Constant tree species richness along an elevational gradient of Mt. Bokor,a table-shaped mountain in southwestern Cambodia

Some previous studies along an elevational gradient on a tropical mountain documented that plant species richness decreases with increasing elevation. However, most of studies did not attempt to standardize the amount of sampling effort. In this paper, we employed a standardized sampling effort to study tree species richness along an elevational gradient on Mt. Bokor, a table-shaped mountain in southwestern Cambodia, and examined relationships between tree species richness and environmental factors. We used two methods to record tree species richness: first, we recorded trees taller than 4 m in 20 uniform plots (5 × 100 m) placed at 266–1048-m elevation; and second, we collected specimens along an elevational gradient from 200 to 1048 m. For both datasets, we applied rarefaction and a Chao1 estimator to standardize the sampling efforts. A generalized linear model (GLM) was used to test the relationship of species richness with elevation. We recorded 308 tree species from 20 plots and 389 tree species from the general collections. Species richness observed in 20 plots had a weak but non-significant correlation with elevation. Species richness estimated by rarefaction or Chao1 from both data sets also showed no significant correlations with elevation. Unlike many previous studies, tree species richness was nearly constant along the elevational gradient of Mt. Bokor where temperature and precipitation are expected to vary. We suggest that the table-shaped landscape of Mt. Bokor, where elevational interval areas do not significantly change between 200 and 900 m, may be a determinant of this constant species richness.     

Testing species–stone area and species–bryophyte cover relationships in riverine macroinvertebrates at small scales

1. The species–area relationship is considered amongst the few genuine laws in ecology. Although positive species richness–stone area relationships have been found previously in stream systems, very few studies have simultaneously examined species–individuals, individuals–area, species–bryophyte biomass and individuals–bryophyte biomass relationships. We examined these relationships based on temporally replicated assessments of macroinvertebrates on stones at two river sites. 2. We found only one significant species–area relationship out of six relationship tested, and two significant individuals–area relationships. Even these significant relationships were weak, however. By contrast, we detected significant and rather strong relationships between species richness and the number of individuals at both river sites on all three sampling dates. We also found significant relationships of both species richness and the number of individuals with bryophyte biomass at both river sites on all sampling occasions. One of the river sites was disturbed by a bulldozer, and the species–bryophyte biomass relationships were somewhat stronger after the disturbance event. 3. Our findings are quite surprising, given that there were very weak species–area relationships on stream stones. By contrast, our results suggest a pivotal role for bryophyte biomass in determining the species richness and the number of individuals of stream macroinvertebrates at this small scale. The most probably origin of these relationships begins with bryophyte cover, which determines the number of individuals, and subsequently passively affects species richness. Thus, there is not necessarily a direct mechanism that determines the variability of species richness on stream stones. 4. Experimental studies are needed to disentangle the various mechanisms (e.g. passive sampling, provision of more food, more niche space, flood disturbance refugia) by which bryophyte biomass affects stream macroinvertebrates.     

Competitive exclusion and the No‐Interaction model operate simultaneously in microcosm plant communities

Question: Is the relation between productivity and species richness due to an increase in plant size and hence a reduced plant density? Location: Glasshouse experiment. Methods: Productivity was manipulated with fertilizer and irrigation in a microcosm experiment. The ‘sampling effect’ was removed using rarefaction to a common density of individual plants per pot. Results: Fertility increased community biomass towards an asymptotic maximum, and reduced the light passing through plant canopies towards an asymptotic minimum. As biomass increased, so did species richness. However, this did not seem to be a direct effect of productivity on species richness, but rather one mediated by plant density, since: (1) the richness/ density relation was stronger than the richness/biomass one; (2) adding biomass to the richness/density regression did not increase its predictivity; (3) the richness/biomass relation was removed by rarefaction to 200 individuals per pot. It is therefore concluded that the richness/biomass relation observed was due to the sampling effect. Rarefaction to a small number of plants gave a quite different trend: lower richness at high biomass. This seems to be due to an increased number of subordinate species at high community biomass, and a more uneven distribution of abundance. Conclusion: The Competitive Exclusion and No‐Interaction hypotheses have been seen as alternatives. We suggest that they can operate simultaneously.     

