Staged invasions across disparate grasslands: effects of seed provenance,consumers and disturbance on productivity and species richness

Exotic plant invasions are thought to alter productivity and species richness, yet these patterns are typically correlative. Few studies have experimentally invaded sites and asked how addition of novel species influences ecosystem function and community structure and examined the role of competitors and/or consumers in mediating these patterns. We invaded disturbed and undisturbed subplots in and out of rodent exclosures with seeds of native or exotic species in grasslands in Montana, California and Germany. Seed addition enhanced aboveground biomass and species richness compared with no‐seeds‐added controls, with exotics having disproportionate effects on productivity compared with natives. Disturbance enhanced the effects of seed addition on productivity and species richness, whereas rodents reduced productivity, but only in Germany and California. Our results demonstrate that experimental introduction of novel species can alter ecosystem function and community structure, but that local filters such as competition and herbivory influence the magnitude of these impacts.     

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.     

Native and alien plant species richness in relation to spatial heterogeneity on a regional scale in Germany

Aim The aim of our study was to reveal relationships between richness patterns of native vs. alien plant species and spatial heterogeneity across varying landscape patterns at a regional scale. Location The study was carried out in the administrative district of Dessau (Germany), covering around 4000 km². Methods Data on plant distribution of the German vascular flora available in grid cells covering 5′ longitude and 3′ latitude (c. 32 km²) were divided into three status groups: native plants, archaeophytes (pre 1500 AD aliens) and neophytes (post 1500 AD aliens). Land use and abiotic data layers were intersected with 125 grid cells comprising the selected area. Using novel landscape ecological methods, we calculated 38 indices of landscape composition and configuration for each grid cell. Principal components analysis (PCA) with a set of 29 selected, low correlated landscape indices was followed by multiple linear regression analysis. Results PCA reduced 29 indices to eight principal components (PCs) that explained 80% cumulative variance. Multiple linear regression analysis was highly significant and explained 41% to 60% variance in plant species distribution (adjusted R²) with three significant PCs (tested for spatial autocorrelation) expressing moderate to high disturbance levels and high spatial heterogeneity. Comparing the significance of the PCs for the species groups, native plant species richness is most strongly associated with riverine ecosystems, followed by urban ecosystems, and then small‐scale rural ecosystems. Archaeophyte and neophyte richness are most strongly associated with urban ecosystems, followed by small‐scale rural ecosystems and riverine ecosystems for archaeophytes, and riverine ecosystems and small‐scale rural ecosystems for neophytes. Main conclusions Our overall results suggest that species richness of native and alien plants increases with moderate levels of natural and/or anthropogenic disturbances, coupled with high levels of habitat and structural heterogeneity in urban, riverine, and small‐scale rural ecosystems. Despite differences in the order of relevance of PCs for the three plant groups, we conclude that at the regional scale species richness patterns of native plants as well as alien plants are promoted by similar factors.     

Impact of Oreochromis niloticus (Linnaeus, 1758) (Pisces: Cichlidae) invasion on taxonomic and functional diversity of native fish species in the upper Kabompo River,northwest of Zambia

Invasive alien species have been revealed to drastically alter the structure of native communities; however, there is scarce information on whether taxonomic and functional spaces occupied by native species are equally filled by exotic species. We investigated the diversity of native species to understand the impact of exotic Oreochromis niloticus in the upper Kabompo River, northwest of Zambia using taxonomic and functional diversity indices. To achieve this, two tests were performed (Test 1, compared natives in invaded and uninvaded sections; Test 2, compared natives in invaded section). A total of 17 species were collected for functional diversity computation, out of which fourteen (14) functional trait measurements linked to feeding, locomotion, and life history strategy were taken. Findings revealed that taxonomic and functional diversity values changed with invasion in both tests. Taxonomic diversity was 15% more in invaded than uninvaded sections in Test 1 and was not consistent across sampling points of invaded section in Test 2. Invaded areas were taxonomically less diverse, but functionally diverse in both tests. The analysis of similarity and nonmetric multidimensional scaling revealed no difference in Bray–Curtis similarity assemblages in both tests. Our findings revealed that exotic species more often occupy unfilled gaps in the communities often occupied by the native species; this is achieved by occupying functional spaces. Overall, changes in taxonomic and functional diversity of native species documented here partially confirmed impacts of O. niloticus invasion. Therefore, we recommend a multifaceted approach to assess cumulative impacts of invasion on native species.     

