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.     

Shape matters in sampling plant diversity: Evidence from the field

The identification of shape and size of sampling units that maximises the number of plant species recorded in multiscale sampling designs has major implications in conservation planning and monitoring actions. In this paper we tested the effect of three sampling shapes (rectangles, squared, and randomly shaped sampling units) on the number of recorded species. We used a large dataset derived from the network of protected areas in the Siena Province, Italy. This dataset is composed of plant species occurrence data recorded from 604 plots (10 m × 10 m), each divided in a grid of 16 contiguous subplot units (2.5 m × 2.5 m). Moreover, we evaluated the effect of plot orientation along the main environmental gradient, to examine how the selection of plot orientation (when elongated plots are used) influences the number of species collected. In total, 1041 plant species were recorded from the study plots. A significantly higher species richness was recorded by the random arrangement of 4 subplots within each plot in comparison to the ‘rectangle’ and ‘square’ shapes. Although the rectangular shape captured a significant larger number of species than squared ones, plot orientation along the main environmental gradient did not show a systematic effect on the number of recorded species. We concluded that the choice of whether or not using elongated (rectangular) versus squared plots should dependent upon the objectives of the specific survey with squared plots being more suitable for assessing species composition of more homogeneous vegetation units and rectangular plots being more suited for recording more species in the pooled sample of a large area.     

Effect of inventory plot patterns in the floristic analysis of tropical woodland and dense forest

This study was set up to examine the effect of plot patterns on the accuracy of phytosociological characterization of tropical vegetation. Fifteen and twenty square plots of 1 ha were demarcated, respectively, in woodland and dense forest in Bénin. Each 1 ha plot was divided into 100 quadrats of one 100 m². Species of trees in each quadrat were identified and recorded. The cost in terms of time required to record tree species in each 1 ha plot and five random quadrats in a 1 ha plot were also recorded to compute the mean inventory effort for a team of three foresters. From the 100 quadrats in a 1 ha plot, fourteen independent subplots of square and rectangular plots with different sizes were considered by grouping together adjacent quadrats of 100 m². Eigenanalysis was carried out to compare the subplots. Moreover, the relationship between the relative loss of accuracy (RLA) and the size of subplots was modelled. Plot size highly influenced the RLA (P < 0.05). Findings indicated that the square plots of 1500 and 1000 m² with an inventory effort of 0.35 and 0.20 man‐days per subplot, respectively in tropical dense forests and woodlands appeared to be the most efficient in the phytosociological characterization of woody vegetation.     

Native and naturalized plant diversity are positively correlated in scrub communities of California and Chile

Abstract. An emerging body of literature suggests that the richness of native and naturalized plant species are often positively correlated. It is unclear, however, whether this relationship is robust across spatial scales, and how a disturbance regime may affect it. Here, I examine the relationships of both richness and abundance between native and naturalized species of plants in two mediterranean scrub communities: coastal sage scrub (CSS) in California and xeric-sloped matorral (XSM) in Chile. In each vegetation type I surveyed multiple sites, where I identified vascular plant species and estimated their relative cover. Herbaceous species richness was higher in XSM, while cover of woody species was higher in CSS, where woody species have a strong impact upon herbaceous species. As there were few naturalized species with a woody growth form, the analyses performed here relate primarily to herbaceous species. Relationships between the herbaceous cover of native and naturalized species were not significant in CSS, but were nearly significant in XSM. The herbaceous species richness of native and naturalized plants were not significantly correlated on sites that had burned less than one year prior to sampling in CSS, and too few sites were available to examine this relationship in XSM. In post 1-year burn sites, however, herbaceous richness of native and naturalized species were positively correlated in both CSS and XSM. This relationship occurred at all spatial scales, from 400 m² to 1 m² plots. The consistency of this relationship in this study, together with its reported occurrence in the literature, suggests that this relationship may be general. Finally, the residuals from the correlations between native and naturalized species richness and cover, when plotted against site age (i.e. time since the last fire), show that richness and cover of naturalized species are strongly favoured on recently burned sites in XSM; this suggests that herbaceous species native to Chile are relatively poorly adapted to fire.     