Models and estimators linking individual-based and sample-based rarefaction,extrapolation and comparison of assemblages

Aims In ecology and conservation biology, the number of species counted in a biodiversity study is a key metric but is usually a biased underestimate of total species richness because many rare species are not detected. Moreover, comparing species richness among sites or samples is a statistical challenge because the observed number of species is sensitive to the number of individuals counted or the area sampled. For individual-based data, we treat a single, empirical sample of species abundances from an investigator-defined species assemblage or community as a reference point for two estimation objectives under two sampling models: estimating the expected number of species (and its unconditional variance) in a random sample of (i) a smaller number of individuals (multinomial model) or a smaller area sampled (Poisson model) and (ii) a larger number of individuals or a larger area sampled. For sample-based incidence (presence–absence) data, under a Bernoulli product model, we treat a single set of species incidence frequencies as the reference point to estimate richness for smaller and larger numbers of sampling units.Methods The first objective is a problem in interpolation that we address with classical rarefaction (multinomial model) and Coleman rarefaction (Poisson model) for individual-based data and with sample-based rarefaction (Bernoulli product model) for incidence frequencies. The second is a problem in extrapolation that we address with sampling-theoretic predictors for the number of species in a larger sample (multinomial model), a larger area (Poisson model) or a larger number of sampling units (Bernoulli product model), based on an estimate of asymptotic species richness. Although published methods exist for many of these objectives, we bring them together here with some new estimators under a unified statistical and notational framework. This novel integration of mathematically distinct approaches allowed us to link interpolated (rarefaction) curves and extrapolated curves to plot a unified species accumulation curve for empirical examples. We provide new, unconditional variance estimators for classical, individual-based rarefaction and for Coleman rarefaction, long missing from the toolkit of biodiversity measurement. We illustrate these methods with datasets for tropical beetles, tropical trees and tropical ants.Important findings Surprisingly, for all datasets we examined, the interpolation (rarefaction) curve and the extrapolation curve meet smoothly at the reference sample, yielding a single curve. Moreover, curves representing 95% confidence intervals for interpolated and extrapolated richness estimates also meet smoothly, allowing rigorous statistical comparison of samples not only for rarefaction but also for extrapolated richness values. The confidence intervals widen as the extrapolation moves further beyond the reference sample, but the method gives reasonable results for extrapolations up to about double or triple the original abundance or area of the reference sample. We found that the multinomial and Poisson models produced indistinguishable results, in units of estimated species, for all estimators and datasets. For sample-based abundance data, which allows the comparison of all three models, the Bernoulli product model generally yields lower richness estimates for rarefied data than either the multinomial or the Poisson models because of the ubiquity of non-random spatial distributions in nature.     

The species richness–biomass relationship in herbaceous plant communities: what difference does the incorporation of root biomass data make?

So far, in all studies on the much-discussed hump-backed relationship between plant community productivity and species richness, productivity has been assessed through plant shoot biomass, i.e. it has been ignored that frequently most of the biomass is produced below ground. We revisited the 27 grassland and forest field-layer communities, studied earlier by Zobel and Liira, to sample root biomass, plant total biomass and root/shoot allocation, and learn how the incorporation of below-ground biomass data would affect the shape of the hump-backed relationship. In order to avoid scaling artefacts we estimated richness as the average count of species per 500 plant ramets (absolute richness). We also included relative richness measures. Relative richness was defined as richness per 500 ramets/size of the actual species pool (the set of species present in the community), relative pool size was defined as size of the actual species pool/size of the regional species pool (the set of species available in the region and capable of growing in the given community).
The biomass-absolute richness relationship was humped, irrespective of the biomass measure used, the hump being most obvious when plant total biomass was used as the independent variable. Evidently, the unimodal richness–productivity curve is not a sampling artefact, as suspected by Oksanen. However, relative richness was not related to community biomass (above-ground, below-ground or total). The hump-backed curve is shaped by the sizes of actual species pools in communities, implying that processes which are responsible for small-scale diversity pattern mainly operate on the community level.
Neither absolute nor relative richness were significantly related to root/shoot allocation. The presumably stronger (asymmetric) shoot competition at greater allocation to shoots appears not to suppress small-scale richness. However, there is a significant relationship between relative pool size and root/shoot allocation. Relatively more species from regional species pools are able to enter and persist in communities with more biomass allocated into roots.     

