Phenology of the dioecious shrub Baccharis halimifolia along a salinity gradient: Consequences for the invasion of Atlantic subhalophilous communities

Atlantic subhalophilous communities are habitats of high conservation value that are seriously threatened by plant invasions. The dioecious alien shrub Baccharis halimifolia L. (Asteraceae) is among the most aggressive invaders in estuarine communities in Europe, especially in sea rush communities of Juncus maritimus. We investigated the growth and flowering phenology of B. halimifolia through fine-scale environmental gradients, and we assessed the intersexual differentiated responses. We monitored 120 individuals of B. halimifolia growing in rush communities differing in salinity and waterlogging through one growing season in Urdaibai Biosphere Reserve. Ground water salinity and water table level were also recorded. Overall, B. halimifolia shoot growth occurred from May to December and flowering in late August, but males flowered earlier than females. However, we found high intrapopulation and intersexual variation in the phenology of B. halimifolia, related to fine-scale variations in edaphic conditions. Growth rate and leaf production were lower in high-salinity rush communities. Females and males showed different phenological responses to environmental conditions. Females appeared to be more sensitive to salinity than males, but males experienced higher levels of attack by natural enemies. The implications for the invasiveness and management of sea rush communities are discussed.     

Replacement of estuarine communities by an exotic shrub: distribution and invasion history of Baccharis halimifolia in Europe

Baccharis halimifolia L. (Asteraceae) is a shrub native to North America which is invading estuarine communities in Europe. We report the invasion history and the distribution limits of B. halimifolia in Europe, with particular emphasis on the frequency of its presence in estuarine communities in Spain. B. halimifolia has been cultivated in Europe since the 17th century to present. It was first recorded as naturalized in the Bay of Biscay in 1906, where it forms currently stable and locally abundant populations in almost all the estuaries of Northern Spain and Western France. The ongoing invasion to the west could reach well conserved estuarine communities in Galicia (Spain). B. halimifolia also forms scattered populations in Northern and Southern France, Belgium, Netherlands, United Kingdom and Italy. In these countries it has experienced a rapid expansion during the last years. In Northern Spain, subhalophilous communities dominated by rush (Juncus maritimus) and/or sea couch (Elytrigia atherica), common reed stands (Phragmites australis) and ungrazed wet meadows are the most vulnerable to invasion. The subhalophilous communities are part of natural habitats of community interest according to the habitats directive 92/43/EEC. In some areas of Northern Spain these communities have been totally replaced by monospecific stands of B. halimifolia. In contrast, halophilous communities of the low marsh are resistant to invasion, suggesting that the survival of B. halimifolia may be limited by threshold values of salinity and waterlogging. With this study we want to raise awareness about the risk of replacement of estuarine subhalophilous communities by the ongoing invasion of B. halimifolia in Europe.     

An unfortunate alliance: Native shrubs increase the abundance,performance, and apparent impacts of Bromus tectorum across a regional aridity gradient

Interspecific facilitation contributes to the assembly of desert plant communities. However, we know little of how desert communities invaded by exotic species respond to facilitation along regional-scale aridity gradients. These measures are essential for predicting how desert plant communities might respond to concomitant plant invasion and environmental change. Here, we evaluated the potential for Bromus tectorum (a dominant invasive plant species) and the broader herbaceous plant community to form positive associations with native shrubs along a substantial aridity gradient across the Great Basin, Mojave, and San Joaquin Deserts in North America. Along this gradient, we sampled metrics of abundance and performance for B. tectorum, all native herbaceous species combined, all exotic herbaceous species combined, and the total herbaceous community using 180 pairs of shrub and open microsites. Across the gradient, B. tectorum formed strong positive associations with native shrubs, achieving 1.6–2.2 times greater abundance, biomass, and reproductive output under native shrubs than away from shrubs, regardless of relative aridity. In contrast, the broader herbaceous community was not positively associated with native shrubs. Interestingly, increasing B. tectorum abundance corresponded to decreasing native abundance, native species richness, exotic species richness, and total species richness under but not away from shrubs. Taken together, these findings suggest that native shrubs have considerable potential to directly (by increasing abundance and performance) and indirectly (by increasing competitive effects on neighbors) facilitate B. tectorum invasion across a large portion of the non-native range.     

