A nested-intensity design for surveying plant diversity

Managers of natural landscapes need cost-efficient, accurate, and precise systems to inventory plant diversity. We investigated a nested-intensity sampling design to assess local and landscape-scale heterogeneity of plant species richness in aspen stands in southern Colorado, USA. The nested-intensity design used three vegetation sampling techniques: the Modified-Whittaker, a 1000-m² multiple-scale plot (n = 8); a 100-m² multiple-scale Intensive plot (n = 15); and a 100-m² single-scale Extensive plot (n = 28). The large Modified-Whittaker plot (1000 m²) recorded greater species richness per plot than the other two sampling techniques (P < 0.001), estimated cover of a greater number of species in 1-m² subplots (P < 0.018), and captured 32 species missed by the smaller, more numerous 100-m² plots of the other designs. The Intensive plots extended the environmental gradient sampled, capturing 17 species missed by the other techniques, and improved species–area calculations. The greater number of Extensive plots further expanded the gradient sampled, and captured 18 additional species. The multi-scale Modified-Whittaker and Intensive designs allowed quantification of the slopes of species–area curves in the single-scale Extensive plots. Multiple linear regressions were able to predict the slope of species–area curves (adj R ² = 0.64, P < 0.001) at each Extensive plot, allowing comparison of species richness at each sample location. Comparison of species–accumulation curves generated with each technique suggested that small, single-scale plot techniques might be very misleading because they underestimate species richness by missing locally rare species at every site. A combination of large and small multi-scale and single-scale plots greatly improves our understanding of native and exotic plant diversity patterns.     

Scale-dependent relationships between the spatial distribution of a limiting resource and plant species diversity in an African grassland ecosystem

One cornerstone of ecological theory is that nutrient availability limits the number of species that can inhabit a community. However, the relationship between the spatial distribution of limiting nutrients and species diversity is not well established because there is no single scale appropriate for measuring variation in resource distribution. Instead, the correct scale for analyzing resource variation depends on the range of species sizes within the community. To quantify the relationship between nutrient distribution and plant species diversity, we measured NO₃^- distribution and plant species diversity in 16 paired, modified Whittaker grassland plots in Serengeti National Park, Tanzania. Semivariograms were used to quantify the spatial structure of NO₃^- from scales of 0.4–26 m. Plant species diversity (Shannon-Weiner diversity index; H ) was quantified in 1-m² plots, while plant species richness was measured at multiple spatial scales between 1 and 1,000 m². Small-scale variation in NO₃^- (<0.4 m) was positively correlated with 1-m² H , while 1,000-m² species richness was a log-normal function of average NO₃^- patch size. Nine of the 16 grassland plots had a fractal (self-similar across scales) NO₃^- spatial distribution; of the nine fractal plots, five were adjacent to plots that had a non-fractal distribution of NO₃^-. This finding offered the unique opportunity to test predictions of Ritchie and Olff (1999): when the spatial distribution of limiting resources is fractal, communities should display a left-skewed log-size distribution and a log-normal relationship between net primary production and species richness. These predictions were supported by comparisons of plant size distributions and biomass-richness relationships in paired plots, one with a fractal and one with a non-fractal distribution of NO₃^-. In addition, fractal plots had greater large-scale richness than paired non-fractal plots (1,0–1000 m²), but neither species diversity (H ) nor richness was significantly different at small scales (1 m²). This result is most likely explained by differences in the scale of resource variation among plots: fractal and non-fractal plots had equivalent NO₃^- variation at small scales but differed in NO₃^- variation at large scales (as measured by the fractal dimension). We propose that small-scale variation in NO₃^- is largely due to the direct effects of plants on soil, while patterns of species richness at large scales is controlled by the patch size and fractal dimension of NO₃^- in the landscape. This study provides an important empirical step in understanding the relationship between the spatial distribution of resources and patterns of species diversity across multiple spatial scales.     

Are Plant Censuses Carried Out on Small Quadrats More Reliable than on Larger Ones?

