Exotic Plant Species Richness Research Articles

Assessing exotic plant species invasions and associated soil characteristics: A case study in eastern Rocky Mountain National Park, Colorado, USA, using the pixel nested plot design

Rocky Mountain National Park (RMNP), Colorado, USA, contains a diversity of plant species. However, many exotic plant species have become established, potentially impacting the structure and function of native plant communities. Our goal was to quantify patterns of exotic plant species in relation to native plant species, soil characteristics, and other abiotic factors that may indicate or predict their establishment and success. Our research approach for field data collection was based on a field plot design called the pixel nested plot. The pixel nested plot provides a link to multi-phase and multi-scale spatial modeling-mapping techniques that can be used to estimate total species richness and patterns of plant diversity at finer landscape scales. Within the eastern region of RMNP, in an area of approximately 35,000 ha, we established a total of 60 pixel nested plots in 9 vegetation types. We used canonical correspondence analysis (CCA) and multiple linear regressions to quantify relationships between soil characteristics and native and exotic plant species richness and cover. We also used linear correlation, spatial autocorrelation and cross correlation statistics to test for the spatial patterns of variables of interest. CCA showed that exotic species were significantly ( P < 0.05) associated with photosynthetically active radiation ( r = 0.55), soil nitrogen ( r = 0.58) and bare ground ( r = −0.66). Pearson's correlation statistic showed significant linear relationships between exotic species, organic carbon, soil nitrogen, and bare ground. While spatial autocorrelations indicated that our 60 pixel nested plots were spatially independent, the cross correlation statistics indicated that exotic plant species were spatially associated with bare ground, in general, exotic plant species were most abundant in areas of high native species richness. This indicates that resource managers should focus on the protection of relatively rare native rich sites with little canopy cover, and fertile soils. Using the pixel nested plot approach for data collection can facilitate the ecological monitoring of these vulnerable areas at the landscape scale in a time- and cost-effective manner.

Successful biological control of mist flower (Ageratina riparia) in New Zealand: Agent establishment, impact and benefits to the native flora

The white smut fungus (Entyloma ageratinae) and the gall Xy (Procecidochares alani) were released in New Zealand in 1998 and 2001 respectively to suppress mist Xower (Ageratina riparia). The fungus established and spread rapidly, crossing 80 km of sea to Great Barrier Island within 2 years. The mean number of P. alani galls increased exponentially to 1.96/stem at release sites, but dispersal was slow. The impact of the biocontrol agents was monitored once annually from 1998/99 to 2003/04, at up to 51 sites in the North Island. The mean percentage of live leaves infected with fungus rapidly reached nearly 60%. Maximum plant height declined signiWcantly. In heavy infestations, mean percentage cover of mist Xower declined from 81 to 1.5%. Galls were only recorded towards the end of the impact study, and at low mean numbers. As mist Xower declined, the species richness and mean percentage cover of native plants increased. In contrast, the species richness and mean percentage cover of exotic plants (excluding mist Xower) did not change signiWcantly. Many plant species colonizing the plots were important native mid- or late-successional shrubs or trees. With few exceptions, the exotic plant species common in the plots were not weeds that appeared to threaten native forest habitats. There was only a weak “replacement weed eVect” from the potentially serious invader African club moss (Selaginella kraussiana). These data, together with reports of reduced threats to rare endemic plants from mist Xower, suggest this rapid, well-monitored weed biocontrol program was very successful.

Opposing Effects of Native and Exotic Herbivores on Plant Invasions

Exotic species are widely assumed to thrive because they lack natural enemies in their new ranges. However, a meta-analysis of 63 manipulative field studies including more than 100 exotic plant species revealed that native herbivores suppressed exotic plants, whereas exotic herbivores facilitated both the abundance and species richness of exotic plants. Both outcomes suggest that plants are especially susceptible to novel, generalist herbivores that they have not been selected to resist. Thus, native herbivores provide biotic resistance to plant invasions, but the widespread replacement of native with exotic herbivores eliminates this ecosystem service, facilitates plant invasions, and triggers an invasional "meltdown."