Species richness and phylogenetic diversity of native and non‐native species respond differently to area and environmental factors

Aim

To test whether native and non‐native species have similar diversity–area relationships (species–area relationships [SARs] and phylogenetic diversity–area relationships [PDARs]) and whether they respond similarly to environmental variables.

Location

United States.

Methods

Using lists of native and non‐native species as well as environmental variables for >250 US national parks, we compared SARs and PDARs of native and non‐native species to test whether they respond similarly to environmental conditions. We then used multiple regressions involving climate, land cover and anthropogenic variables to further explore underlying predictors of diversity for plants and birds in US national parks.

Results

Native and non‐native species had different slopes for SARs and PDARs, with significantly higher slopes for native species. Corroborating this pattern, multiple regressions showed that native and non‐native diversity of plants and birds responded differently to a greater number of environmental variables than expected by chance. For native species richness, park area and longitude were the most important variables while the number of park visitors, temperature and the percentage of natural area were among the most important ones for non‐native species richness. Interestingly, the most important predictor of native and non‐native plant phylogenetic diversity, temperature, had positive effects on non‐native plants but negative effects on natives.

Main conclusions

SARs, PDARs and multiple regressions all suggest that native and non‐native plants and birds responded differently to environmental factors that influence their diversity. The agreement between diversity–area relationships and multiple regressions with environmental variables suggests that SARs and PDARs can be both used as quick proxies of overall responses of species to environmental conditions. However, more importantly, our results suggest that global change will have different effects on native and non‐native species, making it inappropriate to apply the large body of knowledge on native species to understand patterns of community assembly of non‐native species.

     

Contrasting alien and native plant species–area relationships: the importance of spatial grain and extent

Aim The proportion of alien plant species in floras is increasingly being used to indicate the threat of invasions to native species and/or the homogenization of biodiversity. However, this indicator is only valuable if it is independent of the spatial extent and grain of observation. This study tested the equivalence of native and alien species–area relationships (SARs) in order to assess the support for scale invariance in the proportion of alien species in floras. Location England, UK. Methods Nested SARs were generated by assessing the richness of native and alien plant species drawn from the New atlas of the British and Irish flora for six areas comprising 100, 400, 900, 1600, 2500 and 3600 km² with each larger area containing all smaller areas. Five replicate sets of nested areas encompassing northern, southern, eastern, western and central regions were chosen. For each set of nested areas, the log‐transformed species richness was regressed on log‐transformed area to fit a power function to the SAR. Results Native and alien plant SARs reveal consistent differences in slope, highlighting that the proportion of alien species is a function of spatial grain. Aliens are more rare than natives and have higher spatial turnover leading to faster accumulation of species as area increases. However, equivalent samples drawn from a larger spatial extent reveal similar alien and native SARs. Main conclusions The significant differential scale dependence in native and alien species richness observed in this study reflects dissimilar influences of regional drivers such as habitat, but potentially also propagule pressure and introduction history, that leads to the relative rarity and high spatial turnover of alien species. Maps of invasion hotspots that identify areas where the proportion of the alien flora is particularly high should therefore be treated with considerable caution since patterns across most grains used for species monitoring will be scale dependent.     

Assessing the vulnerability of riparian vegetation to invasion by Mimulus guttatus: relative importance of biotic and abiotic variables in determining species occurrence and abundance

Riparian habitats are particularly susceptible to invasion by non‐native plants. At present, attempts to build consensus as to what the primary drivers of plant invasion in riparian ecosystems might be is hindered by the absence of common standards for data collected on plant species (e.g. occurrence, or relative abundance). Mimulus guttatus L., a non‐native riparian plant species, was used as a model to determine how environmental drivers influence two aspects of invasibility: species occurrence and abundance (assessed in relation to three variables number of patches, patch area and number of stems per patch). Mimulus occurrence and abundance, together with 20 environmental variables, were surveyed in almost 700 contiguous 50‐m‐long riverbank segments within a catchment in north‐east Scotland. More than half of the segments had been colonized by Mimulus. Occurrence and number of patches responded to similar environmental gradients, particularly bare sediment, boulders, high soil moisture, short‐statured ruderal communities, and open canopies, and tended to be highest downstream where the river was widest. In contrast to occurrence and patch number, patch area and stem number per patch were higher in the upper reaches of the catchment and were positively associated with low tree canopy and vegetation dominated by light‐demanding species and smaller‐statured species. Patch area and stem number per patch were also positively related to grazing. This study has highlighted the importance of assessing more than one measure of invasion success (occurrence or patch number and either patch area or stem number per patch), as they are each determined by a different suite of environmental variables. Abiotic factors, such as sediment availability and presence of boulders, appeared to be the major determinants of occurrence and patch number, whereas biotic factors, such as interspecific competition and grazing, were more important ecological determinants underlying area and stem number per patch.     