Impacts of a woody invader vary in different vegetation communities

The impact of an exotic species in natural systems may be dependent not only on invader attributes but also on characteristics of the invaded community. We examined impacts of the invader bitou bush, Chrysanthemoides monilifera ssp. rotundata , in fore and hind dune communities of coastal New South Wales, Australia. We compared invader impacts on vegetation structure, richness of both native and exotic growth forms and community variability in fore and hind dunes. We found that impacts of bitou invasion were context specific: in fore dune shrublands, functionally distinct graminoid, herb and climber rather than shrub growth forms had significantly reduced species richness following bitou invasion. However, in forested hind dunes, the functionally similar native shrub growth form had significantly reduced species richness following bitou invasion. Density of vegetation structure increased at the shrub level in both fore and hind dune invaded communities compared with non-invaded communities. Fore dune ground-level vegetation density declined at invaded sites compared with non-invaded sites, reflecting significant reductions in herb and graminoid species richness. Hind dune canopy-level vegetation density was reduced at invaded compared with non-invaded sites. Bitou bush invasion also affected fore dune community variability with significant increases in variability of species abundances observed in invaded compared with non-invaded sites. In contrast, variability among all hind dune sites was similar. The results suggest that effects of bitou bush invasion are mediated by the vegetation community. When bitou bush becomes abundant, community structure and functioning may be compromised.     

Grassland invaders and their mycorrhizal symbionts: a study across climate and invasion gradients

Controlled experiments show that arbuscular mycorrhizal fungi (AMF) can increase competitiveness of exotic plants, potentially increasing invasion success. We surveyed AMF abundance and community composition in Centaurea stoebe and Potentilla recta invasions in the western USA to assess whether patterns were consistent with mycorrhizal-mediated invasions. We asked whether (1) AMF abundance and community composition differ between native and exotic forbs, (2) associations between native plants and AMF shift with invading exotic plants, and (3) AMF abundance and/or community composition differ in areas where exotic plants are highly invasive and in areas where they are not. We collected soil and roots from invaded and native forb communities along invasion gradients and in regions with different invasion densities. We used AMF root colonization as a measure of AMF abundance and characterized AMF communities in roots using 454-sequencing of the LSU-rDNA region. All plants were highly colonized (>60%), but exotic forbs tended to be more colonized than natives (P < 0.001). We identified 30 AMF operational taxonomic units (OTUs) across sites, and community composition was best predicted by abiotic factors (soil texture, pH). Two OTUs in the genera Glomus and Rhizophagus dominated in most communities, and their dominance increased with invasion density (r = 0.57, P = 0.010), while overall OTU richness decreased with invasion density (r = −0.61, P = 0.006). Samples along P. recta invasion gradients revealed small and reciprocal shifts in AMF communities with >45% fungal OTUs shared between neighboring native and P. recta plants. Overall, we observed significant, but modest, differences in AMF colonization and communities between co-occurring exotic and native forbs and among exotic forbs across regions that differ in invasion pressure. While experimental manipulations are required to assess functional consequences, the observed patterns are not consistent with those expected from strong mycorrhizal-mediated invasions.     