The effects of protection from fishing on species richness: distinguishing between alternative explanations

Marine reserves that prohibit fishing often result in greater densities of individuals and more species than adjacent fished areas. However, simple conclusions about their effects on species richness are confounded, because more species are expected to occur wherever there are more individuals. Here, there is an important distinction between the number of species per sampling unit (species density), and species richness measured as the number of species per given number of individuals. When conservation of species richness is an important goal, analyses need to discriminate between the alternative explanations for differences in the number of species. We used rarefaction to test whether species richness was higher in two ‘no-take’ marine reserves after controlling for differences in the density of individuals. We surveyed each reserve in three different years. There was a higher density of individuals and species in each reserve than in adjacent fished areas. However, rarefaction analyses indicated that effects on species richness were weak after controlling for the number of individuals: slightly higher species richness was recorded inside each reserve in one of three surveys, but the difference was small, and was apparent only when the maximum number of individuals was approached. Our results therefore indicate that patterns in species density were not reflected by patterns in species richness—the application of rarefaction methods is needed to determine the responses of species richness to protection elsewhere. The distinction between species density and species richness will not be important in all situations, but when it is important, inferences about species richness cannot be reliably deduced from measurements of species density.     

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.     

Species pool size and rainfall account for the relationship between biodiversity and biomass production in natural forests of China

The strength of biodiversity–biomass production relationships increases with increasing environmental stress and time. However, we know little about the effects of abiotic (e.g., climate) and biotic (e.g., species pool and community composition) factors on this trend. Whether variation in biomass production is best explained by phylogenetic diversity metrics or traditional measures of species richness also remains elusive. We compiled estimates of community composition and biomass production for tree species in 111 permanent quadrats spanning three natural forests (tropical, subtropical, and temperate) in China. Based on ~10 years of data, we compared temperature, rainfall, species pool size, and community composition in each forest each year. We estimated species richness and phylogenetic diversity in each quadrat each year; the latter metric was based on the sum of branch lengths of a phylogeny that connects species in each quadrat each year. Using generalized linear mixed‐effect models, we found that top‐ranked models included the interaction between forest and biodiversity and the interaction between forest and year for both biodiversity metrics. Variation in biomass production was best explained by phylogenetic diversity; biomass production generally increased with phylogenetic diversity, and the relationship was stronger in subtropical and temperate forests. Increasing species pool size, temperature, and rainfall and decreasing inter‐quadrat dissimilarity range shifted the relationship between biomass production and phylogenetic diversity from positive to neutral. When considered alone, species pool size had the strongest influence on biomass production, while species pool size, rainfall, and their interaction with phylogenetic diversity constituted the top‐ranked model. Our study highlights the importance of species pool size and rainfall on the relationship between phylogenetic diversity and biomass production in natural forest ecosystems.     

Is breaking up hard to do? Breakage, growth, and survival in the parasitic clonal plant Cuscuta corymbosa (Convolvulaceae)

In a tripartite exploration of the effects of traumatic breakage in the parasitic clonal plant Cuscuta corymbosa, experimental breakage had significant negative effects upon stem extension of ramets distal to a break and the level of that effect depended upon the number of ramets in the severed fragment. Regular monitoring of clonal individuals transplanted onto naturally occurring, native host species revealed that breakage in individuals growing under natural conditions ranged from 2 to 66% of all interramet connections and significantly affected distance between ramets and stolon generation in ramets associated with a break. Simulations revealed that these responses to breakage would significantly change the "shape" of a clonal individual in a manner capable of affecting probabilities of encounter with patchily distributed resources. Nonetheless, the observed levels of breakage had no discernible effect on biomass accumulation, although individuals that survived into the dry season and thus gained the potential to reproduce in additional seasons had a significantly lower rate of breakage than those that did not. We suggest that the lack of a relationship between breakage and within-season biomass accumulation is an indication that parasite response to breakage is gauged to expected levels of breakage for the habitat. We note that the capacity of ramets to survive disconnection significantly decreased the loss of tissue that would occur if subunits did not possess the potential to function independently.     

取样尺度对亚高寒草甸物种多样性与生产力关系的影响

植物物种多样性与生产力之间的关系是群落生态学的一个热点问题, 目前仍存在着很多争议。为探究自然群落中二者之间的关系, 对青藏高原亚高寒草甸3个样地的自然植物群落分别进行了不同取样面积的抽样调查。结果显示, 取样样地和取样尺度均对物种丰富度有显著影响, 取样样地而非取样尺度对群落地上生物量有显著性影响。在某一时刻对某一样地进行取样, 其单位面积生产力并不因取样面积的增加而提高, 而是保持恒定的, 尽管物种数随取样面积的增加而有明显增多。物种多样性与生产力之间的回归关系因样地与取样尺度不同而不同, 有U型、单峰型、正线性相关和无相关性, 其中无相关性出现的最多。据此推测, 亚高寒草甸群落物种多样性与生产力之间不存在某种确定性关系, 或者说, 亚高寒草甸物种多样性和生产力之间不存在必然的因果联系。     

Evaluating detectability of freshwater fish assemblages in tropical streams: Is hand-seining sufficient?