Soil phosphorus and tree cover modify the effects of livestock grazing on plant species richness in Australian grassy woodland

The effects of grazing on the richness of understorey plant communities are predicted to vary along gradients of resources and tree cover. In temperate Australia livestock management has involved phosphorus addition and tree removal but little research has examined how the effects of grazing on plant species richness may vary with these management regimes. Patterns of understorey plant species richness were examined in 519, 0.09 ha quadrats in grazed pastures and remnant grassy forests and woodlands in southern Australia. Sheep grazing was the primary land use and sites varied widely in grazing frequency and density, tree cover and phosphorus fertiliser history. Using an information theoretic approach the available data provides strong evidence that the effect of grazing on total species richness varies according to available phosphorus and tree cover. Intermittent grazing and no grazing were associated with high total and native plant richness, but only at low phosphorus concentrations. Phosphorus was strongly negatively correlated with richness, particularly at low grazing frequency. Total species richness was positively correlated with tree cover except under frequent grazing at high stocking rates, suggesting that heavy grazing eliminates spatial and temporal heterogeneity imposed by trees. Native plant species richness was negatively correlated with a history of cultivation, positively correlated with tree cover and varied according to landscape position and geological substrate. Frequent high density grazing, particularly when associated with clearing, cultivation and fertiliser addition, was associated with the persistence of very few native plant species. In contrast, the richness of exotic plant species was relatively invariant and performance of the best model was low. While several studies have highlighted the importance of the grazed and cleared matrix for the conservation of native plant species, this benefit may be limited in landscapes where intensive grazing management systems dominate. Strong evidence for interactions between grazing, phosphorus and tree cover suggest that failure to consider other land use practices associated with grazing management systems could lead to erroneous conclusions regarding vegetation responses to livestock grazing.

Patterns of exotic and native plant species richness and cover along a semi-arid Iberian river and across its floodplain

Patterns of native and exotic species richness and cover were examined together with the composition of species assemblages along a typical Mediterranean river and across its floodplain. Major goals were to characterize species distribution regarding diverse environmental and habitat features, and to explore floodplain landforms relations using plant assemblages and invasion responses as biological indicators of aquatic–terrestrial connectivity. A floristic survey was undertaken during May and early June 1998 in six sites of a southern Iberian basin. Vegetation data were gathered from three transects in three landforms: inundated and frequently flooded banks (FF), infrequently flooded valley floors (IF), and rarely flooded higher floodplains (RF). We used 2×2 m plots to record each species present and visual estimates of percentage cover of each species. Cover of native and exotic species assemblages were also estimated for each plots, for the transects, and for the landforms. The total number of vascular plant species found was 287 of which only 26 were exotic. Despite the high environmental and geomorphological variability, the hierarchical classification showed that species composition varied across the floodplain, rather than along the river. The IF and RF landforms were the most species-rich, both for native and exotic species group, whereas the cover trends of exotic and native plant species were opposed, with exotics being consistently more abundant in FF, and native cover increasing towards RF landforms. We found positive correlations between native and exotic species richness on the floodplain scale, and on a smaller scale in FF and IF landforms, thus supporting the hypothesis that in aquatic and riparian ecosystems of semi-arid landscapes, species-rich communities are preferentially colonized by exotic species. However, this pattern was not consistently coupled with a high cover of exotic species, and it was not sustained for the dry areas of the RF landforms.

Human impacts, energy availability and invasion across Southern Ocean Islands

ABSTRACTAim Ongoing biological invasions will enhance the impacts of humans on biodiversity. Nonetheless, the effects of exotic species on diversity are idiosyncratic. Increases in diversity might be a consequence of similar responses by species to available energy, or because of positive relationships between human density, energy and propagule pressure. Here we use data from the Southern Ocean island plants and insects to investigate these issues.Location The Southern Ocean Islands ranging from Tristan da Cunha to Heard Island and South Georgia.Methods Generalized linear models are used to explore the relationships between indigenous and exotic species richness for plants and insects on two different islands. Similar models are used to examine interactions between indigenous and exotic species richness, energy availability and propagule pressure at the regional scale.Results Positive relationships were found between indigenous and exotic species richness at local scales, although for plants, the relationship was partially triangular. Across the Southern Ocean Islands, there was strong positive covariation between indigenous and exotic plant species richness and insect species richness, even taking spatial autocorrelation into account. Both exotic and indigenous plant and insect species richness covaried with energy availability, as did human visitor frequency. When two islands with almost identical numbers of human visits were contrasted, it was clear that energy availability, or perhaps differences in climate‐matching, were responsible for differences in the extent of invasion.Conclusion In plants and insects, there are positive relationships between indigenous and exotic diversity at local and regional scales across the Southern Ocean islands. These relationships are apparently a consequence of similar responses by both groups and by human occupants to available energy. When visitor frequency is held constant, energy availability is the major correlate of exotic species richness, though the exact mechanistic cause of this relationship requires clarification.