Plant censuses are known to be significantly affected by observers’ biases. In this study, we checked whether the magnitude of observer effects (defined as the % of total variance) varied with quadrat size: we expected the census repeatability (% of the total variance that is not due to measurement errors) to be higher for small quadrats than for larger ones. Variations according to quadrat size of the repeatability of species richness, Simpson equitability and reciprocal diversity indices, Ellenberg indicator values, plant cover and plant frequency were assessed using 359 censuses of vascular plants. These were carried out independently by four professional botanists during spring 2002 on the same 18 forest plots, each comprising one 400-m² quadrat, four 4-m² and four 2-m² quadrats. Time expenditure was controlled for. General Linear Models using random effects only were applied to the ecological indices to estimate variance components and magnitude of the following effects (if possible): plot, quadrat, observer, plant species and two-way interactions. High repeatability was obtained for species richness and Ellenberg indicator values. Species richness and Ellenberg indicator values were generally more accurate but also more biased in large quadrats. Simpson reciprocal diversity and equitability indices were poorly repeatable (especially equitability) probably because plant cover estimates varied widely among observers, irrespective of quadrat size. Grouping small quadrats usually increased the repeatability of the variable considered (e.g. species richness, Simpson diversity, plant cover) but the number of plant species found on those pooled 16 m² was much lower than if large plots were sampled. We therefore recommend to use large, single quadrats for forest vegetation monitoring.     

The Historic Square Foot Dataset – Outstanding small-scale richness in Swiss grasslands around the year 1900

Grasslands host a significant share of Europe's species diversity but are among the most threatened vegetation types of the continent. Resurvey studies can help to understand patterns and drivers of changes in grassland diversity and species composition. However, most resurveys are based on local or regional data, and hardly reach back more than eight decades. Here, we publish and describe the Historic Square Foot Dataset, comprising 580 0.09-m² and 43 1-m² vegetation plots carefully sampled between 1884 and 1931, covering a wide range of grassland types across Switzerland. We provide the plots as an open-access data set with coordinates, relocation accuracy and fractional aboveground biomass per vascular plant species. We assigned EUNIS habitat types to most plots. Mean vascular plant species richness in 0.09 m² was 19.7, with a maximum of 47. This is considerably more than the present-day world record of 43 species for this plot size. Historically, species richness did not vary with elevation, differing from the unimodal relationship found today. The data set provides unique insight into how grasslands in Central Europe looked more than 100 years ago, thus offering manifold options for studies on the development of grassland biodiversity and productivity.     

N<Subscript>2</Subscript> fixation in three perennial <Emphasis Type="Italic">Trifolium</Emphasis> species in experimental grasslands of varied plant species richness and composition

This study is the first to investigate quantitative effects of plant community composition and diversity on N₂ fixation in legumes. N₂ fixation in three perennial Trifolium species grown in field plots with varied number of neighbouring species was evaluated with the ¹⁵N natural abundance method (two field sites, several growing seasons, no N addition) and the isotope dilution method (one site, one growing season, 5 g N m⁻²). The proportion of plant N derived from N₂ fixation, pNdfa, was generally high, but the N addition decreased pNdfa, especially in species-poor communities. Also following N addition, the presence of grasses in species-rich communities increased pNdfa in T. hybridum and T. repens L., while legume abundance had the opposite effect. In T. repens, competition for light from grasses appeared to limit growth and thereby the amount of N₂ fixed at the plant level, expressed as mg N₂ fixed per sown seed. We conclude that the occurrence of diversity effects seems to be largely context dependent, with soil N availability being a major determinant, and that species composition and functional traits are more important than species richness regarding how neighbouring plant species influence N₂ fixation in legumes.     

Grain size affects the relationship between species richness and above-ground biomass in semi-arid rangelands

ABSTRACT

Background: Discrepancies in the shape of the productivity–diversity relationship may arise from differences in spatial scale. We hypothesised that there is a grain size effect on the productivity–diversity relationship.