Pteridophyte diversity and species composition in four Amazonian rain forests

Abstract. Local variation in individual density, species composition, species richness and species diversity of terrestrial pteridophytes were studied at four sites in the tropical lowland rain forest of western Amazonia. 15 568 pteridophyte individuals representing 40 species were recorded in four plots. The variability among subplots within the same plot was considerable in all the characteristics measured (number of individuals, number of species, species diversity); the square 1‐ha plot was more homogeneous in these respects than any of the three 5 m by 1300 m transects. Species richness was affected by the density of individuals both within and among plots. Density of individuals was not affected by topographical position within any of the plots, whereas in some of the plots both species richness and species diversity were. Clustering and ordination analyses showed that floristically similar subplots could be found in different plots: although there was a tendency for subplots from the same plot to be floristically similar and therefore to group together, many recognized groups included subplots from two or more plots. Both within and among plots, the floristic differences corresponded to topographic position and were probably related to soil drainage. This was also evident in that the abundance patterns of many species followed the topography.     

Riparian zones as havens for exotic plant species in the central grasslands

In the Central Grasslands of the United States, we hypothesized that riparian zones high in soil fertility would contain more exotic plant species than upland areas of low soil fertility. Our alternate hypothesis was that riparian zones high in native plant species richness and cover would monopolize available resources and resist invasion by exotic species. We gathered nested-scale vegetation data from 40 1 m²subplots (nested in four 1000 m² plots) in both riparian and upland sites at four study areas in Colorado, Wyoming, and South Dakota (a total of 320 1 m² subplots and 32 1000 m² plots). At the 1 m² scale, mean foliar cover of native species was significantly greater (P<0.001) in riparian zones (36.3% ± 1.7%) compared to upland sites (28.7% ± 1.5%), but at this small scale there were no consistent patterns of native and exotic species richness among the four management areas. Mean exotic species cover was slightly higher in upland sites compared to riparian sites (9.0% ± 3.8% versus 8.2% ± 3.0% cover). However, mean exotic species richness and cover were greater in the riparian zones than upland sites in three of four management areas. At the 1000 m² scale, mean exotic species richness was also significantly greater (P<0.05) in riparian zones (7.8 ± 1.0 species) compared to upland sites (4.8 ± 1.0 species) despite the heavy invasion of one upland site. For all 32 plots combined, 21% of the variance in exotic species richness was explained by positive relationships with soil % silt (t =1.7, P=0.09) and total foliar cover (t = 2.4, P=0.02). Likewise, 26% of the variance in exotic species cover (log₁₀ cover) was explained by positive relationships with soil % silt (t =2.3, P=0.03) and total plant species richness (t = 2.5, P=0.02). At landscape scales (four 1000 m² plots per type combined), total foliar cover was significantly and positively correlated with exotic species richness (r=0.73, P<0.05) and cover (r=0.74, P<0.05). Exotic species cover (log₁₀ cover) was positively correlated with log₁₀% N in the soil (r=0.61, P=0.11) at landscape scales. On average, we found that 85% (±5%) of the total number of exotic species in the sampling plots of a given management area could be found in riparian zones, while only 50% (±8%) were found in upland plots. We conclude that: (1) species-rich and productive riparian zones are particularly invasible in grassland ecosystems; and (2) riparian zones may act as havens, corridors, and sources of exotic plant invasions for upland sites and pose a significant challenge to land managers and conservation biologists.     