Contrasting impacts of a native and an invasive exotic shrub on flood‐plain succession

Abstract. Question: How do Coriaria arborea, an N‐fixing native shrub, and Buddleja davidii, a non‐N‐fixing exotic shrub, affect N:P stoichiometry in plants and soils during early stages of primary succession on a flood‐plain?Location: Kowhai River Valley, northeast South Island, New Zealand.Methods: We measured soil and foliar nutrient concentrations, light levels, plant community composition and the above‐ground biomass of Coriaria and Buddleja in four successional stages: open, young, vigorous and mature.Results: Coriaria occurred at low density but dominated above‐ground biomass by the vigorous stage. Buddleja occurred at 5.3 ± 1.0 stems/m2 in the young stage and reached a maximum biomass of 520–535 g.m‐2 during the young and vigorous stages. Mineral soil N increased with above‐ground Coriaria biomass (r2= 0.45), but did not vary with Buddleja biomass. In contrast, soil P increased with Buddleja biomass (r2= 0.35), but not with Coriaria biomass. In early successional stages, 70–80% of the species present were exotic, but this declined to about 15% by the mature stage. Exotic plant species richness declined with increasing Coriaria biomass, but no other measures of diversity varied with either Coriaria or Buddleja biomass.Conclusion: These results demonstrate that Buddleja dominates early succession and accumulates P whereas Coriaria dominates later succession and accumulates N. A key ecosystem effect of the invasive exotic Buddleja is alteration of soil N:P stoichiometry.

Contrasting impacts of a native and an invasive exotic shrub on flood-plain succession

Question: How do Coriaria arborea, an N-fixing native shrub, and Buddleja davidii, a non-N-fixing exotic shrub, affect N:P stoichiometry in plants and soils during early stages of primary succession on a flood-plain? Location: Kowhai River Valley, northeast South Island, New Zealand. Methods: We measured soil and foliar nutrient concentrations, light levels, plant community composition and the aboveground biomass of Coriaria and Buddleja in four successional stages: open, young, vigorous and mature. Results: Coriaria occurred at low density but dominated above-ground biomass by the vigorous stage. Buddleja occurred at 5.3 ± 1.0 stems/m 2 in the young stage and reached a maximum biomass of 520-535 g.m –2 during the young and vigorous stages. Mineral soil N increased with above-ground Coriaria biomass (r 2 = 0.45), but did not vary with Buddleja biomass. In contrast, soil P increased with Buddleja biomass (r 2 = 0.35), but not with Coriaria biomass. In early successional stages, 70-80% of the species present were exotic, but this declined to about 15% by the mature stage. Exotic plant species richness declined with increasing Coriaria biomass, but no other measures of diversity varied with either Coriaria or Buddleja biomass. Conclusion: These results demonstrate that Buddleja dominates early succession and accumulates P whereas Coriaria dominates later succession and accumulates N. A key ecosystem effect of the invasive exotic Buddleja is alteration of soil N:P stoichiometry.