Aims: To determine the effect of three sampling grain sizes on the productivity–diversity relationship.

Methods: We applied generalised linear mixed effect models on community data from 735 vegetation plots in the Taleghan rangelands, Iran, sampled at three grain sizes (0.25, 1 and 2 m²) to ascertain plant productivity-diversity patterns, while accounting for the effects of site, plant community type, disturbance, and life form.

Results: Overall, relationships between biomass and plant species richness were unimodal at grain sizes of 0.25 and 1 m², and asymptotical at 2 m². The spurious occurrence of a single large shrub may overwhelm a small-sized sampling unit, resulting in a high estimate of the sample’s biomass relative to species richness. However, the relationship between biomass and species richness at larger grain sizes is more likely to reach an asymptote.

Conclusions: Shrubs are partly responsible for driving the relationship between plant biomass and species richness. Given that the frequency of shrubs is highly variable between small plots but not so in large plots, their presence may result in unimodal productivity–diversity relationships at small but not at large grain sizes.     

Plant species richness and diversity along an altitudinal gradient in the Sierra Nevada, Mexico

This article presents an analysis of plant species richness and diversity and its association with climatic and soil variables along a 1300‐m elevation gradient on the Cerro Tláloc Mountain in the northern Sierra Nevada in Mexico. Two 1000‐m² tree sampling plots were created at each of 21 selected sampling sites, as well as two 250‐m² plots for shrubs and six 9‐m² plots for herbaceous plants. Species richness and diversity were estimated for each plant life form, and beta diversity between sites was estimated along the gradient. The relationship between species richness and diversity and environmental variables was modelled using simple linear correlation and regression trees. Species richness and diversity showed a unimodal pattern with a bias towards high values in the lower half of the elevation gradient under study. This response was consistent for all three life forms. Beta diversity increased steadily along the elevation gradient, being lower between contiguous sites at intermediate elevations and high – the species replacement rate was nearly 100%– between sites at the extremes of the gradient. Few species were adapted to the full spectrum of environmental variation along the elevation gradient studied. The regression tree suggests that differences in species richness are mainly influenced by elevation (temperature and humidity) and soil variables, namely A₂ permanent wilting point, organic matter and horizon field capacity and A₁ horizon Mg²⁺.     

A Modified-Whittaker nested vegetation sampling method

A standardized sampling technique for measuring plant diversity is needed to assist in resource inventories and for monitoring long-term trends in vascular plant species richness. The widely used Whittaker plot (Shmida 1984) collects species richness data at multiple spatial scales, using 1 m², 10 m², and 100 m² subplots within a 20 m × 50 m (1000 m²) plot, but it has three distinct design flaws involving the shape and placement of subplots. We modified and tested a comparable sampling design (Modified-Whittaker plot) that minimizes the problems encountered in the original Whittaker design, while maintaining many of its attractive attributes. We overlaid the two sampling methods in forest and prairie vegetation types in Larimer County, Colorado, USA (n=13 sites) and Wind Cave National Park, South Dakota, USA (n=19 sites) and showed that the modified design often returned significantly higher (p<0.05) species richness values in the 1 m², 10 m², and 100 m² subplots. For all plots, except seven ecotone plots, there was a significant difference (p<0.001) between the Whittaker plot and the Modified-Whittaker plot when estimating the total number of species in the 1000 m² plots based on linear regressions of the subplot data: the Whittaker plot method, on average, underestimated plant species richness by 34%. Species-area relationships, using the Modified-Whittaker design, conformed better to published semilog relationships, explaining, on average, 92% of the variation. Using the original Whittaker design, the semilog species-area relationships were not as strong, explaining only 83% of the variation, on average. The Modified-Whittaker plot design may allow for better estimates of mean species cover, analysis of plant diversity patterns at multiple spatial scales, and trend analysis from monitoring a series of strategically-placed, long-term plots.     

Would adding scallop shells (Chlamys islandica) to the sea bottom enhance recruitment of commercial species?