Species–area relationships in Mediterranean-climate plant communities

Aim To determine the best‐fit model of species–area relationships for Mediterranean‐type plant communities and evaluate how community structure affects these species–area models. Location Data were collected from California shrublands and woodlands and compared with literature reports for other Mediterranean‐climate regions. Methods The number of species was recorded from 1, 100 and 1000 m² nested plots. Best fit to the power model or exponential model was determined by comparing adjusted r² values from the least squares regression, pattern of residuals, homoscedasticity across scales, and semi‐log slopes at 1–100 m² and 100–1000 m². Dominance–diversity curves were tested for fit to the lognormal model, MacArthur's broken stick model, and the geometric and harmonic series. Results Early successional Western Australia and California shrublands represented the extremes and provide an interesting contrast as the exponential model was the best fit for the former, and the power model for the latter, despite similar total species richness. We hypothesize that structural differences in these communities account for the different species–area curves and are tied to patterns of dominance, equitability and life form distribution. Dominance–diversity relationships for Western Australian heathlands exhibited a close fit to MacArthur's broken stick model, indicating more equitable distribution of species. In contrast, Californian shrublands, both postfire and mature stands, were best fit by the geometric model indicating strong dominance and many minor subordinate species. These regions differ in life form distribution, with annuals being a major component of diversity in early successional Californian shrublands although they are largely lacking in mature stands. Both young and old Australian heathlands are dominated by perennials, and annuals are largely absent. Inherent in all of these ecosystems is cyclical disequilibrium caused by periodic fires. The potential for community reassembly is greater in Californian shrublands where only a quarter of the flora resprout, whereas three quarters resprout in Australian heathlands. Other Californian vegetation types sampled include coniferous forests, oak savannas and desert scrub, and demonstrate that different community structures may lead to a similar species–area relationship. Dominance–diversity relationships for coniferous forests closely follow a geometric model whereas associated oak savannas show a close fit to the lognormal model. However, for both communities, species–area curves fit a power model. The primary driver appears to be the presence of annuals. Desert scrub communities illustrate dramatic changes in both species diversity and dominance–diversity relationships in high and low rainfall years, because of the disappearance of annuals in drought years. Main conclusions Species–area curves for immature shrublands in California and the majority of Mediterranean plant communities fit a power function model. Exceptions that fit the exponential model are not because of sampling error or scaling effects, rather structural differences in these communities provide plausible explanations. The exponential species–area model may arise in more than one way. In the highly diverse Australian heathlands it results from a rapid increase in species richness at small scales. In mature California shrublands it results from very depauperate richness at the community scale. In both instances the exponential model is tied to a preponderance of perennials and paucity of annuals. For communities fit by a power model, coefficients z and log c exhibit a number of significant correlations with other diversity parameters, suggesting that they have some predictive value in ecological communities.     

Do vine species in neotropical forests see the forest or the trees?

Questions: Is the occurrence of vine species in neotropical rain forests primarily determined by properties of the forest (environmental factors), by properties of the trees (tree species or tree size) or are vines randomly distributed? Location: Maya Biosphere Reserve, Guatemala. Methods: In five 1‐ha plots that span variation from unlogged forest to forest impacted by recurrent human disturbance we recorded the presence of all climbing vine species on every tree. The presence of all free standing vine species and 11 environmental variables were recorded in 100‐m² subplots. The relationship of host tree diameter and host tree identity on single tree vine species richness was investigated by GLM modelling. Partial redundancy analyses were used to partition the variation in vine species composition on two sources: environmental factors and tree species identity. Results: Single tree vine richness increased with increasing host tree DBH and differed significantly among host species. For climbing vines, the ratio of variation in subplot presence explained by tree species and by environmental variables was ca. 4:1 (in the most disturbed logged plots slightly lower), for free standing vines this ratio varied from 1:2 in the most disturbed logged plots to 9:1 in reserve plots, while a ratio of ca. 1:1 was found for all plots analysed together. Conclusion: Different tree species have different probabilities of being infested by vines. Vines see both the forest and the trees; the environment is more important in earlier developmental stages, properties of individual trees become more important from the time vines start to climb.     

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.     

Space-time heterogeneity in the recovery after experimental burning and cutting in a Cistus laurifolius shrubland

Recovery after experimental burning and cutting in a shrubland of Cistus laurifolius in NW Spain has been studied. The community was homogeneous prior to the disturbances, and tended to recover through a process of autosuccession. It was tested whether in a small space (two 100 m² plots) there was a greater similarity among individual subplots (1²) in five consecutive years, or among the five subplots considered in each plot in the same year. By comparing space and time beta diversity using analysis of variance, no significant differences were observed, which indicates that temporal changes are not of a greater magnitude than space heterogeneity, even on such a small scale. Changes in time are characterized by an increase in cover by woody species, mainly Cistus laurifolius, or a decrease in the diversity and richness of species. Space heterogeneity (differences between subplots) does not seem to be determined by environmental gradients, since the sampling surface is very small, and may be due to the effect of some annual or perennial species, which are not dominant and only appear in some subplots, probably due to random dispersal.     