VARIABLE EFFECTS OF FERAL PIG DISTURBANCES ON NATIVE AND EXOTIC PLANTS IN A CALIFORNIA GRASSLAND

Biological invasions are a global phenomenon that can accelerate disturbance regimes and facilitate colonization by other nonnative species. In a coastal grassland in northern California, we conducted a four-year exclosure experiment to assess the effects of soil disturbances by feral pigs (Sus scrofa) on plant community composition and soil nitrogen availability. Our results indicate that pig disturbances had substantial effects on the community, although many responses varied with plant functional group, geographic origin (native vs. exotic), and grassland type. (''Short patches'' were dominated by annual grasses and forbs, whereas ''tall patches'' were dominated by perennial bunchgrasses.) Soil disturbances by pigs increased the richness of exotic plant species by 29% and native taxa by 24%. Although native perennial grasses were unaffected, disturbances reduced the bio- mass of exotic perennial grasses by 52% in tall patches and had no effect in short patches. Pig disturbances led to a 69% decrease in biomass of exotic annual grasses in tall patches but caused a 62% increase in short patches. Native, nongrass monocots exhibited the opposite biomass pattern as those seen for exotic annual grasses, with disturbance causing an 80% increase in tall patches and a 56% decrease in short patches. Native forbs were unaffected by disturbance, whereas the biomass of exotic forbs increased by 79% with disturbance in tall patches and showed no response in short patches. In contrast to these vegetation results, we found no evidence that pig disturbances affected nitrogen mineral- ization rates or soil moisture availability. Thus, we hypothesize that the observed vegetation changes were due to space clearing by pigs that provided greater opportunities for colo- nization and reduced intensity of competition, rather than changes in soil characteristics. In summary, although responses were variable, disturbances by feral pigs generally pro- moted the continued invasion of this coastal grassland by exotic plant taxa.

Citizens as Propagules for Exotic Plants: Measurement and Management Implications1

Abstract There is a significant positive association between exotic plant species richness of an area and the human population size of that area. This association is notable not only for its high correlation (r2 = 0.69) but also because it spans five orders of magnitude for human population size. The correlation is apparently caused by the introduction of increasing numbers of exotic species as human population size grows. As population grows, so does exotic propagule importation and habitat disturbance. The regression is clearly asymptotic, indicating that per capita rates of exotic species introductions decline as human population continues to increase. A main implication for exotic species control is that efforts to limit exotic species will be most effective and least costly in areas (at many scales) where human population size is still relatively small. Another implication is that because human population size will continue to increase in many areas, educating the general public about harmful exotic ...

Applicability of landscape and island biogeography theory to restoration of riparian understorey plants

Summary Ecosystem recovery is influenced by processes at different spatial scales, yet land managers lack specific predictions on the relative importance of such processes that might guide management decisions. We tested whether ideas from landscape ecology (local vs. landscape scales) and island biogeography theory (patch size and isolation) predict restoration success for understorey plant communities in a highly fragmented riparian landscape, in an effort to provide guidance on how to allocate scarce restoration resources. We sampled naturally colonizing riparian forest understorey plant communities in 15 riparian forests restored by planting native woody species along a 150‐km stretch of the Sacramento River in central California. We analysed native and exotic understorey species richness and cover as a function of biotic and abiotic local and landscape variables. Cover and species richness of exotic understorey plants decreased strongly with increasing overstorey cover, and were lower in quadrats closer to river base flow. Native understorey species richness and cover were negatively related to exotic cover and positively related to connectivity with remnant forest. Cover of native wind‐dispersed species was best explained by higher percentage forest cover surrounding a site within a 1000‐m buffer, whereas cover of native water‐dispersed species was higher closer to the river. Neither patch size nor time since restoration explained a significant amount of native or exotic species richness or cover. Synthesis and applications. Local factors explained more of the variance in understorey plant communities, but much of the variance remained unexplained. Our results provide weak support for the predictions of island biogeography theory and the importance of landscape‐scale variables. These theories did not have strong predictive power in this applied restoration context at this temporal scale. Given limited resources, efforts to restore understorey plant communities in this highly fragmented system should focus on local‐scale restoration methodologies, such as increasing cover of native overstorey species and reducing cover of exotic plants.

What determines disturbance‐productivity‐diversity relationships? The effect of scale, species and environment on richness patterns in an Australian woodland

Much of the observed variation in relationships between diversity and disturbance or productivity may be attributed to scale, species characteristics, or environment. We used exclusion fences to create gradients of grazing (by native and introduced herbivores), cover, and standing crop in temperate Eucalypt woodlands. We investigated patterns of native, exotic, and total plant species richness at two scales (1 m2 and 625 m2). Richness patterns were similar at both scales, though species richness at 1m2 was more strongly affected by our grazing treatments. Season and rainfall explained more variation in richness than did surrogate measures of productivity or disturbance by herbivores. The richness‐herbivory relationship depended strongly on rainfall, season, and species origin, and altering these factors produced the entire range of observed diversity‐disturbance relationships. Richness‐biomass and richness‐cover relationships were consistently hump‐shaped, and related to species origin with native richness negatively related and exotic richness positively related. The ability of weedy annuals to pre‐empt space after death may have contributed to the observed unimodal responses.

