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.     

Scale,disturbance and productivity control the native‐exotic richness relationship

The relationship between native and exotic species richness may be highly context‐dependent. Spatial scale, including both plot size (grain) and study area (extent), is likely to influence this relationship, as are environmental conditions such as resource availability and disturbance intensity. We used experimental manipulations of soil fertility and disturbance in a California coastal grassland to directly examine the sensitivity of the native–exotic richness relationship (NERR) to these factors across five grain sizes and two spatial extents. The slope of the NERR was a function of grain size, extent and treatment. Over small spatial extents, native and exotic richness were usually uncorrelated. Across a larger extent, NERRs were negative in control plots, neutral in disturbance plots, and positive in plots with experimentally reduced soil fertility. These patterns were strongest for small grain sizes. We verify the importance of spatial grain in determining the NERR, and emphasize the role of spatial extent.     

Relationship between plant species richness and biomass in a coastal Maine Quercus-Pinus forest

Abstract. Several researchers have hypothesized that plant species richness has a unimodal relationship with biomass, while others have argued for a linear relationship. Data from various types of herbaceous communities show some support for the unimodal hypothesis, but this has not been tested extensively for forests and questions remain concerning its generality. We used linear and quadratic regression models to examine the relationship between overstory biomass and richness in a coastal Maine Quercus-Pinus forest across and within cover types using data from two plot sizes (2500-m² quadrats and 625-m² sub-quadrats). Understory data from 1-m² plots were also analysed. Richness was quadratically related to biomass at both plot sizes for all cover types combined, but the amount of variation explained by the models was very low (R²s < 0.09). Richness and biomass were not significantly related at either plot size for the mixed mesic cover type, the most common type in the forest. The best fit (R²= 0.43) was obtained with a quadratic model for the conifer cover type at the sub-quadrat level, with the quadratic model for the 1-m² data having the second highest R² (0.24). Across all six data sets, the quadratic model was the only one with a significant fit in two cases, and had considerably higher R²s (1.3–1.9 ×) in two others. The remaining two data sets could not be fit with a significant model of either type. For this forest, these results suggest little support for a linear relationship between plant species richness and biomass and variable, often weak, support for a unimodal relationship. Density, a potentially confounding variable in this type of analysis, was only weakly correlated with richness and was not found to alter the relationship between biomass and richness.     

Relationship between plant species richness and biomass in an arid sub-alpine grassland of the central Himalayas,Nepal

The hump-shaped relationship between plant species richness and biomass is commonly observed at fine scale for herbaceous vegetation in temperate climates. This relationship predicts that herbaceous species richness is highest at an intermediate level of biomass that corresponds to moderate competition or disturbance. However, this relationship has not previously been investigated in high arid sub-alpine mountain grasslands. We tested the humped-back prediction in the arid Trans-Himalayan mountain grassland with a seasonal grazing system. The study area is located in the bottom of a U-shaped valley, in the Manang district (3500 m a.s.l.). We sampled two hundred plots (1m × 1m) in two different types of pastures: common pasture and old field, which both have similar grazing practices. There was a significant unimodal relationship between species richness and biomass only in the common pasture, and when the two sites were analyzed together. The species turnover is estimated by DCA in standard deviation unit. The turnover was lower in the old field than in the common pasture. The unimodal relationship between plant species richness and biomass did not disappear after accounting for unknown environmental gradients expressed as DCA (detrended correspondence analysis) axes and spatial variables. The species richness is highest at 120 ± 40 g/m². The results indicate that a hump-shaped relationship is also found in arid Trans-Himalayan grasslands.     

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.     

Linear declines in exotic and native plant species richness along an increasing altitudinal gradient in the Snowy Mountains,Australia