Ecological filtering of exotic plants in an Australian sub‐alpine environment

We investigated some of the factors influencing exotic invasion of native sub‐alpine plant communities at a site in southeast Australia. Structure, floristic composition and invasibility of the plant communities and attributes of the invasive species were studied. To determine the plant characteristics correlated with invasiveness, we distinguished between roadside invaders, native community invaders and non‐invasive exotic species, and compared these groups across a range of traits including functional group, taxonomic affinity, life history, mating system and morphology. Poa grasslands and Eucalyptus‐Poa woodlands contained the largest number of exotic species, although all communities studied appeared resilient to invasion by most species. Most community invaders were broad‐leaved herbs while roadside invaders contained both herbs and a range of grass species. Over the entire study area the richness and cover of native and exotic herbaceous species were positively related, but exotic herbs were more negatively related to cover of specific functional groups (e.g. trees) than native herbs. Compared with the overall pool of exotic species, those capable of invading native plant communities were disproportionately polycarpic, Asteracean and cross‐pollinating. Our data support the hypothesis that strong ecological filtering of exotic species generates an exotic assemblage containing few dominant species and which functionally converges on the native assemblage. These findings contrast with those observed in the majority of invaded natural systems. We conclude that the invasion of closed sub‐alpine communities must be viewed in terms of the unique attributes of the invading species, the structure and composition of the invaded communities and the strong extrinsic physical and climatic factors typical of the sub‐alpine environment.     

Canopy effects of the invasive shrub Pyracantha angustifolia on seed bank composition,richness and density in a montane shrubland (Córdoba,Argentina)

Abstract Invasive woody species frequently change the composition of the established vegetation and the properties of the soil under their canopies. Accordingly, invasion may well affect regenerative phases of the community, especially at the seed bank level, likely influencing community restoration. Pyracantha angustifolia (Rosaceae) is an invasive shrub in central Argentina that affects woody recruitment, particularly enhancing the recruitment of other exotic woody species. There is though no information regarding its effect on the soil seed bank within the invaded community. The present study was set up to gain further insight into the canopy effects of P. angustifolia. We aimed to assess whether the invasive shrub affects seed bank composition, richness and seed density as compared with the dominant native shrub Condalia montana (Rhamnaceae), and to relate the observed seed bank patterns with those of the established vegetation. We evaluated the composition of the germinable seed bank and the established vegetation under the canopy of 16 shrubs of P. angustifolia, 16 shrubs of C. montana, and in 16 control plots (10 m²) without shrub cover. The floristic composition of the seed bank differed among canopy treatments. However, seed bank richness did not differ significantly. There was an overall high seed density of exotic species throughout the study site, though exotic forbs showed significantly lower seed densities under the invasive shrub. Pyracantha angustifolia would not promote the incorporation of new species into the seed bank of the invaded community but rather favour the establishment of woody species that do not depend on seed banks. The absence of dominant woody species in the seed bank, the dominance of exotic forbs, and the high similarity between established exotic species and those present in the seed bank may surely affect community restoration following the main disturbances events observed in the region.     

Invasion impacts local species turnover in a successional system

Exotic plant invasions are often associated with declines in diversity within invaded communities. However, few studies have examined the local community dynamics underlying these impacts. Changes in species richness associated with plant invasions must occur through local changes in extinction and/or colonization rates within the community. We used long‐term, permanent plot data to evaluate the impacts of the exotic vine Lonicera japonica. Over time, species richness declined with increasing L. japonica cover. L. japonica reduced local colonization rates but had no effect on extinction rates. Furthermore, we detected significant reductions in the immigration of individual species as invasion severity increased, showing that some species are more susceptible to invasion than others. These findings suggest that declines in species richness associated with L. japonica invasion resulted from effects on local colonization rates only and not through the competitive displacement of established species.     

Negative Effects of an Exotic Grass Invasion on Small-Mammal Communities

Exotic invasive species can directly and indirectly influence natural ecological communities. Cheatgrass (Bromus tectorum) is non-native to the western United States and has invaded large areas of the Great Basin. Changes to the structure and composition of plant communities invaded by cheatgrass likely have effects at higher trophic levels. As a keystone guild in North American deserts, granivorous small mammals drive and maintain plant diversity. Our objective was to assess potential effects of invasion by cheatgrass on small-mammal communities. We sampled small-mammal and plant communities at 70 sites (Great Basin, Utah). We assessed abundance and diversity of the small-mammal community, diversity of the plant community, and the percentage of cheatgrass cover and shrub species. Abundance and diversity of the small-mammal community decreased with increasing abundance of cheatgrass. Similarly, cover of cheatgrass remained a significant predictor of small-mammal abundance even after accounting for the loss of the shrub layer and plant diversity, suggesting that there are direct and indirect effects of cheatgrass. The change in the small-mammal communities associated with invasion of cheatgrass likely has effects through higher and lower trophic levels and has the potential to cause major changes in ecosystem structure and function.     