Sampling Techniques Influence Understory Plant Trajectories After Restoration: An Example from Ponderosa Pine Restoration

Abstract Although there is no one correct technique for sampling vegetation, the sampling design chosen may greatly influence the conclusions researchers can draw from restoration treatments. Considerations when designing vegetation sampling protocol include determining what sampling attributes to measure, the size and shape of the sampling plot, the number of replicates and their location within the study area, and the frequency of sampling. We installed 20 point‐intercept transects (50‐m long), 8 belt transects (10 × 50 m), 10 adapted Daubenmire transects (four 0.5 × 2‐m plots), and 4 modified‐Whittaker plots (20 × 50 m with smaller nested plots) in treatment and control units to measure understory herbaceous response in a forest restoration experiment that tested different treatments. Point‐intercept transects on average recorded at least twice as much plant cover as did adapted Daubenmire transects and modified‐Whittaker plots taken at the same location for all control and treatment units. Point‐intercept transects and adapted Daubenmire plots on average captured fewer rare and exotic species in the control and treatment units in comparison with the belt transects and modified‐Whittaker plots. Modified‐Whittaker plots captured the highest species richness in all units. Early successional understory response to restoration treatments was likely masked by the response of the herbaceous community to yearly climatic variation (dry vs. wet years). Species richness and abundance were higher in wet years than dry years for all control and treatment units. Our results illustrate that sampling techniques can greatly influence perceptions of understory plant trajectories and therefore the interpretation of whether restoration goals have been achieved. In addition, our results suggest that restoration monitoring needs to be conducted for a sufficient length of time so that restoration treatment responses can be detected.     

A meta‐analysis of species–abundance distributions

The species–abundance distribution (SAD) describes the abundances of all species within a community. Many different models have been proposed to describe observed SADs. Best known are the logseries, the lognormal, and a variety of niche division models. They are most often visualized using either species richness – log abundance class (Preston) plots or abundance – species rank order (Whittaker) plots. Because many of the models predict very similar shapes, model distinction and testing become problematic. However, the variety of models can be classified into three basic types: one that predicts a double S‐shape in Whittaker plots and a unimodal distribution in Preston plots (the lognormal type), a second that lacks the mode in Preston plots (the logseries type), and a third that predicts power functions in both plotting types (the power law type). Despite the interest of ecologists in SADs no formal meta‐analysis of models and plotting types has been undertaken so far. Here we use a compilation of 558 species–abundance distributions from 306 published papers to infer the frequency of the three SAD shapes in dependence of environmental variables and type of plotting. Our results highlight the importance of distinguishing between fully censused and incompletely sampled communities in the study of SADs. We show that completely censused terrestrial or freshwater animal communities tend to follow lognormal type SADs more often than logseries or power law types irrespective of species richness, spatial scale, and geographic position. However, marine communities tend to follow the logseries type, while plant communities tend to follow the power law. In incomplete sets the power law fitted best in Whittaker plots, and the logseries in Preston plots. Finally our study favors the use of Whittaker over Preston plots.     

Climate‐driven diversity change in annual grasslands: Drought plus deluge does not equal normal

Climate forecasts agree that increased variability and extremes will tend to reduce the availability of water in many terrestrial ecosystems. Increasingly severe droughts may be exacerbated both by warmer temperatures and by the relative unavailability of water that arrives in more sporadic and intense rainfall events. Using long‐term data and an experimental water manipulation, we examined the resilience of a heterogeneous annual grassland community to a prolonged series of dry winters that led to a decline in plant species richness (2000–2014), followed by a near‐record wet winter (2016–2017), a climatic sequence that broadly resembles the predicted future in its high variability. In our 80, 5‐m² observational plots, species richness did not recover in response to the wet winter, and the positive relationship of richness to annual winter rainfall thus showed a significant weakening trend over the 18‐year time period. In experiments on 100, 1‐m² plots, wintertime water supplementation increased and drought shelters decreased the seedling survival and final individual biomass of native annual forbs, the main functional group contributing to the observed long‐term decline in richness. Water supplementation also increased the total cover of native annual forbs, but only increased richness within nested subplots to which seeds were also added. We conclude that prolonged dry winters, by increasing seedling mortality and reducing growth of native forbs, may have diminished the seedbank and thus the recovery potential of diversity in this community. However, the wet winter and the watering treatment did cause recovery of the community mean values of a key functional trait (specific leaf area, an indicator of drought intolerance), suggesting that some aggregate community properties may be stabilized by functional redundancy among species.     