We examined the impact of adding scallop shells (byproduct of the fisheries) to sandy and rocky sea bottoms in the northern Gulf of St. Lawrence. The effect of adding shells was greatest on sandy bottoms where species richness increased 3.7-fold and species diversity 1.9-fold. The increase in most species was due to immigration rather than new settlement. Trials examining the effect of different densities of shells in plots of the same size (4 m²) showed that species diversity increased rapidly with shell abundance and levelled off when shells covered half of the bottom, whereas species richness only levelled off when shells almost completely covered the bottom. Trials examining the effect of the size of the shell patches (shell density being kept constant) showed that species diversity was already maximal in 1-m² plots, whereas species richness only attained a plateau at 4 m². Our small-scale trials indicate that the addition of shells would have a positive impact, increasing numerous invertebrates, including commercial species (scallops, whelks and urchins).     

Invasion of an intact plant community: the role of population versus community level diversity

To improve the understanding of how native plant diversity influences invasion, we examined how population and community diversity may directly and indirectly be related to invasion in a natural field setting. Due to the large impact of the dominant C₄ grass species (Andropogon gerardii) on invasion resistance of tallgrass prairie, we hypothesized that genetic diversity and associated traits within a population of this species would be more strongly related to invasion than diversity or traits of the rest of the community. We added seeds of the exotic invasive C₄ grass, A. bladhii, to 1-m² plots in intact tallgrass prairie that varied in genetic diversity of A. gerardii and plant community diversity, but not species richness. We assessed relationships among genetic diversity and traits of A. gerardii, community diversity, community aggregated traits, resource availability, and early season establishment and late-season persistence of the invader using structural equation modeling (SEM). SEM models suggested that community diversity likely enhanced invasion indirectly through increasing community aggregated specific leaf area as a consequence of more favorable microclimatic conditions for seedling establishment. In contrast, neither population nor community diversity was directly or indirectly related to late season survival of invasive seedlings. Our research suggests that while much of diversity–invasion research has separately focused on the direct effects of genetic and species diversity, when taken together, we find that the role of both levels of diversity on invasion resistance may be more complex, whereby effects of diversity may be primarily indirect via traits and vary depending on the stage of invasion.     

Effects of tree species diversity and genotypic diversity on leafminers and parasitoids in a tropical forest plantation

相似文献   

Microtopographic heterogeneity constrains alpine plant diversity, Glacier National Park, MT

Theoretical and empirical evidence exists for a positive relationship between environmental heterogeneity and species diversity. Alpine plant communities can exhibit exceptional diversity at a fine scale, which niche theory would suggest is the result of fine scale spatial heterogeneity of the environment. To test if species diversity of alpine plants is driven by environmental heterogeneity, we sampled vascular plant species composition, microtopography, and ground cover within 1?m² plots with and without solifluction forms in Glacier National Park, MT. We analyzed the relationship between microtopographic heterogeneity and species richness at the plot and sub-plot scale with linear and quantile regression, respectively. Species richness does not differ between the plots varying in cover type. Species richness is negatively related to the fractal dimension (D) of the ground surface and non-vegetated ground cover within 1?m² plots. At a finer scale, the standard deviation of elevation and slope appear to impose a limit on species richness such that more variable sub-plots have lower species richness. Contrary to our expectations, microtopographic heterogeneity does not promote the diversity of alpine plants. The negative relationship between topographic heterogeneity and species richness is contrary to the theoretical prediction that environmental heterogeneity generally results in greater species diversity. It is possible that microtopographic variability represents a measure of soil disturbance, which would be expected to have a negative effect on species diversity in alpine tundra due to its low productivity.     

