Phylogenetic relatedness and plant invader success across two spatial scales

Aim  Successful invaders often possess similar ecological traits that contribute to success in new regions, and thus under niche conservatism, invader success should be phylogenetically clustered. We asked if the degree to which non-native plant species are phylogenetically related is a predictor of invasion success at two spatial scales.
Location  Australia – the whole continent and Royal National Park (south-eastern Australia).
Methods  We used non-native plant species occupancy in Royal National Park, as well as estimated continental occupancy of these species from herbarium records. We then estimated phylogenetic relationships using molecular data from three gene sequences available on GenBank ( matK , rbcL and ITS1 ). We tested for phylogenetic signals in occupancy using Blomberg's K .
Results  Whereas most non-native plants were relatively scarce, there was a strong phylogenetic signal for continental occupancy, driven by the clustering of successful species in Asteraceae, Caryophyllaceae, Poaceae and Solanaceae. However, we failed to detect a phylogenetic signal at the park scale.
Main Conclusions  Our results reveal that at a large spatial scale, invader success is phylogenetically clustered where ecological traits promoting success appear to be shared among close relatives, indicating that phylogenetic relationships can be useful predictors of invasion success at large spatial scales. At a smaller, landscape scale, there was no evidence of phylogenetic clustering of invasion success, and thus, relatedness plays a much reduced role in determining the relative success of invaders.     

A latitudinal gradient of beta diversity for exotic vascular plant species in North America

Determining relationships between the ranges of introduced species and geographical and environmental factors is an important step in understanding the mechanisms and processes of the spread of introduced species. In this study, I examined the beta diversity and latitude relationship for all naturalized exotic species of vascular plants in North America at a continental scale. Beta diversity was calculated as the absolute value of the slope of the relationship between the natural logarithm of the Simpson index of similarity (lnS) and spatial distance between pairs of state‐level exotic floras within four latitudinal zones examined. Relative contributions of spatial distance and environmental difference to species turnover between exotic floras were examined. I found that beta diversity decreased monotonically from low to high latitudes: beta diversity for the southernmost zone was shallower than that for the northernmost zone by a factor of 2.6. Regression models of lnS in relation to spatial distance and environmental (climatic and topographical) difference for each latitudinal zone demonstrated that the explanatory power of these variables diminishes monotonically with latitude: the explained variance in lnS is 70.4%, 62.1%, 53.9%, and 33.9%, respectively, for the four latitudinal zones from south to north. For the southernmost zone, 58.3% of the variance in lnS is explained by climate variables and topography, and spatial distance explains only 2.3% of the variance. In contrast, for the northernmost zone, more than half the amount (22.5%) of the explained variance in lnS is attributable to spatial distance, and the remaining (18.9%) of the explained variance is attributable to climate variables and topography.     

Stream communities across a rural–urban landscape gradient

Rapid urbanization throughout the world is expected to cause extensive loss of biodiversity in the upcoming decades. Disturbances associated with urbanization frequently operate over multiple spatial scales such that local species extirpations have been attributed both to localized habitat degradation and to regional changes in land use. Urbanization also may shape stream communities by restricting species dispersal within and among stream reaches. In this patch-dynamics view, anthropogenic disturbances and isolation jointly reduce stream biodiversity in urbanizing landscapes. We evaluated predictions of stream invertebrate community composition and abundance based on variation in environmental conditions at five distinct spatial scales: stream habitats, reaches, riparian corridors and watersheds and their spatial location within the larger three-river basin. Despite strong associations between biodiversity loss and human density in this study, local stream habitat and stream reach conditions were poor predictors of community patterns. Instead, local community diversity and abundance were more accurately predicted by riparian vegetation and watershed landscape structure. Spatial coordinates associated with instream distances provided better predictions of stream communities than any of the environmental data sets. Together, results suggest that urbanization in the study region was associated with reduced stream invertebrate diversity through the alteration of landscape vegetation structure and patch connectivity. These findings suggest that maintaining and restoring watershed vegetation corridors in urban landscapes will aid efforts to conserve freshwater biodiversity.     

