community requires to reach equilibrium. Simulations with varying transition probabilities mimicked a changing climate, producing four main results: 1. Unless the duration of climate states is very long or very short, forest composition is in a continual state of disequilibrium. 2. Species vary in their response times to changing climate. 3. The mean abundance of each species under a varying climate scenario is different from that expected from the mean climate state. 4. The rare, long-lived species was favored by climatic fluctuations at the expense of more common shorter lived species. Differential mortality rates provide an equilibrium-based mechanism for coexistence, and temporally fluctuating recruitment probabilities a non-equilibrium mechanism. Composition could be maintained by differential longevity among species and climatic fluctuations allowing periodic recruitment of the less common species.     

The role of mycorrhizal fungi and microsites in primary succession on Mount St. Helens

This study was designed to examine the role of vesicular-arbuscular mycorrhizae (VAM) and microsites on the growth of pioneer species. Flat, rill, near-rock, and dead lupine microsites were created in plots in barren areas of the Pumice Plain of Mount St. Helens. VAM propagules were added to the soil in half of the plots. Six pioneer species were planted into both VAM and non-VAM inoculated microsites. Plants in dead lupine microsites were greater in biomass than those in flat, rill, and near-rock microsites. Significant effects of VAM on plant biomass did not occur. Microsites continue to be important to plant colonization on the Pumice Plain, but VAM do not yet appear to play an important role. This may be due to limited nutrient availability and the facultatively mycotrophic nature of the colonizing plant species. It is unlikely that VAM play an important role in successional processes in newly emplaced nutrient-poor surfaces.     

Spatial factors affecting primary succession on the Muddy River Lahar,Mount St. Helens,Washington

Primary succession is controlled by a combination of landscape and habitat factors whose actions may be stochastic or deterministic. The 1980 eruption of Mount St. Helens, Washington spawned a massive lahar that now supports a mosaic of vegetation. Our goals were to describe vegetation patterns after 28 growing seasons, determine the factors associated with these patterns, and to contrast the effects of stochastic and deterministic processes. We described species composition and explanatory factors that included location and habitat features in one hundred and fifty one 200-m² plots. We classified these plots into nine community types (CTs) that were distinguished quantitatively by variations in dominant species. We used multiple regressions, redundancy analysis (RDA), and Mantel tests to compare the vegetation relationships with explanatory factors. Plots in different CTs mingled spatially and in multivariate space. Species patterns were weakly related to explanatory variables by RDA (31.6% of the species variation). RDA indicated that vegetation was most strongly related to elevation, latitude, and isolation, which are primarily landscape factors. Mantel tests confirmed that factors associated with elevation were most closely associated with vegetation. The effects of arrival order were suggested by the dominance of different colonizers in similar environment and by plots with similar vegetation found in different habitats. We concluded that species composition cannot be predicted well from the data available, suggesting that there were no prominent deterministic assembly rules.     

Vegetation development on pumice at Mount St. Helens, USA

This study explores early vegetation development on pumice at Mount St.Helens. We monitored species annually in a grid of 200 contiguous100-m² plots between 1989 and 1999. Of interestwere how vegetation changed and if it became more homogeneous over time.Speciesrichness and cover increased annually. Diversity(H) stabilized by 1996 and began to decline aslong-livedstress-tolerant species such as Agrostis pallens, Carexmertensii and Penstemon cardwellii began todominate. Protected sites had more species and higher cover than did exposedones. Plots next to relict vegetation had more species and cover than diddistant plots. The vegetation initially was dominated by species with gooddispersal, but subsequently those with poor dispersal became dominant. Wecompared species expansion patterns to a model based on random colonization.Theresults implied that populations with poor dispersal derived from a fewcolonists that then produced seeds for local expansion. Detrendedcorrespondenceanalysis showed a pronounced shift in species composition. This analysis alsoshowed that species composition was becoming more homogeneous over time.However, significant heterogeneity remained and some plots are diverging fromothers. As yet, this vegetation is not developing towards a regional vegetationtype. Rather, an unusual community with Agrostis spp.,Carex spp., Penstemon cardwellii, Lupinuslepidus, Anaphalis margaritacea and Salixcommutata has developed. The accumulation phase of primarysuccessionis nearly complete. The next phase, in which vertical structure develops asSalix and conifers mature, has scarcely begun. It shouldbemarked by the invasion of forest understory species and loss of subalpinemeadowspecies. Assembly rules based on biotic interactions may then become evident.     

Gradients in compositional variation on lahars,Mount St. Helens,Washington, USA

How isolation affects primary succession remains unresolved. Our hypotheses are: 1) the rate of succession is slowed within 50 m from sources and 2) dominance increases more rapidly if the growing season is longer. We sampled lahar vegetation near conifer forests on Mount St. Helens 23 years after the eruption using transects (Muddy) and grids (Butte Camp, BC). Transects were compared to isolated plots of the same age, while grids were compared to vegetation that was 400 years older. Cover declined with distance and with elevation on the Muddy due to denser seed rain near forests and shorter growing seasons at higher elevation. On BC-1, next to a forest, richness measures increased with distance, while cover decreased with elevation. On BC-2, more exposed than BC-1, mean richness and cover declined with elevation, but increased towards seed sources. Equitability increased with distance and with elevation on more stressful transects and on BC-1. Percent similarity (PS), a measure of floristic heterogeneity, declined with elevation on Muddy-SW. BC-1 PS decreased with both distance and elevation. Data from grids collected since 1987 showed that both sites became more homogeneous through time. Directional changes on BC-1 were greater than on BC-2, while annual variation of DCA scores declined only on BC-2. These differences are related to conifer dominance patterns. Succession is influenced by isolation, which controls the seed rain, and by stress, which controls establishment, the rate of biomass accumulation, and the spread of immigrants. Although community development is governed by environmental factors, this study shows that the effects of dispersal that result from distance factors can persist and may be a source of unexplained variation in mature communities.     

A multi-scale assessment of the occurrence of exotic plants on the Olympic Peninsula,Washington

Abstract. Exotic plants were surveyed in 208 plots within the Dungeness and Hoh river watersheds on the Olympic Peninsula, Washington, USA. Landscape patch types included uplands (clearcuts, young and mature forests) and riparian zones (cobble bars, shrub patches, riparian forests, and alder flats). Patterns of exotic plant invasions were assessed between watersheds, between riparian and upland areas, among patch types, and within clearcuts. 52 exotic plant species were encountered, accounting for 23% of the flora in each watershed. In both watersheds, exotic species richness was approximately 33% greater in riparian zones than in uplands, and mean number and cover of exotic species were > 50% greater in riparian zones than in uplands. Among landscape patch types, richness and mean number and cover of exotics was highest in young riparian patches, intermediate in clearcuts and riparian forests, and lowest in young and mature forests. The exception to this was Hoh alder flats, which had the highest mean cover of exotic plants. Cover of exotic plants peaked in uplands 3 to 7 yr after clearcutting, then decreased with increased canopy closure. Disturbance type and time since disturbance were major factors influencing invasibility. Landscape patch size, position within watershed (distance from patch to human population centers, major highway, or river mouth), and environmental variables (slope, aspect, and elevation) were not important indicators of landscape patch invasibility. Riparian zones facilitated movement of exotic plants through landscapes, but did not appear to act as sources of exotic plants for undisturbed upland areas.