Giant invasive Heracleum persicum: Friend or foe of plant diversity?

The impact of invasion on diversity varies widely and remains elusive. Despite the considerable attempts to understand mechanisms of biological invasion, it is largely unknown whether some communities’ characteristics promote biological invasion, or whether some inherent characteristics of invaders enable them to invade other communities. Our aims were to assess the impact of one of the massive plant invaders of Scandinavia on vascular plant species diversity, disentangle attributes of invasible and noninvasible communities, and evaluate the relationship between invasibility and genetic diversity of a dominant invader. We studied 56 pairs of Heracleum persicum Desf. ex Fisch.‐invaded and noninvaded plots from 12 locations in northern Norway. There was lower native cover, evenness, taxonomic diversity, native biomass, and species richness in the invaded plots than in the noninvaded plots. The invaded plots had nearly two native species fewer than the noninvaded plots on average. Within the invaded plots, cover of H. persicum had a strong negative effect on the native cover, evenness, and native biomass, and a positive association with the height of the native plants. Plant communities containing only native species appeared more invasible than those that included exotic species, particularly H. persicum. Genetic diversity of H. persicum was positively correlated with invasibility but not with community diversity. The invasion of a plant community by H. persicum exerts consistent negative pressure on vascular plant diversity. The lack of positive correlation between impacts and genetic diversity of H. persicum indicates that even a small founder population may cause high impact. We highlight community stability or saturation as an important determinant of invasibility. While the invasion by H. persicum may decrease susceptibility of a plant community to further invasion, it severely reduces the abundance of native species and makes them more vulnerable to competitive exclusion.     

A role for the sampling effect in invaded ecosystems

Species loss and invasion of exotic species are two components of global biodiversity change that are expected to influence ecosystem functioning. Yet how they interact in natural settings remains unclear. Experiments have revealed two major mechanisms for the observed increase in primary productivity with plant species richness. Plant productivity may rise with species richness due to the increased amount of resources used by more diverse communities (niche complementarity) or through the increased probability of including a highly productive, dominant species in the community (sampling effect). Current evidence suggests that niche complementarity is the most relevant mechanism, whereas the sampling effect would only play a minor and transient role in natural systems. In turn, exotic species can invade by using untapped resources or because they possess a fitness advantage over resident species allowing them to dominate the community. We argue that the sampling effect can be a significant biodiversity mechanism in ecosystems invaded by dominant exotic species, and that the effect can be persistent even after decades of succession. We illustrate this idea by analyzing tree species richness–productivity relationships in a subtropical montane forest (NW Argentina) heavily invaded by Ligustrum lucidum, an evergreen tree from Asia. We found that the forest biomass increased along a natural gradient of tree species richness whether invaded by L. lucidum or not. Consistent with the sampling effect, L. lucidum invasion tripled total tree biomass irrespective of species richness, and monocultures of L. lucidum were more productive than any of the most species‐rich, uninvaded communities. Hence, the sampling effect may not be restricted to randomly assembled, synthetic communities. We emphasize that studying invaded ecosystems may provide novel insights on the mechanisms underlying the effect of biodiversity on ecosystem function.     