Scale-dependent relationships between native richness, resource stability and exotic cover in dock fouling communities of Washington, USA

Aim In terrestrial plant communities, the relationship between native species diversity and exotic success is typically scale‐dependent. It is often proposed that within local neighbourhoods, high native diversity limits resources, thereby inhibiting exotic success. However, environmental variation that manifests over space or time can create positive correlations between native diversity and exotic success at larger scales. In marine habitats, there have been few multi‐scale surveys of this pattern, so it is unclear how diversity, resource limitation and the environment influence the success of exotic species in these systems. Location Washington, USA. Methods I analysed nested spatial and temporal surveys of fouling communities, which are assemblages of sessile marine invertebrates, to test whether the relationships between native richness, resource availability and exotic cover supported the diversity‐stability and diversity‐resistance theories, to test whether these relationships changed with spatio‐temporal scale, and to explore the temperature preferences of native and exotic fouling species. Results Survey data failed to support diversity‐stability theory: space availability actually increased with native richness at the local neighbourhood scale, and neither space availability nor variability decreased with native richness across larger spatio‐temporal scales. I did find support for diversity‐resistance theory, as richness negatively correlated with exotic cover in local neighbourhoods. Unexpectedly, this negative correlation disappeared at intermediate scales, but emerged again at the regional scale. This scale‐dependent pattern could be partially explained by contrasting water temperature preferences of native and exotic species. Main conclusions Within local neighbourhoods, native diversity may inhibit exotic abundance, but the mechanism is unlikely related to resource limitation. At the largest scale, correlations suggest that native richness is higher in cooler environments, whereas exotic richness is higher in warmer environments. This large‐scale pattern contrasts with the typical plant community pattern, and has important implications for coastal management in the face of global climate change.     

Phylogenetic diversity metrics for ecological communities: integrating species richness, abundance and evolutionary history

Phylogenetic information is increasingly being used to understand the assembly of biological communities and ecological processes. However, commonly used metrics of phylogenetic diversity (PD) do not incorporate information on the relative abundances of individuals within a community. In this study, we develop three indices of PD that explicitly consider species abundances. First, we present a metric of phylogenetic-abundance evenness that evaluates the relationship between the abundance and the distribution of terminal branch lengths. Second, we calculate an index of hierarchical imbalance of abundances at the clade level encapsulating the distribution of individuals across the nodes in the phylogeny. Third, we develop an index of abundance-weighted evolutionary distinctiveness and generate an entropic index of phylogenetic diversity that captures both information on evolutionary distances and phylogenetic tree topology, and also serves as a basis to evaluate species conservation value. These metrics offer measures of phylogenetic diversity incorporating different community attributes. We compare these new metrics to existing ones, and use them to explore diversity patterns in a typical California annual grassland plant community at the Jasper Ridge biological preserve.
Ecology Letters (2010) 13: 96–105     

BIODIVERSITY RESEARCH: Native‐exotic species richness relationships across spatial scales and biotic homogenization in wetland plant communities of Illinois,USA

Aim To examine native‐exotic species richness relationships across spatial scales and corresponding biotic homogenization in wetland plant communities. Location Illinois, USA. Methods We analysed the native‐exotic species richness relationship for vascular plants at three spatial scales (small, 0.25 m² of sample area; medium, 1 m² of sample area; large, 5 m² of sample area) in 103 wetlands across Illinois. At each scale, Spearman’s correlation coefficient between native and exotic richness was calculated. We also investigated the potential for biotic homogenization by comparing all species surveyed in a wetland community (from the large sample area) with the species composition in all other wetlands using paired comparisons of their Jaccard’s and Simpson’s similarity indices. Results At large and medium scales, native richness was positively correlated with exotic richness, with the strength of the correlation decreasing from the large to the medium scale; at the smallest scale, the native‐exotic richness correlation was negative. The average value for homogenization indices was 0.096 and 0.168, using Jaccard’s and Simpson’s indices, respectively, indicating that these wetland plant communities have been homogenized because of invasion by exotic species. Main Conclusions Our study demonstrated a clear shift from a positive to a negative native‐exotic species richness relationship from larger to smaller spatial scales. The negative native‐exotic richness relationship that we found is suggested to result from direct biotic interactions (competitive exclusion) between native and exotic species, whereas positive correlations likely reflect the more prominent influence of habitat heterogeneity on richness at larger scales. Our finding of homogenization at the community level extends conclusions from previous studies having found this pattern at much larger spatial scales. Furthermore, these results suggest that even while exhibiting a positive native‐exotic richness relationship, community level biotas can/are still being homogenized because of exotic species invasion.     