Soil characteristics and plant exotic species invasions in the Grand Staircase—Escalante National Monument, Utah, USA

The Grand Staircase—Escalante National Monument (GSENM) contains a rich diversity of native plant communities. However, many exotic plant species have become established, potentially threatening native plant diversity. We sought to quantify patterns of native and exotic plant species and cryptobiotic crusts (mats of lichens, algae, and mosses on the soil surface), and to examine soil characteristics that may indicate or predict exotic species establishment and success. We established 97 modified-Whittaker vegetation plots in 11 vegetation types over a 29,000 ha area in the Monument. Canonical correspondence analysis (CCA) and multiple linear regressions were used to quantify relationships between soil characteristics and associated native and exotic plant species richness and cover. CCA showed that exotic species richness was significantly ( P<0.05) associated with soil P ( r=0.84), percentage bare ground ( r=0.71), and elevation ( r=0.67). Soil characteristics alone were able to predict 41 and 46% of the variation in exotic species richness and cover, respectively. In general, exotic species invasions tend to occur in fertile soils relatively high in C, N and P. These areas are represented by rare mesic high-elevation habitats that are rich in native plant diversity. This suggests that management should focus on the protection of the rare but important vegetation types with fertile soils.

Changes to plant species richness in forest fragments: fragment age, disturbance and fire history may be as important as area

AimThe impact of fragmentation on a eucalypt forest was investigated by examining the effects of fragment size, time since fragmentation, degree of anthropogenic disturbance to fragment interiors, and time since fire, on native and exotic plant species richness per unit area.LocationTwo areas of dry open‐forest were studied on the central coast of New South Wales in south‐eastern Australia. Fifty forest fragments were located at Tomago, an area progressively fragmented over the last 60 years, most recently by clearing for sand‐mining. Also at Tomago were six very large blocks of forest that were used as reference sites. The second area at Myall Lakes National Park (50 km north of Tomago) had four very large areas of intact forest that were also used as reference sites.MethodsFragments were allocated into (1) three size classes: small (&lt;1 ha), medium (1 to &lt;10 ha) and large (10 to &lt;100 ha); (2) two classes of age since fragmentation: young (up to 10 years) and old (10 years or more); (3) two classes of disturbance (minor, major) based on the degree and extent of human‐induced disturbance to the fragment interior; and (4) two classes of time since fire (between 5 and 6 years; 10 years or more). Reference sites belonged to a very large size class (&gt;100 ha). Mean plant species richness per 25 m2 area was determined for each fragment and analysed separately for native and exotic species.ResultsThe most significant effects observed resulted from anthropogenic disturbance. In fragments with major disturbance, native species richness per unit area was significantly reduced in small‐sized young and medium‐sized old fragments. A significant increase was also observed for exotic species richness in fragments with major disturbance. With minor disturbance, native species richness in small fragments declined significantly with time since fragmentation, in contrast to medium and large fragments. Among recently created fragments, those recently burned had significantly more native species per unit area than those burned 10 or more years ago.Main conclusionsAnthropogenic disturbance coupled with fragmentation had a stronger and more immediate effect in reducing native species richness and increasing exotic species richness than did fragmentation alone. In the absence of major disturbance, small fragments had fewer native species than larger size classes, but only after 10 or more years since fragmentation, confirming the importance of controlling for age of fragments when examining species–area relationships. This study has not tested whether differences in area were the direct cause of this loss of species over time; other factors that are correlated with area (such as edge effects) may also be involved. The increase in native species richness following fire was consistent with other studies of fire in unfragmented eucalypt forest. This study thus shows that in addition to the factors emphasized in classical island biogeography models, fragment age, disturbance and fire history are important in explaining species richness in fragmented eucalypt forests.