Exotic plant species of the St Lawrence River wetlands: a spatial and historical analysis

Aim To evaluate the importance (number of species, plant cover) of the exotic flora in seven well‐defined sectors of one of the most important transportation waterways in North America. To determine the impact of exotic species on wetland plant diversity and reconstruct the spread of some invasive species. Location St Lawrence River, southern Québec. Methods The exotic flora (vascular plants) of wetlands bordering the St Lawrence River was studied using 713 sampling stations (25 m²) along a 560‐km long corridor. Results Exotic species represent 13.7% of the vascular flora of the St Lawrence wetlands. The relative plant cover occupied by exotic species is high in some of the fluvial sectors (42–44%), but low (6–10%) in the estuarine sectors. Wetlands (marshes) surrounding islands were particularly susceptible to invasion by exotic plants. Historical, abiotic and landscape factors may explain the differences observed between sites. Purple loosestrife (Lythrum salicaria L.) is the most common exotic species of the St Lawrence wetlands, but other species, namely flowering‐rush (Butomus umbellatus L.) and reed canary grass (Phalaris arundinacea L.) are much more invasive. There is no linear relationship between the exotic species cover and the diversity of wetland plants; low diversity sites can be dominated by either exotic or native plant species. In the other sites, exotic species generally have little impact on plant communities and can contribute to increase diversity. Common reed (Phragmites australis (Cav.) Trin. ex Steudel) and reed canary grass, both considered as exotic species in this study, clearly have a stronger impact on plant diversity than flowering‐rush and purple loosestrife. Main conclusions This study shows that the global impact of an invader cannot be adequately evaluated with only a few highly invaded sites. While nationwide strategies have been developed to control exotic species, large surveys are essential to adapt them to regional particularities.     

Impact of exotic invasion on the temporal stability of natural annual plant communities

Much work in ecology has focused on understanding how changes in community diversity and composition will affect the temporal stability of communities (the degree of fluctuations in community abundance or biomass over time). While theory suggests diversity and dominant species can enhance temporal stability, empirical work has tended to focus on testing the effect of diversity, often using synthetic communities created with high species evenness. We use a complementary approach by studying the temporal stability of natural plant communities invaded by a dominant exotic, Erodium cicutarium. Invasion was associated with a significant decline in community diversity and change in the identity of the dominant species allowing us to evaluate predictions about how these changes might affect temporal stability. Community temporal stability was not correlated with community richness or diversity prior to invasion. Following invasion, community stability was again not correlated with community richness but was negatively correlated with community diversity. Before and after invasion, community stability was positively correlated with the stability of the most dominant species in the community, even though the identity of the dominant species changed from a native (prior to invasion) to an exotic species. Our results demonstrate that invasion by a dominant exotic species may reduce diversity without negatively affecting the temporal stability of natural communities. These findings add support to the idea that dominant species can strongly affect temporal stability, independent of community diversity.     

Exotic shrub invasion in an undisturbed wetland has little community-level effect over a 15-year period

In this long-term study, we examined the invasion by the exotic shrub glossy buckthorn (Rhamnus frangula L.) and the response of co-occurring plants in a large, undisturbed wetland. We first sampled the vegetation in 1991 and repeated the sample 15 years later using the same, permanently located sample units (n = 165). Despite dramatic increases in the abundance of buckthorn, the invasion elicited little apparent response by the resident plant community. Species richness and cover in the herbaceous plant stratum had no apparent relationship with change in buckthorn cover. The number of shrub species other than buckthorn showed no relationship with change in buckthorn cover, but the cover of other shrubs decreased as buckthorn cover increased. Species composition changed independently of changes in buckthorn cover. These results show that dramatic increases in the abundance of an invasive species do not necessarily cause large changes in the native plant community and suggest disturbance history influences community response to invasion.     

Shrubs as ecosystem engineers across an environmental gradient: effects on species richness and exotic plant invasion

Ecosystem-engineering plants modify the physical environment and can increase species diversity and exotic species invasion. At the individual level, the effects of ecosystem engineers on other plants often become more positive in stressful environments. In this study, we investigated whether the community-level effects of ecosystem engineers also become stronger in more stressful environments. Using comparative and experimental approaches, we assessed the ability of a native shrub (Ericameria ericoides) to act as an ecosystem engineer across a stress gradient in a coastal dune in northern California, USA. We found increased coarse organic matter and lower wind speeds within shrub patches. Growth of a dominant invasive grass (Bromus diandrus) was facilitated both by aboveground shrub biomass and by growing in soil taken from shrub patches. Experimental removal of shrubs negatively affected species most associated with shrubs and positively affected species most often found outside of shrubs. Counter to the stress-gradient hypothesis, the effects of shrubs on the physical environment and individual plant growth did not increase across the established stress gradient at this site. At the community level, shrub patches increased beta diversity, and contained greater rarified richness and exotic plant cover than shrub-free patches. Shrub effects on rarified richness increased with environmental stress, but effects on exotic cover and beta diversity did not. Our study provides evidence for the community-level effects of shrubs as ecosystem engineers in this system, but shows that these effects do not necessarily become stronger in more stressful environments.     