Pollution reduces native diversity and increases invader dominance in marine hard-substrate communities

Anthropogenic disturbance is considered a risk factor in the establishment of non‐indigenous species (NIS); however, few studies have investigated the role of anthropogenic disturbance in facilitating the establishment and spread of NIS in marine environments. A baseline survey of native and NIS was undertaken in conjunction with a manipulative experiment to determine the effect that heavy metal pollution had on the diversity and invasibility of marine hard‐substrate assemblages. The study was repeated at two sites in each of two harbours in New South Wales, Australia. The survey sampled a total of 47 sessile invertebrate taxa, of which 15 (32%) were identified as native, 19 (40%) as NIS, and 13 (28%) as cryptogenic. Increasing pollution exposure decreased native species diversity at all study sites by between 33% and 50%. In contrast, there was no significant change in the numbers of NIS. Percentage cover was used as a measure of spatial dominance, with increased pollution exposure leading to increased NIS dominance across all sites. At three of the four study sites, assemblages that had previously been dominated by natives changed to become either extensively dominated by NIS or equally occupied by native and NIS alike. No single native or NIS was repeatedly responsible for the observed changes in native species diversity or NIS dominance at all sites. Rather, the observed effects of pollution were driven by a diverse range of taxa and species. These findings have important implications for both the way we assess pollution impacts, and for the management of NIS. When monitoring the response of assemblages to pollution, it is not sufficient to simply assess changes in community diversity. Rather, it is important to distinguish native from NIS components since both are expected to respond differently. In order to successfully manage current NIS, we first need to address levels of pollution within recipient systems in an effort to bolster the resilience of native communities to invasion.     

Native-species seed additions do not shift restored prairie plant communities from exotic to native states

Exotic- and native-dominated communities can exist as alternate states in landscapes, but whether exotic-dominated states are persistent in the face of propagule pressure from native species is not well known. Here, we asked whether adding native seeds to low diversity, exotic-dominated patches would shift them to a more diverse, native state by using a long-term experiment with tallgrass prairie species in Iowa, USA. Previous work established that community assembly history led to alternate exotic or native states of perennial species. We added native seeds to plots in the spring after removing aboveground biomass with fire. We found that an experimental seed addition did not cause a shift from exotic to native states. Plots seeded eight years earlier in spring and without a priority effect continued to have the highest abundance and diversity of native species and lowest proportion of exotics. Our results suggest that exotic-dominated states in restorations can persist in the face of native species propagule pressure. Thus, assembly history can play a strong role in generating and maintaining alternate states over long time frames that are relevant to restoration. New restoration projects in exotic-dominated landscapes should maximize effort toward establishing native species during initial stages of restoration.     