Unprecedented threats to natural ecosystems mean that accurate quantification of biodiversity is a priority, particularly in the tropics which are underrepresented in monitoring schemes. Data from a freshwater fish assemblage in Trinidad were used to evaluate the effectiveness of hand-seining as a survey method in tropical streams. We uncovered large differences in species detectability when hand-seining was used alone, in comparison with when hand-seining and electrofishing were used together. The addition of electrofishing increased the number of individuals caught threefold, and increased the biomass fivefold. Some species were never detected using hand-seining, resulting in significant underestimates of species richness; rarefaction curves suggest that even when hand-seining effort increases, species richness is still underestimated. Diversity indices (Shannon and Simpson index) reveal that diversity was also significantly lower for hand-seined samples. Furthermore, the results of multivariate analyses investigating assemblage structure also differed significantly depending on whether they were based on hand-seined data alone, or a combination of hand-seining and electrofishing. Despite the extra equipment and maintenance required, these findings underline the value of including electrofishing when sampling tropical freshwater streams.     

百里香无性系的克隆生长特征

植物克隆生长及其与生态适应性的关系是当今植物种群生态学研究的热点和前沿课题,但目前小半灌木克隆生长的研究开展不多。百里香(Thymus serpyllum var. asiaticus)是一种具有地面匍匐茎的草本状小半灌木,可在土壤侵蚀剧烈、基岩大面积裸露的砒砂岩区形成百里香单优群落,在维持生态系统稳定方面具有重要的生态学作用。皇甫川流域是砒砂岩大面积分布的典型区域,在这一地区对百里香无性系的克隆生长进行研究,不仅具有重要的学术价值,而且在生态环境建设方面也具有一定的现实意义。在皇甫川流域选择含三级分株的百里香无性系,对其各级分株的总生物量、各构件生物量及数量、各构件生物量占总生物量的百分比及其月变化进行了研究。结果表明: 1)母株与子代相比,在总生物量、构件生物量及数量上占有绝对优势,而且具有体型大、结构复杂的特点; 2)对生物量分配格局的研究显示,母株根的生物量在总生物量中所占的比例最大,其叶所占的比例较低。子代叶的生物量在总生物量中所占的比例最大,其根所占的比例较低;3)不同级别分株在生物量分配上的差异,揭示了相互连接的分株在功能上的差别,母株可能更侧重于养分和水分的吸收,子株则更侧重于光合生产;4)构件枝、茎、花生物量分配比月变化显示,子1代各构件的生长规律与母株的基本一致,子2代与母株和子1代的相比差异较大,分析认为这可能是分株间不同程度的生理整合作用造成的结果。     

Community assessment techniques and the implications for rarefaction and extrapolation with Hill numbers

Diversity estimates play a key role in ecological assessments. Species richness and abundance are commonly used to generate complex diversity indices that are dependent on the quality of these estimates. As such, there is a long‐standing interest in the development of monitoring techniques, their ability to adequately assess species diversity, and the implications for generated indices. To determine the ability of substratum community assessment methods to capture species diversity, we evaluated four methods: photo quadrat, point intercept, random subsampling, and full quadrat assessments. Species density, abundance, richness, Shannon diversity, and Simpson diversity were then calculated for each method. We then conducted a method validation at a subset of locations to serve as an indication for how well each method captured the totality of the diversity present. Density, richness, Shannon diversity, and Simpson diversity estimates varied between methods, despite assessments occurring at the same locations, with photo quadrats detecting the lowest estimates and full quadrat assessments the highest. Abundance estimates were consistent among methods. Sample‐based rarefaction and extrapolation curves indicated that differences between Hill numbers (richness, Shannon diversity, and Simpson diversity) were significant in the majority of cases, and coverage‐based rarefaction and extrapolation curves confirmed that these dissimilarities were due to differences between the methods, not the sample completeness. Method validation highlighted the inability of the tested methods to capture the totality of the diversity present, while further supporting the notion of extrapolating abundances. Our results highlight the need for consistency across research methods, the advantages of utilizing multiple diversity indices, and potential concerns and considerations when comparing data from multiple sources.     