Native–Exotic Species Richness Relationships Across Spatial Scales in a Prairie Restoration Matrix

In highly invaded ecosystems, restoration of native plant communities is dependent upon reducing exotic species relative to native species. Even so, in monitoring, the native–exotic species richness ratio has been shown to be scale‐dependent. Measurement at small spatial scales (<1 m²) can reveal a negative native–exotic richness relationship, where niche occupation may prevent invasion. Conversely, at larger scales, a positive correlation may exist, where environmental heterogeneity and equally favorable conditions may drive native–exotic relationships. Here, we compare slopes of native–exotic relationships across spatial scales in a prairie undergoing active restoration. The observed native–exotic richness ratios varied considerably over scales ranging from 1 to 1,000 m², emphasizing the importance of choosing a measurement scale that is most pertinent to the treatment and ecological mechanism used to evaluate restoration success. Our native–exotic richness slopes were positive over all scales, but lower than would be expected in a random community assembly, suggesting the influence of niche‐based competition. Correspondingly, our native–exotic cover slope was more negative than a null model; however, areas of frequent fire treatments showed a significant deviation from null only for richness, indicating that burning may enhance native–exotic competitive dynamics for number of species but not cover. The negative native–exotic cover relationships appear to be driven in this system mainly by exotic graminoids, across burn treatments and native functional groups, supporting the concept that frequent burning can alter the dominant competitive mechanism from coverage of these exotic grasses to an improved environment for germination and dispersal of more native species.     

Nitrogen limitation, 15N tracer retention, and growth response in intact and Bromus tectorum-invaded Artemisia tridentata ssp. wyomingensis communities

Annual grass invasion into shrub-dominated ecosystems is associated with changes in nutrient cycling that may alter nitrogen (N) limitation and retention. Carbon (C) applications that reduce plant-available N have been suggested to give native perennial vegetation a competitive advantage over exotic annual grasses, but plant community and N retention responses to C addition remain poorly understood in these ecosystems. The main objectives of this study were to (1) evaluate the degree of N limitation of plant biomass in intact versus B. tectorum-invaded sagebrush communities, (2) determine if plant N limitation patterns are reflected in the strength of tracer ¹⁵N retention over two growing seasons, and (3) assess if the strength of plant N limitation predicts the efficacy of carbon additions intended to reduce soil N availability and plant growth. Labile C additions reduced biomass of exotic annual species; however, growth of native A. tridentata shrubs also declined. Exotic annual and native perennial plant communities had divergent responses to added N, with B. tectorum displaying greater ability to use added N to rapidly increase aboveground biomass, and native perennials increasing their tissue N concentration but showing little growth response. Few differences in N pools between the annual and native communities were detected. In contrast to expectations, however, more ¹⁵N was retained over two growing seasons in the invaded annual grass than in the native shrub community. Our data suggest that N cycling in converted exotic annual grasslands of the northern Intermountain West, USA, may retain N more strongly than previously thought.     