Incorporating functional dissimilarities into sample‐based rarefaction curves: from taxon resampling to functional resampling

Question: Indices of functional diversity have been seen as the key for integrating information on species richness with measures that focus on those components of community composition related to ecosystem functioning. For comparing species richness among habitats on an equal‐effort basis, so‐called sample‐based rarefaction curves may be used. Given a study area that is sampled for species presence and absence in N plots, sample‐based rarefaction generates the expected number of accumulated species as the number of sampled plots increases from 1 to N. Accordingly, the question for this study is: can we construct a ‘functional rarefaction curve’ that summarizes the expected functional dissimilarity between species when n plots are drawn at random from a larger pool of N plots? Methods: In this paper, we propose a parametric measure of functional diversity that is obtained by combining sample‐based rarefaction techniques that are usually applied to species richness with Rao's quadratic diversity. For a given set of N presence/absence plots, the resulting measure summarizes the expected functional dissimilarity at an increasingly larger cumulative number of plots n (n≤N). Results and Conclusions: Due to its parametric nature, the proposed measure is progressively more sensitive to rare species with increasing plot number, thus rendering this measure adequate for comparing the functional diversity of species assemblages that have been sampled with variable effort.     

A flexible multi-scale approach for standardised recording of plant species richness patterns

A sound monitoring of appropriate biodiversity indicators is necessary in order to assess the progress towards the internationally agreed target of halting the loss of biodiversity by 2010. However, existing monitoring schemes often do not address species richness as a key component of biodiversity directly or do so with insufficient methods. I provide an overview and assessment of the large variety of different sampling approaches for small-scale plant species richness. Major shortcomings of many of these are (i) non-uniform plot sizes or shapes; (ii) analysis of only one spatial scale despite the scale dependence of nearly all biodiversity parameters; (iii) lack of replication of smaller subplots; and (iv) exclusion of bryophytes and lichens despite their often large contribution to total plant diversity. Based on this review, I propose a new standardised sampling approach for plant diversity patterns at small scales that is applicable for a multitude of purposes and in any biome. In its basic variant, species composition is recorded on nested squares of 0.01 m², 0.1 m², 1 m², 10 m², and 100 m², with all smaller subplots being replicated at least 3-fold and evenly spaced within the next larger plot. Not only terricolous vascular plants, but also bryophytes, lichens, macro-algae as well as non-terricolous taxa should be recorded with the any-part system, i.e. those plants are counted within a plot whose superficial parts reach over it. This approach can be used to assess plant diversity patterns (i) of individual plots of interest, (ii) along environmental gradients, (iii) within specific vegetation types, or (iv) for landscape sectors. In the latter case, the series of nested plots must be placed randomly or systematically, but irrespective of plot homogeneity. The proposed approach allows the calculation of many meaningful biodiversity indicators, while being well compatible with a range of other sampling schemes, but avoiding their shortcomings. As this approach is not very time-consuming in its basic variant, but can easily be extended for specific purposes, I suggest its use for any kind of biodiversity studies and particularly for monitoring.     

Changes in the understory of Canadian southern boreal forest after fire

Abstract. We investigated changes in the composition and abundance of understory species after fire in the southern boreal forest around Lake Duparquet, Québec. Ten plots of 100 m² were sampled in each of eight sites varying in post-fire age from 26 to 230 yr, with 20 1-m² quadrats in each of these 80 plots. Variation in the understory was described by DCA ordination and interpreted as a regeneration succession series. Thickness of the organic layers, stand age and canopy composition were all correlated with vegetational change. This change was not constant throughout succession; some old sites showed an increase in the diversity and abundance of certain pioneer species. This was partly related to openings in the canopy resulting from a major outbreak of spruce budworm, which affected sites dominated by Abies balsamea. The ordinations were performed on both the 100-m² plots and the 1-m² quadrats. Heterogeneity within sites was larger at the 1-m² scale and there was a great deal of overlap in the position of the quadrats in ordination space. At the smaller scale of analysis, stand age and thickness of the organic layers were not correlated with the changes observed in the understory.     