Geographical patterns of species turnover in aquatic plant communities

1. A classic theory in biogeography predicts that high latitude communities are unstable. This may be because of decreased species richness or decreased environmental predictability and productivity towards the poles.
2. We studied latitudinal patterns in long-term community persistence of aquatic vascular plants in 112 Finnish lakes, situated within a 1000-km range from the northernmost to the southernmost lake.
3. Contrary to theoretical predictions, we found that the turnover rate of plant species in 45 years was inversely related to latitude. That is, plant communities in northern lakes were more persistent than communities in southern lakes. When we used multiple regression to find the best predictors of species turnover rate (TR), latitude was the only variable that was highly significantly related to species turnover rate. Area, species number, water transparency, pH and change in transparency did not notably explain the gradient observed.
4. The latitudinal trend was mainly because of lower species immigration rates at higher latitudes, whereas extinction rate did not so strongly decrease with increasing latitude. Immigrations and extinctions in the lakes were not in balance: the species numbers between the 1930s and 1980s increased more strongly in the southern than northern lakes.
5. We suggest that the inverse relationship between latitude and plant species TR in Finland is most probably caused by human influence on lakes, especially eutrophication and immigration of new species in southern latitudes. In addition, although species richness per lake did not decrease towards the north, the total species pool probably does, which means that in the north there are fewer species that can actually immigrate.     

Phylogenetic beta diversity: linking ecological and evolutionary processes across space in time

A key challenge in ecological research is to integrate data from different scales to evaluate the ecological and evolutionary mechanisms that influence current patterns of biological diversity. We build on recent attempts to incorporate phylogenetic information into traditional diversity analyses and on existing research on beta diversity and phylogenetic community ecology. Phylogenetic beta diversity (phylobetadiversity) measures the phylogenetic distance among communities and as such allows us to connect local processes, such as biotic interactions and environmental filtering, with more regional processes including trait evolution and speciation. When combined with traditional measures of beta diversity, environmental gradient analyses or ecological niche modelling, phylobetadiversity can provide significant and novel insights into the mechanisms underlying current patterns of biological diversity.     

BIODIVERSITY RESEARCH: Experimental introduction of the alien plant Hieracium lepidulum reveals no significant impact on montane plant communities in New Zealand

Aim There is debate over whether alien plants necessarily alter the communities they invade or can coexist with native species without discernable impacts. We followed the fate of montane plant communities in response to the experimental sowing of the alien weed Hieracium lepidulum, looking for changes in plant community composition and structure over 6 years. Location Craigieburn Range, New Zealand. Methods We used a replicated randomised block design, with 30 × 30 cm plots (n = 756) subdivided into 5 × 5 cm cells to examine and compare the effects of H. lepidulum at 0.09 m² (plot) and 0.0025 m² (cell) scales. Plots were sown with between 0 and 15,625 H. lepidulum seeds in 2003, forming gradients of invader density and cover. Measurements comprised community richness, evenness and diversity along with H. lepidulum density and cover at both scales. The relationships between the invader and local community attributes were modelled using hierarchical mixed‐effect models. Results Plant communities differed in the extent to which they became invaded, with H. lepidulum cover in the plots ranging from 0% to 52%, with a mean of only 1.89%. Plot species richness increased from 2003 to 2009, with a component of this increase (+0.002 species per year) associated with increasing H. lepidulum density. Other relationships between the plant community and H. lepidulum were generally non‐significant. Main conclusions In these montane plant communities, it appears H. lepidulum coexists with the native community with no measurable negative effects after 6 years on species richness, evenness or diversity, even where density and cover of the invader are highest. We suggest H. lepidulum has persisted preferentially at those sites with abiotic conditions sufficient to support a species‐rich assemblage.     

The role of dominant prairie species ecotypes on plant diversity patterns of restored grasslands across a rainfall gradient in the US Great Plains

Questions

A robust ecosystem requires a functionally heterogeneous community of organisms with ecological traits that permit broad resource partitioning. Understanding community diversity patterns can help investigate drivers of community assembly and assess restoration success. Do biodiversity patterns differ among grassland communities sown with different ecotypes of dominant species during restoration along a rainfall gradient in the tallgrass prairie of the central US Great Plains?

Location

Four field sites across a rainfall gradient within the North American Great Plains: Colby, Kansas (39°23′17.8″N, 101°04′57.4″W), Hays, Kansas (38°51′13.2″N, 99°19′08.6″W), Manhattan, Kansas (39°08′22.3″N, 96°38′23.3″W), and Carbondale, Illinois (IL, 37°41′47.0″N, 89°14′19.2″W).