Multi-scale analysis of plant species richness in Serengeti grasslands

Aim To assess scale dependence between environmental factors and plant species richness. Additionally, we aimed to identify the scales at which niche relations and habitat heterogeneity, as hypothesized by A. Shmida & M.V. Wilson (1985) Journal of Biogeography, 12 , 1–20, operate in the savanna grasslands that were the focus of this study. Location Savanna grassland plant communities of Serengeti National Park, Tanzania. Methods Plant species richness was sampled in 102 modified Whittaker plots and tested for associations with two climate factors, mean annual rainfall (MAP) and potential evapotranspiration (PET), and two landscape variables, plot aspect (ASP) and topographic variation (TOPO), using multiple regressions. Scale dependence was assessed by conducting regressions after altering three aspects of spatial scale: grain, extent and focus. Grain was altered by analysing plant richness at 1, 10, 10² and 10³ m²; extent was investigated by restricting the maximum distance between samples to 75, 100, 125 and 150 km; and focus was manipulated by averaging samples spatially according to geographical land regions. Within the context of our data, we assumed that niche relations were represented by climate factors and habitat heterogeneity by landscape factors. Results Across all 102 plots, plant species richness between 1 and 10² m² had a negative relation to PET and a weak positive relation to MAP. Plant species richness at 10³ m² had a positive association with TOPO and weaker associations with climate factors. ASP stayed in the model between grains of 10 and 10³ m², but had a very weak positive association with richness. When the focus was changed to land regions, associations between plant species richness and explanatory variables strengthened, but were not qualitatively different. At spatial extents of 75 and 100 km, PET was the strongest correlate of plant species richness across all spatial grains. At spatial extents ≥ 125 km, PET explained the majority of the model variance at spatial grains ≤ 10² m², whereas TOPO explained equal amounts or more of the model variance at spatial grains of 10³ m². Main conclusions Both climate and topographic variation explained plant species richness in Serengeti grasslands, but specific patterns depended on grain, extent and, to a lesser degree, focus. Consistent with the ideas of Shmida & Wilson (1985) , determinants of plant species richness shifted from niche relations to habitat heterogeneity between spatial grains of 1 and 10³ m², although this occurred only at relatively large spatial extents (≥ 150 km). Finally, the signs, strength and shape of plant species richness relationships in Serengeti closely match those that describe macro‐scale patterns of woody plant species richness across the entire African continent.     

Species richness in deciduous forests: Effects of species pools and environmental variables

Abstract. The study was conducted in deciduous forests of two Swedish regions, Öland and Uppland. It had two objectives: to (1) test the species pool hypothesis by examining if differences in small‐scale species richness are related to differences in large‐scale species richness and the size of the regional species pool, and (2) to examine the relationship between species richness and productivity and its scale‐dependence. The first data set comprised 36 sites of moderate to high productivity. In each site, we recorded the presence of vascular plant species in nested plots ranging from 0.001 to 1000 m² and measured several environmental variables. Soil pH and Ellenberg site indicator scores for nitrogen were used as estimators of productivity. The second data set included 24 transects (each with 20 1‐m² plots) on Öland in sites with low to high productivity. Species number, soil pH and relative light intensity were determined in each plot. The forest sites on Öland were more species‐rich than the Uppland sites on all spatial scales, although environmental conditions were similar. Small‐scale and large‐scale species richness were positively correlated. The results present evidence in favour of the species pool hypothesis. In the nested‐plots data set, species number was negatively correlated with pH and nitrogen indicator scores, whereas a unimodal relationship between species number and pH was found for the transect data set. These results, as well as previously published data, support the hump‐shaped relationship between species richness and productivity in Swedish deciduous forests. Two explanations for the higher species richness of the sites with moderate productivity are given: first, these sites have a higher environmental heterogeneity and second, they have a larger ‘habitat‐specific’ species pool.     

Could we obtain better estimates of plot species richness from multiple‐observer plant censuses?