Spatial heterogeneity of soil nutrients and plant species in herb-dominated communities of contrasting land use

Recent interest in spatial pattern in terrestrial ecosystems has come from an awareness of the intimate relationship between spatial heterogeneity of soil resources and maintenance of plant species diversity. Soil and vegetation can vary spatially in response to several state factors of the system. In this study, we examined fine-scale spatial variability of soil nutrients and vascular plant species in contrasting herb-dominated communities (a pasture and an old field) to determine degree of spatial dependence among soil variables and plant community characteristics within these communities by sampling at 1-m intervals. Each site was divided into 25 1-m² plots. Mineral soil was sampled (2-cm diameter, 5-cm depth) from each of four 0.25-m² quarters and combined into a single composite sample per plot. Soil organic matter was measured as loss-on-ignition. Extractable NH₄ and NO₃ were determined before and after laboratory incubation (28 days at 27°C) to determine potential net N mineralization and nitrification. Cations were analyzed using inductively coupled plasma emission spectrometry. Vegetation was assessed using estimated percent cover. Most soil and plant variables exhibited sharp contrasts between pasture and old-field sites, with the old field having significantly higher net N mineralization/nitrification, pH, Ca, Mg, Al, plant cover, and species diversity, richness, and evenness. Multiple regressions revealed that all plant variables (species diversity, richness, evenness, and cover) were significantly related to soil characteristics (available nitrogen, organic matter, moisture, pH, Ca, and Mg) in the pasture; in the old field only cover was significantly related to soil characteristics (organic matter and moisture). Both sites contrasted sharply with respect to spatial pattern of soil variables, with the old field exhibiting a higher degree of spatial dependence. These results demonstrate that land-use practices can exert profound influence on spatial heterogeneity of both soil properties and vegetation in herb-dominated communities.     

Larger patches of diverse floral resources increase insect pollinator density,diversity, and their pollination of native wildflowers

Native wildflower plantings can be used to provide nutritional resources to support pollinating insects, yet the effects of planting size and bloom richness on the density, diversity, and function of these insects are not well understood. We established stands of twelve native flowering perennial plant species in replicated plots ranging in size from 1 to 100 m². These plots were sampled for insect pollinators, bloom richness, and seed production by three wildflower species. Honeybees, wild bees, and hoverflies all responded positively to increasing flower richness, whereas particular insect pollinator groups responded differently to the size of the flowering plant area. The density of honeybees and hoverflies was not affected by increasing flowering patch size, whereas in general, wild bees were observed at higher density and diversity in the 30 and 100 m² patches. Increasing wildflower patch size, and thus wild bee density, resulted in greater seed set in the sampled wildflowers. These results indicate that wild bees are sensitive to the area and richness of floral resources in patches, even at relatively small scales. Therefore, larger wildflower plantings with more diverse flower species mixes are more suitable for the conservation of wild pollinators and reproduction of sown species.     

Diversity of forest plant species at the community and landscape scales in Switzerland

Abstract

Conservation strategies increasingly refer to indicators derived from large biological data. However, such data are often unique with respect to scale and species groups considered. To compare richness patterns emerging from different inventories, we analysed forest species richness at both the landscape and the community scales in Switzerland. Numbers of forest species were displayed using nationwide distributional species data and referring to three different definitions of forest species. The best regression models on a level of four predictor variables ranged between adj. R ² = 0.50 and 0.66 and revealed environmental heterogeneity/energy, substrate (rocky outcrops) and precipitation as best explanatory variables of forest species richness at the landscape scale. A systematic sample of community data (n = 729; 30 m², 200 m², 500 m²) was examined with respect to nationwide community diversity and plot species richness. More than 50% of all plots were assigned to beech forests (Eu-Fagion, Cephalanthero-Fagion, Luzulo-Fagion and Abieti-Fagion), 14% to Norway spruce forests (Vaccinio-Piceion) and 13% to silver fir forests (Piceo-Abietion). Explanatory variables were derived from averaged indicator values per plot, and from biophysical and disturbance factors. The best models for plot species richness using four predictor variables ranged between adj. R ² = 0.31 and 0.34. Light (averaged L-indicator, tree canopy) and substrate (averaged R-indicator and pH) had the highest explanatory power at all community scales. By contrast, the influence of disturbance variables was very small, as only a small portion of plots were affected by this factor. The effects of disturbances caused by extreme events or by management would reduce the tree canopies and lead to an increase in plant species richness at the community scale. Nevertheless, such community scale processes will not change the species richness at the landscape scale. Instead, the variety of different results derived from different biological data confirms the diversity of aspects to consider. Therefore, conservation strategies should refer to value systems.     