Predicting plant species diversity in response to disturbance magnitude in grassland remnants of central Alberta

The Intermediate Disturbance Hypothesis states that the greatest species diversity occurs at intermediate levels of disturbance because species coexistence is maintained at a nonequilibrium state and no strong competitor can dominate completely. On the other hand, diversity of exotic species is expected to increase with the disturbance magnitude. These patterns were tested for in this study. The cover of all vascular plants, mosses, and lichens in 1 × 1 m plots across a range of disturbance levels was sampled in 11 remnant grasslands within the Aspen Parkland Ecoregion of central Alberta, western Canada. The Intermediate Disturbance Hypothesis was supported for small-scale within-patch diversity for total species richness and Simpson's diversity index. Lower species diversity was found in undisturbed and lightly grazed as well as in highly disturbed plots. Intermediate levels of disturbance had reduced dominance of Festuca hallii (Vasey) Piper and increased abundance of most other species; this gave the highest species diversity. The species richness and diversity of exotic plant species showed a significant positive relationship with the magnitude of the disturbance. Understanding relationships between disturbance and plant species diversity, especially exotic plants, can influence management decisions on what disturbance regime is conducive to maintaining natural plant communities.Key words: Intermediate Disturbance Hypothesis, diversity, exotic species, grassland.

Size of the local species pool determines invasibility of a C4‐dominated grassland

The size of the local species pool (i.e., species surrounding a community capable of dispersal into that community) and other dispersal limitations strongly influence native plant community composition. However, the role that the local species pool plays in determining the invasibility of communities by exotic plants remains to be evaluated. We hypothesized that the richness and abundance of exotic species would be greater in C4‐dominated grassland communities if the local species pool included a larger proportion of exotic species. We also predicted that an increase in the exotic species pool would increase the invasibility of sites thought to be resistant to invasion (annually burned grassland). To test these hypotheses, study plots were established within two long‐term (&gt;20 yr) fire experiments at a tallgrass prairie preserve in NE Kansas (USA). Study plots were surrounded by either a small pool of exotic species (small species pool (SSP) plots; six species) or a larger exotic species pool (large species pool (LSP) plots; 18 species). We found that richness and absolute cover of exotic species was significantly (P&lt;0.001) lower (∼70 and 90%, respectively) in annually burned compared to unburned plots, regardless of the size of the exotic species pool. As predicted, exotic species richness was higher (P&lt;0.001) for LSP plots (3.9 per 250 m2) than for SSP plots (0.7 per 250 m2); however, absolute cover was unaffected by the size of the exotic species pool. In the absence of fire, plots with a LSP had four times as many exotic species than SSP plots. An increase in the local exotic species pool also increased the invasibility of annually burned grassland. Indeed, richness of exotic plant species in annually burned LSP plots did not differ from unburned plots with a SSP, indicating that a larger pool of exotic species countered the negative effects of fire. These findings have important implications for predicting how the invasion of plant communities may respond to human‐induced global changes, such as habitat fragmentation. Community characteristics or factors such as frequent fires in grasslands may impart resistance to invasions by exotic species in large, intact ecosystems. However, when a large pool of exotic species is present, frequent fire may not be sufficient to limit the invasions of exotic plants in fragmented landscapes.

Rapid assessment of butterfly diversity in a montane landscape

We present the results of a rapid assessment of butterfly diversity in the 754 ha Beaver Meadows study area in Rocky Mountain National Park, Larimer County, Colorado. We measured butterfly species richness and relative abundance as part of a landscape-scale investigation of diversity patterns involving several groups of organisms. A stratified random sampling design was used to include replication in both rare and common vegetation types. We recorded 49 butterfly species from the twenty-four 0.1 ha plots that were sampled four times during June, July, and August 1996. Butterfly species richness, diversity, and uniqueness were highest in quaking aspen (Populus tremuloides Michaux) groves and wet meadows, which occupy only a small proportion of the studied landscape. This result supports the suggestion that aspen areas represent ‘hotspots’ of biological diversity in this montane landscape. Patterns of butterfly species richness were positively correlated with total vascular plant species richness (r = 0.69; P < 0.001), and native plant species richness (r = 0.64; P < 0.001). However, exotic plant species richness (r = 0.70; P < 0.001) and the cover of exotic plant species (r = 0.70; P < 0.001) were the best predictors of butterfly species richness.