Patterns of woody plant invasion in an Argentinean coastal grassland

Coastal dune grasslands are fragile ecosystems that have historically been subjected to various types of uses and human activities. In Buenos Aires Province (Argentina), these areas are frequently afforested for urban and touristic development. The introduction and subsequent spread of exotic tree species is one of the main threats to conservation of natural grasslands as invasive trees strongly transform their structure and composition. The aim of this study was to identify patterns of woody plant invasion comparing plant communities and environmental variables between invaded and non-invaded areas surrounding the coastal village of Mar Azul, Argentina. Coastal grasslands in this area are being invaded by Populus alba (white poplar) and Acacia longifolia (coast wattle). The height of the saplings and the richness of the accompanying vegetation were evaluated in relation to the distance from the edge of the mature tree patches. Also, the cover, richness and diversity of all species in the invaded and non-invaded areas were measured, as well as soil pH, temperature and particle size. Negative correlations were found between the height of the saplings and distance to mature tree patches in all areas. The richness of the accompanying vegetation was negatively and positively correlated with the distance from the poplar and acacia area, respectively. The most abundant native species was Cortaderia selloana. Less cover, richness and diversity of native plant species and greater soil particle size were found in invaded areas, where the proportion of bare soil was higher. Also, a higher proportion of leaf litter in the invaded areas was registered. The results emphasize the invasive capacity of P. alba and A. longifolia advancing on the native communities and reducing their richness. Knowledge of the impact of invasive woody plants in coastal grasslands is important to design active management strategies for conservation purposes.     

Effects of exotic annual grass litter and local environmental gradients on annual plant community structure

Exotic annual grasses have been introduced into many semi-arid ecosystems worldwide, often to the detriment of native plant communities. The accumulation of litter from these grasses (i.e. residual dry biomass) has been demonstrated to negatively impact native plant communities and promote positive feedbacks to exotic grass persistence. More targeted experiments are needed, however, to determine the relative impact of exotic grass litter on plant community structure across local environmental gradients. We experimentally added exotic grass litter to annual forb-dominated open woodland communities positioned along natural canopy cover gradients in southwest Western Australia. These communities are an important component of this region’s plant biodiversity hotspot and are documented to be under threat from exotic annual grasses. After a one-year treatment period, we measured the effects of exotic grass litter, soil properties, and canopy cover on native and exotic species richness and abundance, as well as common species’ biomass and abundances. Plant community structure was more strongly influenced by soil properties and canopy cover than by grass litter. Total plant abundances per plot, however, were significantly lower in litter addition plots than control plots, a trend driven by native species. Exotic grass litter was also associated with lower abundances of one very common native species: Waitzia acuminata. Our results suggest that exotic grass litter limits the establishment of some native species in this system. Over multiple years, these subtle impacts may contribute substantially to the successful advancement of exotic species into this system, particularly in certain microenvironments.     

Estuarine and scalar patterns of invasion in the soft-bottom benthic communities of the San Francisco Estuary

The spatial patterns of nonindigenous species in seven subtidal soft-bottom communities in the San Francisco Estuary were quantified. Sixty nonindigenous species were found out of the 533 taxa enumerated (11%). Patterns of invasion across the communities were evaluated using a suite of invasion metrics based on the abundance or species richness of nonindigenous species. Patterns of invasion along the estuarine gradient varied with the invasion metric used, and the ecological interpretation of the metrics is discussed. Overall, the estuarine transition community located in the estuarine turbidity maximum zone (mean 5 practical salinity unit (psu)), main estuarine community (mean 16 psu), and marine muddy community (mean 28 psu) were more invaded than two fresh-brackish communities (mean < 1 psu) and a marine sandy community (mean 27 psu). Nonindigenous species were numerically dominant over much of the Estuary, making up more than 90% of the individuals in two communities. The percentage of the total species composed of nonindigenous species increased at smaller spatial scales: 11% at the estuary (gamma) scale, 21% at the community (alpha) scale, and 42% at the grab (point) scale. Wider spatial distributions of nonindigenous species and a relatively greater percentage of rare native species may have resulted in this pattern. Because of this scale dependency, comparisons among sites need to be made at the same spatial scale. Native species were positively correlated with nonindigenous species in several of the communities, presumably due to similar responses to small-scale differences in habitat quality. The rate of invasion into the soft-bottom communities of the San Francisco Estuary appears to have increased over the last one to two decades and many of the new introductions have become numerically dominant.