Question: Could we better estimate plot species richness by asking several botanists to survey the same plots and using non‐parametric estimators of richness? Location: Two French deciduous forests. Methods: Using replicated, independent censuses made by 11 professional botanists on the same eight 100‐m² forest plots, the relative performance of different richness estimators (Lincoln‐Petersen, Jackknife 1&2, Chao 1&2, Bootstrap, Chao Mth, Darroch) and the variation in their performance with the number of botanists involved (teams with two to eight botanists) were investigated. The sensitivity of these estimators to the presence of misidentifications in the data was also assessed. Results: When misidentifications are removed, Chao Mth estimators converged fastest to true richness, but none of the tested estimators correctly accounted for differences in exhaustiveness between the teams. Finally, all estimators were highly sensitive to misidentifications. Conclusions: Richness estimators are of little help in the presence of misidentifications and are ineffective at removing between‐team discrepancies, thus strongly limiting their usefulness in practice. Methods are presented to show how surveys can be designed to remove misidentifications and limit between‐team discrepancies. A sensible sampling design for 100‐m² plots in temperate forests would involve triplets of botanists and correcting data with the Chao N1. Pairs of botanists would already significantly improve the richness estimates, but such estimates would still be biased low. However, further research is needed to design new richness estimators that are more robust to observer effects.     

The influence of resprouting forest canopy species on richness in Southern Cape forests, South Africa

We investigated the relationship between species richness and numbers and types of individuals and species present in forests with different physiognomies in the southern Cape Province, South Africa. Data were collected from three different ‘plot’ types: 400 m², canopy‐scaled (plot length is directly proportional to canopy height) and per 100 individuals closest to a point. Plots were designed to control for the effect of scale on local richness. Canopy species richness was inversely proportional to the abundance of resprouting species. The strength of the relationship between the abundance of resprouters and canopy species richness increased progressively from the 400 m² plots to the canopy‐scaled plots and finally to the plots of 100 individuals. Resprouter abundance decreased, while canopy species richness increased, with increasing canopy height. Resprouters are able to retain their in situ position in the forests for longer periods of time than do reseeders. This reduces individual and species turnover, thus reducing species richness in resprouter‐dominated forests.     

Management intensity affects the relationship between non‐native and native species in subtropical wetlands

Question: Does management intensity affect the association between non‐native and native species and between non‐native species and soil nutrients in wetlands? Location: MacArthur Agro‐Ecology Research Center, Florida, USA. Methods: We evaluated native and non‐native plant richness and relative frequency in 15 1‐m² plots in 40 wetlands across two types of pastures, highly managed (fertilized, ditched, planted, heavily grazed by cattle) and semi‐natural (unfertilized, lightly seasonally grazed). Plant biomass was collected in five 0.25‐m² plots per wetland and sorted to species. Soil cores were collected to analyse soil total nitrogen (N) and phosphorus (P). An information‐theoretic approach was used to compare mixed effects models considering the association of non‐native richness, relative frequency, and biomass with native richness, relative frequency, biomass, C₃ grass relative frequency (a dominant native group), N, P and wetland‐type. Results: Non‐native richness was negatively correlated with native richness in semi‐natural wetlands, but there was no evidence of an association between these variables in highly managed wetlands. Non‐native richness increased with increasing soil N in semi‐natural wetlands, but not in the highly managed wetlands. Soil P was positively related to non‐native frequency in semi‐natural wetlands but negatively related in highly managed wetlands. Non‐native frequency and biomass were negatively related to relative frequency of C₃ grasses in both management types. Conclusions: Our results indicate that management intensity influences relationships between native and non‐native richness. Management intensity interacts with abiotic or biotic factors, such as soil nutrients and composition, in predicting where non‐native species will most likely need control.     

Effects of termite mounds on composition,functional types and traits of plant communities in Pendjari Biosphere Reserve (Benin,West Africa)

Understanding the role of termite mounds in biodiversity and ecosystem functioning is a priority for the management of tropical terrestrial protected areas dominated by savannahs. This study aimed to assess the effects of termite mounds on the diversity of plant functional types (PFTs) and herbaceous’ net aboveground primary productivity (NAPP) in plant communities (PCs) of the Pendjari Biosphere Reserve. PCs were identified through canonical correspondence analysis performed on 96 phytosociological ‘relevés’ realized in plots of 900 m². PFTs’ diversity was compared between savannahs and mounds’ plots using generalized linear models. In each plot, 7 m² subplots were harvested and NAPP was determined. Linear mixed models were performed to assess change in herbaceous NAPP regarding species richness, graminoids’ richness, specific leaf area and termite mounds. There is no specific plant community related to mounds. However, the occurrence of termite mounds induced an increase of woody and forbs diversity while the diversity of legumes and graminoids decreased. These diversity patterns led to decreasing of PCs’ NAPP. This study confirms that termite‐induced resource heterogeneity supports niche differentiation theory and increased savannah encroachment by woody species.     

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.