Methods

We applied linear mixed models to assess the effect of dominant species ecotype, year, and location on grassland taxonomic, phylogenetic, and functional diversity.

Results

The non-local grass ecotype (compared to the local ecotype) promoted species richness. In contrast, the effect of the dominant species ecotype on phylogenetic or functional diversity was site-specific over the 10-year restoration. Richness decreased across the rainfall gradient from dry to moist sites, and the wettest site had the highest phylogenetic and functional diversity.

Conclusions

Our results suggest that abiotic filtering by rainfall is a key assembly mechanism that could predict grassland changes in biodiversity in the early restoration phases. Given the community response across the tallgrass prairie, restoration practitioners should consider the impact of regional sources of dominant species used in restoration when biodiversity is a restoration goal. It is recommended for future grassland restoration to detect gaps and limitations in evolutionary and trait structure that will reveal which diversity components to evaluate.     

意大利苍耳入侵对本地植物群落物种多样性和稳定性的影响——以乌鲁木齐市为例

研究外来植物不同程度入侵对区域群落结构和植物群落物种多样性的影响对评价其对生态系统的危害具有重要意义。以入侵植物以意大利苍耳(Xanthium italicum)为主要研究对象,选择意大利苍耳危害较明显且分布面积较大的中国新疆乌鲁木齐市米东区为研究区域进行植物样方调查,以意大利苍耳和其共生的本地植物为切入点,以物种多样性指数、群落稳定性、群落可入侵性、影响程度指数、入侵植物的竞争优势和入侵强度来探究不同入侵程度下的本地植物群落物种多样性和群落稳定性的变化。结果表明:(1)与未入侵地相比,意大利苍耳轻度入侵显著增加了本地植物群落的物种多样性(P ＜ 0.05),而重度入侵显著降低了本地植物群落的物种多样性(P ＜ 0.05)。(2)在意大利苍耳入侵的群落中,群落稳定性由轻度入侵水平到重度入侵水平降低了25%,而群落可入侵性、意大利苍耳的竞争优势指数和入侵强度指数分别增加了77%、79%和83%。(3)在轻度入侵水平下,意大利苍耳对本植物群落物种多样性、群落稳定性和群落物种数的影响程度指数 ＜ 0,产生了积极影响;相反,在轻中度和重度入侵水平下意大利苍耳对本植物群落物种多样性、群落稳定性和群落物种数的影响程度指数 ＞ 0,产生了负面影响。(4)本地植物群落多样性与群落稳定性均呈正相关,但与群落可入侵性呈负相关(P ＜ 0.05)。(5)Shannon-Wiener指数对群落的稳定性的直接通经系数最大,为0.828;Simpson''s指数对群落可入侵性的直接通经系数最大,为0.932。由此可见,意大利苍耳在中度和重度入侵对植物群落结构产生巨大影响,而作为"一带一路"的核心和枢纽地带的新疆,将会对其脆弱的生态系统和土著物种的多样性造成不可估量的影响。应在入侵初期对其进行控制与铲除,防止意大利苍耳等有害外来入侵种的入侵和进一步扩散与蔓延。     

Variation in the impact of exotic grasses on native plant composition in relation to fire across an elevation gradient in Hawaii