Abstract Declines in plant species richness with increasing altitude are common, but the form of the relationship can vary, with both monotonic decreasing relationships and humped relationship recorded. However, these different richness to altitude relationships may be due to methods that used different plot sizes/areas and survey efforts. To explore native and exotic plant richness along an altitudinal gradient in the Snowy Mountains of Australia, we consistently surveyed plots that were 120 m² in area at 39 sites ranging from 540 to 2020 m. To relate exotic plant richness to disturbance, we surveyed plots at 16 sites along main roads and 23 sites along minor roads and also compared these 39 roadside plots to 120‐m² plots located in undisturbed vegetation adjacent to the roadside (native plant richness was only surveyed in 25 of these 39 adjacent plots). We found a negative linear relationship between total, exotic and native species richness and altitude for plots on the side of main roads (16 sites) and minor roads (23 sites). For adjacent plots negative linear relationships were significant for all measures of species richness except for native species adjacent to major roads. As the pattern occurred for exotics it is less likely to be due to historical constraints on the species pools. The pattern could be influenced by difference in levels of disturbance along the gradient, although any such gradient in disturbance would have to apply to roadside and adjacent plots on major and minor roads. Therefore, it may be due to other factors such as changes in climate along the altitudinal gradient, although additional sampling including direct measures of climatic conditions, soil and disturbance factors would be needed to determine if this was the case.     

Grazing effects on species richness depends on scale: a 5-year study in Tenerife pastures (Canary Islands)

The effect of herbivores on species richness is important for the conservation of protected areas under grazing management but research findings on this are far from consistent. The main objective of this study is to analyze how the cessation of grazing by goats affects the diversity parameters at different scales over a 5-year period. The study was conducted in the Teno Rural Park in the northwest corner of Tenerife, Canary Islands. The studied areas have been grazed by livestock since the beginning of the 16th century and currently are used by local farmers, mainly for extensive goat production. In these areas we selected four blocks and in each block four 100 m² plots were established, two excluded from grazing (located in 12 × 12 m² exclosures) and two control plots. The analyses showed an accumulation of species in the control plots significantly higher than in exclosure plots at small scales. Power function parameters such as c and z only showed differences in function for the sampling year and not for the treatment. Although the results showed an increase in species richness due to grazing, this is very small. However, negative effects in native species richness are not detected, so we suggest the promotion of goat grazing as a way to maintain land use, cultural values, and species richness in these pastures.     

Grazing and an invasive grass confound spatial pattern of exotic and native grassland plant species richness

Previous work has shown exotic and native plant species richness are negatively correlated at fine spatial scales and positively correlated at broad spatial scales. Grazing and invasive plant species can influence plant species richness, but the effects of these disturbances across spatial scales remain untested. We collected species richness data for both native and exotic plants from five spatial scales (0.5–3000 m²) in a nested, modified Whittaker plot design from severely grazed and ungrazed North American tallgrass prairie. We also recorded the abundance of an abundant invasive grass, tall fescue (Schedonorus phoenix (Scop.) Holub), at the 0.5-m² scale. We used linear mixed-effect regression to test relationships between plant species richness, tall fescue abundance, and grazing history at five spatial scales. At no scale was exotic and native species richness linearly related, but exotic species richness at all scales was greater in grazed tracts than ungrazed tracts. Native species richness declined with increasing tall fescue abundance at all five spatial scales, but exotic species richness increased with tall fescue abundance at all but the broadest spatial scales. Severe grazing did not reduce native species richness at any spatial scale. We posit that invasion of tall fescue in this working landscape of originally native grassland plants modifies species richness-spatial scale relationships observed in less disturbed systems. Tall fescue invasion constitutes a unique biotic effect on plant species richness at broad spatial scales.     

Multi-scale comparisons of cliff vegetation in Colorado

We developed a nested vegetation sampling protocol to sample the plant diversity on south-facing cliffs and cliff bases in Jefferson County, Colorado. The multi-scale plots included three nested spatial scales, 1 m², 20 m², and 40 m². We compared plant species richness and species diversity among large cliffs, medium cliffs, small cliffs, and non-cliff sites using Hill's diversity numbers (N ₀, N ₁, and N ₂) for the 1-m² quadrats. Species richness (N ₀) was calculated for the 20-m² and 40-m² plots. Our results indicate that plant species diversity on the cliff faces did not increase with increasing cliff area. This pattern was consistent at all three sampling scales. A model selection was run to determine if plant species diversity values on the cliff faces were associated with cliff variables. None of the cliff variables measured were good predictors of diversity at the 1-m² scale. However, at the 20-m² scale, canyon differences and a positive relationship with increasing cliff surface roughness explained 70% of the variability in species richness. Although most plant species sampled on the cliff faces were also found in the base plots, 13 species were sampled only on the cliff faces. Additionally, dry south facing cliffs support a mix of xeric and mesic plants indicating that cliffs may provide unique microenvironments for plant establishment. This revised version was published online in August 2006 with corrections to the Cover Date.     