The impact of suburbanization on remnant coastal vegetation in Hobart, Tasmania

Abstract. Changes in vascular species composition and abundance were examined in coastal vegetation at Hobart, Tasmania, Australia over a period of 13 yr, during which suburbanization extended to cover a large proportion of its hinterland. There were significant increases in the richness and cover of exotic vascular plant species, most of which derived from the Mediterranean basin or southern Africa. While most of the introduced species were confined to vegetation with a high exotic component, and close to domestic gardens, some, including the South African coastal shrub, Chrysanthemoides monilifera, expanded in otherwise native vegetation relatively remote from gardens. The impact of proximity to gardens on the relative abundance of exotics and natives in coastal vegetation in 1997 best correlated with the pattern of distance from gardens in 1966, suggesting that there is, at least, a 30‐yr lag period in transformation from largely native to substantially exotic vegetation. This lag period provides an opportunity to reduce exotic invasion problems before they become severe.

Exotic plant invasion and understorey species richness: a comparison of two types of eucalypt woodland in agricultural Western Australia

Exotic plants are a major threat to native plant diversity in Australia yet a generic model of the invasion of Australian ecosystems by exotic species is lacking because invasion levels differ with vegetation/soil type and environmental conditions. This study compared relative differences in exotic species invasion (percent cover, spp. richness) and the species richness of herbaceous native plants in two structurally very similar vegetation types, Gimlet Eucalyptus salubris and Wandoo E. capillosa woodlands in the Western Australian wheatbelt. For each woodland type, plant variables were measured for relatively undisturbed woodlands, woodlands with &gt;30 years of livestock grazing history, and woodlands in road-verges. Grazed and road-verge Gimlet and Wandoo woodlands had significantly higher cover of exotic species, and lower species richness of native plants, compared with undisturbed Gimlet and Wandoo. Exotic plant invasion was significantly greater in Gimlet woodlands for both grazed (mean 78% cover) and road-verge sites (mean 42% cover) than in comparable sites in Wandoo woodlands (grazed sites 25% cover, road-verge sites 19% cover). There was no significant difference in the species richness of exotic plants between Wandoo and Gimlet sites for any of the three situations. Mean site richness of native plants was not significantly different between undisturbed Wandoo and undisturbed Gimlet woodlands. Undisturbed woodlands were significantly richer in plant species than grazed and road-verge woodlands for both woodland types. Grazed and road-verge Wandoo sites were significantly richer in plant species than communities in grazed and road-verge Gimlet. The percent cover of exotics was negatively correlated with total (native) plant species richness for both woodland types (Wandoo r = ?0.70, Gimlet r = ?0.87). Of the total native species recorded in undisturbed Gimlet, 83% and 61% were not recorded in grazed and road-verge Gimlet, respectively. This compared with 40% and 33% for grazed and road-verge Wandoo, respectively. Grazed Wandoo and grazed Gimlet sites had significantly fewer native plant species than did road-verge Wandoo and road-verge Gimlet sites. Ecosystem implications of differential invasions by exotic species, and the effects of grazing (disturbance) and other factors influencing susceptibility to exotic plant invasion (landscape, competition and allelopathy) on native species decline are discussed. Exclusion of livestock and adequate methods of control and prevention of further invasions by exotic plants are essential requirements for the conservation of these woodland systems.

Promotion of exotic weed establishment by endangered giant kangaroo rats (Dipodomys ingens) in a California grassland

Giant kangaroo rats (Dipodomys ingens) continually modify their burrow precincts by digging tunnels, clipping plants, and other activities. In the valley grasslands of the Carrizo Plain Natural Area (San Luis Obispo County, California), this chronic disturbance to soil and vegetation promoted the establishment of exotic ruderal and early successional plant species. Erodium cicutarium, Bromus madritensis ssp. rubens, and other Mediterranean annuals were found to constitute a very large proportion of the vegetation on giant kangaroo rat precincts. When vegetation on precincts was compared with the vegetation in less disturbed intermediate areas located between precincts, species richness, cover and frequency of exotic plants were significantly greater on precincts. The reverse was found for native species. In addition, exotic species encountered in this study had significantly larger seeds than did native species, suggesting that these granivorous kangaroo rats preferentially cache large weed seeds on their precincts. Since the kangaroo rats depend on exotic plants for food and the exotic plants depend upon the kangaroo rats to disturb their habitat continually, the weed-kangaroo rat relationship is mutualistic. This strong relationship may also inhibit population growth of native grassland plants which occupy disturbed habitats but have difficulty competing with exotic weeds for resources. From a conservation perspective, this mutualism presents an intractable management dilemma. Restoration of valley grasslands where endangered giant kangaroo rats occur, to conditions where native species dominate, may be impossible.