The impact that an exotic species can have on the composition of the community it enters is a function of its abundance, its particular species traits and characteristics of the recipient community. In this study we examined species composition in 14 sites burned in fires fuelled by non‐indigenous C4 grasses in Hawaii Volcanoes National Park, Hawaii. We considered fire intensity, time since fire, climatic zone of site, unburned grass cover, unburned native cover and identity of the most abundant exotic grass in the adjacent unburned site as potential predictor variables of the impact of fire upon native species. We found that climatic zone was the single best variable for explaining variation in native cover among burned sites and between burned and unburned pairs. Fire in the eastern coastal lowlands had a very small effect on native plant cover and often stimulated native species regeneration, whereas fire in the seasonal submontane zone consistently caused a decline in native species cover and almost no species were fire tolerant. The dominant shrub, Styphelia tameiameia, in particular was fire intolerant. The number of years since fire, fire intensity and native cover in reference sites were not significantly correlated with native species cover in burned sites. The particular species of grass that carried the fire did however, have a significant effect on native species recovery. Where the African grass Melinis minutiflora was a dominant or codominant species, fire impacts were more severe than where it was absent regardless of climate zone. Overall, the impacts of exotic grass‐fuelled fires on native species composition and cover in seasonally dry Hawaiian ecosystems was context specific. This specificity is best explained by differences between the climatic zones in which fire occurred. Elevation was the main physical variable that differed among the climatic zones and it alone could explain a large percentage of the variation in native cover among sites. Rainfall, by contrast, did not vary systematically with elevation. Elevation is associated with differences in composition of the native species assemblages. In the coastal lowlands, the native grass Heteropogon contortus, was largely responsible for positive changes in native cover after fire although other native species also increased. Like the exotic grasses, this species is a perennial C4 grass. It is lacking in the submontane zone and there are no comparable native species there and almost all native species in the submontane zone were reduced by fire. The lack of fire tolerant species in the submontane zone thus clearly contributes to the devastating impact of fire upon native cover there.     

Biotic and abiotic constraints to a plant invasion in vegetation communities of Tierra del Fuego

The biotic resistance theory relates invader success to species richness, and predicts that, as species richness increases, invasibility decreases. The relationship between invader success and richness, however, seems to be positive at large scales of analysis, determined by abiotic constraints, and it is to be expected that it is negative at small scales, because of biotic interactions. Moreover, the negative relationship at small scales would be stronger within species of the same functional group, because of having similar resource exploitation mechanisms. We studied the relationship between the cover of a worldwide invader of grasslands, Hieracium pilosella L., and species richness, species diversity and the cover of different growth forms at two different levels of analysis in 128 sites during the initial invasion process in the Fuegian steppe, Southern Patagonia, Argentina. At regional level, the invader was positively correlated to total (r = 0.28, P = 0.003), exotic (r = 0.273, P = 0.004), and native species richness (r = 0.210, P = 0.026), and to species diversity (r = 0.193, P = 0.041). At community level, we found only a weak negative correlation between H. pilosella and total richness (r = ?0.426, P = 0.079) and diversity (r = ?0.658, P = 0.063). The relationship between the invader and other species of the same growth form was positive both at regional (r = 0.484, P < 0.001) and community (r = 0.593, P = 0.012) levels. Consequently, in the period of establishment and initial expansion of this exotic species, our results support the idea that invader success is related to abiotic factors at large scales of analysis. Also, we observed a possible sign of biotic constraint at community level, although this was not related to the abundance of species of the same growth form.     

The unseen invaders: introduced earthworms as drivers of change in plant communities in North American forests (a meta‐analysis)

Globally, biological invasions can have strong impacts on biodiversity as well as ecosystem functioning. While less conspicuous than introduced aboveground organisms, introduced belowground organisms may have similarly strong effects. Here, we synthesize for the first time the impacts of introduced earthworms on plant diversity and community composition in North American forests. We conducted a meta‐analysis using a total of 645 observations to quantify mean effect sizes of associations between introduced earthworm communities and plant diversity, cover of plant functional groups, and cover of native and non‐native plants. We found that plant diversity significantly declined with increasing richness of introduced earthworm ecological groups. While plant species richness or evenness did not change with earthworm invasion, our results indicate clear changes in plant community composition: cover of graminoids and non‐native plant species significantly increased, and cover of native plant species (of all functional groups) tended to decrease, with increasing earthworm biomass. Overall, these findings support the hypothesis that introduced earthworms facilitate particular plant species adapted to the abiotic conditions of earthworm‐invaded forests. Further, our study provides evidence that introduced earthworms are associated with declines in plant diversity in North American forests. Changing plant functional composition in these forests may have long‐lasting effects on ecosystem functioning.