Increasing native, but not exotic, biodiversity increases aboveground productivity in ungrazed and intensely grazed grasslands

Species-rich native grasslands are frequently converted to species-poor exotic grasslands or pastures; however, the consequences of these changes for ecosystem functioning remain unclear. Cattle grazing (ungrazed or intensely grazed once), plant species origin (native or exotic), and species richness (4-species mixture or monoculture) treatments were fully crossed and randomly assigned to plots of grassland plants. We tested whether (1) native and exotic plots exhibited different responses to grazing for six ecosystem functions (i.e., aboveground productivity, light interception, fine root biomass, tracer nitrogen uptake, biomass consumption, and aboveground biomass recovery), and (2) biodiversity-ecosystem functioning relationships depended on grazing or species origin. We found that native and exotic species exhibited different responses to grazing for three of the ecosystem functions we considered. Intense grazing decreased fine root biomass by 53% in exotic plots, but had no effect on fine root biomass in native plots. The proportion of standing biomass consumed by cattle was 16% less in exotic than in native grazed plots. Aboveground biomass recovery was 30% less in native than in exotic plots. Intense grazing decreased aboveground productivity by 25%, light interception by 14%, and tracer nitrogen uptake by 54%, and these effects were similar in native and exotic plots. Increasing species richness from one to four species increased aboveground productivity by 42%, and light interception by 44%, in both ungrazed and intensely grazed native plots. In contrast, increasing species richness did not influence biomass production or resource uptake in ungrazed or intensely grazed exotic plots. These results suggest that converting native grasslands to exotic grasslands or pastures changes ecosystem structure and processes, and the relationship between biodiversity and ecosystem functioning.     

The relationship between species richness and biomass changes from boreal to subtropical forests in China

Diversity‐manipulation experiments suggest a positive effect of biodiversity on ecosystem properties (EPs), but variable relationships between species richness and EPs have been reported in natural communities. An explanation for this discrepancy is that observed richness–EPs relationships in natural communities change with environment and species functional identities. But how the relationships change along broad‐scale climatic gradients has rarely been examined. In this paper, we sampled 848 plots of 20 × 30 m² from boreal to tropical forests across China. We examined plot biomass with respect to environmental factors, tree species richness and functional group identity (FGI, i.e. evergreen vs deciduous, and coniferous vs broadleaf). Variation partitioning was used to evaluate the relative effects of the three classes of factors. We found that, most of the ‘effects’ (percentage of variation explained) of richness and FGI on forest biomass were shared with environmental factors, but species richness and FGI still revealed significant effects in addition to environment for plots across China. Richness and FGI explained biomass mainly through their shared effects instead of independent effects, suggesting that the positive biodiversity effect is closely associated with a sampling effect. The relative effects of richness, FGI and environment varied latitudinally: the independent effects of environment and richness decreased from boreal to subtropical forests, whereas the total effect of FGI increased. We also found that the slope of richness–biomass relationship decreased monotonically from boreal to subtropical forests, possibly because of decreasing complementarity and increasing competition with increasing productivity. Our results suggest that while species richness does have significant effects on forest biomass it is less important than environmental gradients and other biotic factors in shaping large‐scale biomass patterns. We suggest that understanding how and why the diversity–EPs relationships change along climatic gradient would be helpful for a better understanding of real biodiversity effects in natural communities.     

Small-scale species richness and its spatial variation in an alpine meadow on the Qinghai-Tibet Plateau

We investigated how the high small-scale species richness of an alpine meadow on the Qinghai-Tibet Plateau, China, is maintained. This area is characterized by strong wind and severe cold during long winters. In winter, most livestock is grazed on dead leaves in small pastures near farmers’ residences, whereas in the short summer, livestock is grazed in mountainous areas far from farmers’ residences. The number of plant species and the aboveground biomass were surveyed for three adjacent pastures differing in grazing management: a late-winter grazing pasture grazed moderately from 1 February to 30 April, an early-winter grazing pasture grazed lightly from 20 September to late October, and a whole-year grazing pasture grazed intensively throughout the entire year. In each pasture, we harvested the aboveground biomass from 80 or 100 quadrats of 0.01 m² along a transect and classified the contents by species. We observed 15.5–19.7 species per 0.01 m², which is high richness per 0.01 m² on a worldwide scale. The species richness in the two winter grazing pastures was higher than that in the whole-year grazing pasture. The spatial variation in species richness and species composition in the two winter grazing pastures in which species richness was high was greater than that in the whole-year grazing pasture in which species richness was lower. Most of the leaves that are preserved on the winter grazing pastures during summer are blown away by strong winds during winter, and the remaining leaves are completely exhausted in winter by livestock grazing. A pasture with a high richess is accompanied by a high spatial variation in species richness and species composition. There is a high possibility that the characteristic of spatial variation is also caused by traditional grazing practices in this area.     