Loss of plant biodiversity eliminates stimulatory effect of elevated CO2 on earthworm activity in grasslands

Earthworms are among the world’s most important ecosystem engineers because of their effects on soil fertility and plant productivity. Their dependence on plants for carbon, however, means that any changes in plant community structure or function caused by rising atmospheric CO₂ or loss of plant species diversity could affect earthworm activity, which may feed back on plant communities. Production of surface casts measured during three consecutive years in field experimental plots (n = 24, 1.2 m²) planted with local calcareous grassland species that varied in plant species richness (diversity levels: high, 31 species; medium, 12; low, 5) and were exposed to ambient (356 μl CO₂ l^?1) or elevated (600 μl CO₂ l^?1) CO₂ was only consistently stimulated in high diversity plots exposed to elevated CO₂ (+120 %, 31 spp: 603 ± 52 under ambient CO₂ vs. 1,325 ± 204 g cast dwt. m^?2 year^?1 under elevated CO₂ in 1996; +77 %, 940 ± 44 vs. 1,663 ± 204 g cast dwt. m^?2 year^?1 in 1998). Reductions in plant diversity had little effect on cast production in ecosystems maintained at ambient CO₂, but the stimulatory effect of elevated CO₂ on cast production disappeared when plant species diversity was decreased to 12 and 5 species. High diversity plots were also the only communities that included plant species that an earlier field study showed to be among the most responsive to elevated CO₂ and to be most preferred by earthworms to deposit casts near. Further, the +87 % CO₂-induced increase in cast production measured over the 3 years corresponded to a parallel increase in cumulative total nitrogen of 5.7 g N m^?2 and would help explain the large stimulation of aboveground plant biomass production observed in high-diversity communities under elevated CO₂. The results of this study demonstrate how the loss of plant species from communities can alter responses of major soil heterotrophs and consequently ecosystem biogeochemistry.     

Soil nutrients and variation in biomass rather than native species richness influence introduced plant richness in a semi-arid grassland

Several different hypotheses account for the success of introduced species in new environments. Experimental studies show a negative native-exotic richness relationship (NERR), while observational studies suggest that this relationship is usually positive. Increased resource availability and environmental variation can also enable introduced species to establish in new environments. We conducted an observational study in a semi-arid grassland in the Thompson-Nicola District of British Columbia to examine the biotic and abiotic factors that account for variation in introduced and native species richness.In each of 12 sites, an 8 × 8 m area was set up, containing 64, 1-m² plots. We identified and categorized plant species in each site into introduced and native species. We tested the relationship between introduced species richness and native species richness at the 1-m² sampling grain and at sampling grains up to 64 m². We also analysed the relationship between native and introduced species, and within-plot biomass, and between native and introduced species and variation in biomass. For a representative subset of four sites, we tested the relationship between introduced and native species richness and nitrogen, phosphorus and potassium.We found no NERR at the 1 m² sampling grain, nor for the other sampling grains up to 64 m². Introduced species richness increased with phosphorus and nitrogen availability, and was also positively related to biomass heterogeneity.Our results indicate that introduced species richness in these grasslands is likely influenced by phosphorus and nitrogen, and by variation in vegetation biomass, but not by native species. More non-native plants are likely to occupy nutrient-rich plots compared to nutrient-poor plots in these grasslands. Variation in biomass can leave gaps for the establishment of introduced species. These results should inform management considerations for the control of invasive species to optimize preservation of grasslands.