Plants on small islands revisited: the effects of spatial scale and habitat quality on the species–area relationship

Understanding how species diversity is related to sampling area and spatial scale is central to ecology and biogeography. Small islands and small sampling units support fewer species than larger ones. However, the factors influencing species richness may not be consistent across scales. Richness at local scales is primarily affected by small‐scale environmental factors, stochasticity and the richness at the island scale. Richness at whole‐island scale, however, is usually strongly related to island area, isolation and habitat diversity. Despite these contrasting drivers at local and island scales, island species–area relationships (SARs) are often constructed based on richness sampled at the local scale. Whether local scale samples adequately predict richness at the island scale and how local scale samples influence the island SAR remains poorly understood. We investigated the effects of different sampling scales on the SAR of trees on 60 small islands in the Raja Ampat archipelago (Indonesia) using standardised transects and a hierarchically nested sampling design. We compared species richness at different grain sizes ranging from single (sub)transects to whole islands and tested whether the shape of the SAR changed with sampling scale. We then determined the importance of island area, isolation, shape and habitat quality at each scale on species richness. We found strong support for scale dependency of the SAR. The SAR changed from exponential shape at local sampling scales to sigmoidal shape at the island scale indicating variation of species richness independent of area for small islands and hence the presence of a small‐island effect. Island area was the most important variable explaining species richness at all scales, but habitat quality was also important at local scales. We conclude that the SAR and drivers of species richness are influenced by sampling scale, and that the sampling design for assessing the island SARs therefore requires careful consideration.     

Species pools,community completeness and invasion: disentangling diversity effects on the establishment of native and alien species

Invasion should decline with species richness, yet the relationship is inconsistent. Species richness, however, is a product of species pool size and biotic filtering. Invasion may increase with richness if large species pools represent weaker environmental filters. Measuring species pool size and the proportion realised locally (completeness) may clarify diversity‐invasion relationships by separating environmental and biotic effects, especially if species’ life‐history stage and origin are accounted for. To test these relationships, we added seeds and transplants of 15 native and alien species into 29 grasslands. Species pool size and completeness explained more variation in invasion than richness alone. Although results varied between native and alien species, seed establishment and biotic resistance to transplants increased with species pool size, whereas transplant growth and biotic resistance to seeds increased with completeness. Consequently, species pools and completeness represent multiple independent processes affecting invasion; accounting for these processes improves our understanding of invasion.     

The importance of rare species: a trait‐based assessment of rare species contributions to functional diversity and possible ecosystem function in tall‐grass prairies

The majority of species in ecosystems are rare, but the ecosystem consequences of losing rare species are poorly known. To understand how rare species may influence ecosystem functioning, this study quantifies the contribution of species based on their relative level of rarity to community functional diversity using a trait‐based approach. Given that rarity can be defined in several different ways, we use four different definitions of rarity: abundance (mean and maximum), geographic range, and habitat specificity. We find that rarer species contribute to functional diversity when rarity is defined by maximum abundance, geographic range, and habitat specificity. However, rarer species are functionally redundant when rarity is defined by mean abundance. Furthermore, when using abundance‐weighted analyses, we find that rare species typically contribute significantly less to functional diversity than common species due to their low abundances. These results suggest that rare species have the potential to play an important role in ecosystem functioning, either by offering novel contributions to functional diversity or via functional redundancy depending on how rare species are defined. Yet, these contributions are likely to be greatest if the abundance of rare species increases due to environmental change. We argue that given the paucity of data on rare species, understanding the contribution of rare species to community functional diversity is an important first step to understanding the potential role of rare species in ecosystem functioning.     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.