Human impact diminishes seedling species richness in Kakamega Forest, Kenya

Anthropogenic forest fragmentation and other kinds of human disturbance, such as selective logging, can reduce the diversity of plant and animal species. To evaluate the impact of fragmentation and small-scale disturbance on forest regeneration, we assessed species richness and total abundance of adult trees in comparison with seedlings in the heavily fragmented and disturbed Kakamega Forest, western Kenya. In nine differently disturbed 1-ha study blocks distributed across the main forest and fragments, we mapped all trees >10 cm in diameter at breast height. Additionally, we established ninety 1-m² seedling plots within these 1-ha study blocks which were monitored over 2.5 years. We recorded altogether 74 species of adult trees (30–43 per block) and 64 seedling species (24–41 per block). Neither fragmentation nor small-scale disturbance had an impact on adult tree species richness or total tree abundance. Yet, fragmentation and especially small-scale disturbance significantly reduced seedling species richness, particularly of late-successional species. While human impact did not affect diversity of adults, the impoverished species richness of seedlings suggests a reduced potential for regeneration and a loss of tree diversity in the long-term.     

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.     

Competitive dominance mediates the effects of topography on plant richness in a mountain grassland

Small-scale landforms influence plant species richness, but their mechanisms and effects in semi-natural dry grasslands have been poorly investigated. In this study we compared vascular plant richness, species composition, plant traits, soil properties and biomass nutrient content of convex (hillocks) and concave (hollows) karst landforms in a mountain pasture of the Central Apennines (Italy), at a small spatial scale (1 m² plots). We found hillocks had significantly higher species richness than hollows. On hillocks, smaller Specific Leaf Area and Lateral Width, together with greater allocation of resources to Below-Ground Organs, indicated lower water availability, whereas hollows had deeper (thus moister), more acidic and more fertile soils, with aboveground plant biomass displaying higher nutrient levels. Partial correlation and regression tree models suggested that fine-scale richness patterns were not directly determined by abiotic properties, but were rather the result of competition levels associated with the cover of Agrostis capillaris (=A. tenuis) – a calcifuge and drought-sensitive grass able to achieve dominance only in hollows. The higher functional convergence exhibited by hollows suggests that A. capillaris is a strong competitor both above- and below-ground, mediating the effects of topography by imposing a biotic filter. On hillocks, competition is released by lower levels of available soil water in summer and higher soil pH, resulting in higher species richness and a more functionally divergent assemblage.     

Grain-dependent relationships between plant productivity and invertebrate species richness and biomass in calcareous grasslands

The relationships among productivity, species richness and consumer biomass are of fundamental importance for understanding determinants of biodiversity. These relationships may depend on grain size. We examined the relationships between productivity (above-ground phytomass) and plant species richness and between productivity and species richness and biomass of gastropods and grasshoppers using sampling units of different sizes (0.5, 2.75 and 23 m²) in nutrient-poor, calcareous grasslands in north-western Switzerland in two successive years. Species richness of forbs had a unimodal relationship with productivity in sampling units of 0.5 m² and was negatively correlated with productivity at the other two plot sizes in one year. In the other year, forb species richness tended to decrease with productivity in sampling units of 23 m². No similar relationship was found for grasses. Gastropod biomass had a unimodal relationship with productivity at 0.5 m² in the first year. Grasshopper species richness was correlated with forb species richness at plot sizes of 2.75 and 23 m². This study demonstrates that patterns detected between productivity and diversity and between productivity and biomass of consumers depend on the grain size used in the investigation and vary among years.Die Zusammenhänge zwischen Produktivität, Artenreichtum und Biomasse von Konsumenten sind wichtig, um zu verstehen, was Biodiversität beeinflußt. Diese Zusammenhänge können von der Größe der Untersuchungsfläche abhängig sein. Wir untersuchten während zwei aufeinanderfolgenden Jahren die Zusammenhänge zwischen Produktivität (oberirdische Pflanzenbiomasse) und Artenreichtum von Gefäßpflanzen, sowie zwischen Produktivität und Artenreichtum und Biomasse von Schnecken und Heuschrecken bezüglich dreier räumlicher Skalen (0,5, 2,75 und 23 m²) in Kalkmagerrasen in der Nordwestschweiz. Der Zusammenhang zwischen dem Artenreichtum von Kräutern und der Produktivität war unimodal in Flächeneinheiten von 0,5 m² und negativ in Flächeneinheiten von 2,75 und 23 m² im ersten Jahr und war tendenziell negativ in Flächeneinheiten von 23 m² im zweiten Jahr, während kein solcher Zusammenhang bei Gräsern gefunden wurde. Der Zusammenhang zwischen Produktivität und Biomasse von Schnecken war unimodal in Flächeneinheiten von 0,5 m² im ersten Jahr. Außerdem bestand ein Zusammenhang zwischen dem Artenreichtum von Kräutern und Heuschrecken in Flächeneinheiten von 2,75 und 23 m². Diese Arbeit zeigt, daß Zusammenhänge zwischen Produktivität und Diversität sowie zwischen Produktivität und Biomasse von Konsumenten von der Größe der Untersuchungsfläche abhängen und zwischen Jahren variieren.     

Altitude modifies species richness–nutrient indicator value relationships in a country-wide survey of grassland vegetation

Nutrient enrichment is a threat to botanical diversity in Europe, and its assessment is part of biodiversity monitoring schemes. In Switzerland, this is done by calculating the average nutrient (N) indicator value of the vegetation based on a country-wide systematic vegetation survey. However, it is questionable whether N values indicate eutrophication and resulting species loss equally well across an entire country, which includes wide topographic gradients and distinct biogeographic regions. Here we analyze vascular plant species lists from 415 grassland plots (10 m²) between 365 and 2770 m a.s.l. throughout Switzerland to investigate how the relationship between N value and species richness differs with altitude and among regions. The N value strongly decreased with altitude (piecewise regression: r² = 0.77), particularly between 800 and 2000 m a.s.l., where this decrease was related to a decreasing proportion of fertilized grasslands. In the alpine belt, lower N values were associated with a greater frequency of acidic soils and a restricted species pool. Vascular plant species richness was maximal at intermediate altitude (piecewise regression: r² = 0.33) and intermediate N value (polynomial regression: r² = 0.46). When analyzed separately by altitudinal belt, the relationship between species richness and N value was negative in the lowlands and montane belt but unimodal in the subalpine belt. In the alpine belt, soil pH (R indicator values) explained most of the variation in species richness. Two indices of between-plot diversity (floristic dissimilarity and the contribution of individual plots to total species richness) were negatively related to N values from the lowlands to the subalpine belt but not in the alpine belt. All relationships differed little among the biogeographic regions of Switzerland, but they might be modified by changes in management and by the expansion of common lowland species into mountain grasslands.     

Seasonal changes in the relationship between plant species richness and community biomass in early succession

We analysed the relationship between plant species richness and productivity on first-year-old fields at two similar sites in central Europe. At both sites, a wide range of productivity levels was available resulting from different long-term fertilisation. In order to identify underlying mechanisms of the species richness–productivity relationship we included the seasonal dynamics and the number of individuals of each species in our analysis. We sampled 10 and 21 plots, respectively, at the two sites in May, June and July by harvesting all aboveground parts of vascular plants in 0.25 m² subplots. Species richness, number of individuals of each species and community biomass as a surrogate of productivity were recorded in each sample.At one site, the relationship between species richness and biomass was significantly positive in the May and June harvest. This relationship disappeared in the July harvest due to a reduction in species richness at high productivity levels. The relations between species richness and number of individuals and between number of individuals and biomass paralleled the species richness–productivity relation but the individual number–biomass relationship remained positive until the last harvest. Between-species differences in individual number–community biomass relationships and their seasonal dynamics revealed “interspecific competitive exclusion” even though the species richness–biomass relationships were not negative or hump-shaped. At the second site, species richness was not related to productivity or to number of individuals. Our study demonstrated the importance of temporal dynamics and regional processes in understanding species richness–productivity patterns.