Coarse‐scale plant species richness in relation to environmental heterogeneity

Abstract. We test to what extent mean environmental conditions and environmental heterogeneity are related to species richness in a regular geographical grid system (UTM) of 10 km × 10 km in the NE Iberian Peninsula (i.e. Catalonia, ca. 31 900 km²). Species richness of each UTM quadrat was estimated by compiling a large database (more than a million records) from bibliographic references and atlases. Mean environmental conditions of each quadrat were derived from climatic maps. Environmental heterogeneity was estimated from the diversity of geological substrates and climatic classes in each quadrat. The increase in effective (real) area due to topographic complexity was also considered (derived from the digital elevation model). The statistical analysis was performed by a weighted analysis of deviance assuming a negative binomial error distribution. The results suggest that species richness in the study area is a function of both within‐quadrat heterogeneity (specifically, effective area, heterogeneity of geological substrates, heterogeneity of January temperature) and mean environmental conditions (mean annual temperature, Thornthwaite moisture index and aspect). All these parameters showed a positive relationship with species richness. Quadrat heterogeneity accounted for ca. 2/3 of the explained deviance, suggesting the importance of environmental heterogeneity when using a geographical grid system. The study fits well with earlier results on the importance of climatic parameters on plant species richness and provides one of the few rigorous, quantitative, coarse‐scale studies testing environmental heterogeneity in plant species richness.     

A comparison of Passalidae (Coleoptera, Lamellicornia) diversity and community structure between primary and secondary tropical forest in Los Tuxtlas, Veracruz, Mexico

Comparison of the diversity and community structure of Coleoptera (Passalidae) collected in Los Tuxtlas, Veracruz, Mexico, in primary and secondary tropical forest has been carried out. The saproxylophagous beetles studied can be differentiated according to their presence in three distinct microhabitats of rotting logs: underbark, sapwood—heartwood and microhabitat generalists. Over the 2-year study period, 12 passalid species were recorded (six Passalini and six Proculini) represented by a total of 2971 individuals, collected from 234 rotting logs. The rarefaction method, the lognormal species—abundance relationship, and the nonparametric jackknife method were used to compare species richness between the habitats. The data were also fitted to log series, truncated lognormal, geometric, and broken-stick species abundance models to detect changes in community structure. The community composition of Passalidae in Los Tuxtlas did not differ ostensibly between the primary and secondary forests. Neither the mean number of individuals nor the biomass per log differed significantly. Furthermore, there were no significant differences between the two habitats in terms of the number of underbark, sapwood/heartwood, and microhabitat generalist species. Different richness estimators indicated that the primary forest community is only slightly richer. The slight decrease in richness of the secondary forest is related to a decrease in dominance by certain species, as well as to a more balanced abundance distribution, which is adequately described by the broken-stick model. Complementary explanations for this pattern may be: (1) that logging reduces the abundance of dominant species, thus preventing competitive exclusion in the secondary forest; and (2) that passalid diversity is not regulated by the diversity of tree species.     

Species composition of Bromeliaceae and their distribution at the Massambaba restinga in Arraial do Cabo, Rio de Janeiro, Brazil

We studied some ecological parameters such as richness, abundance, density, biomass and variation in species composition in four vegetation zones and in a zone with anthropic disturbance in the Massambaba Restinga in Arraial do Cabo, Rio de Janeiro State. We sampled 100 plots of 100 m(2) (10 x 10 m) recording the bromeliad species and their abundance. We found a total of seven bromeliad species, with Vriesea neoglutinosa (5647 ramets) and Tillandsia stricta (1277 ramets) being the most abundant. The vegetation zone called Clusia shrubs had the highest richness (S = 5) and density (6360 ramets.ha(-1)) of bromeliads. The differences found in abundance and variation in species composition among vegetation zones seems to be related to the vegetation structure of each zone.     

Effects of trait plasticity on aboveground biomass production depend on species identity in experimental grasslands

Experimental studies with grassland species found a positive relationship between species richness and community biomass production, however the response of individual species was highly variable. The mechanisms behind these patterns are poorly understood. Here we studied aboveground biomass production and plasticity of growth characteristics of four legumes with similar morphology (Lotus corniculatus, Medicago lupulina, Onobrychis viciifolia, Trifolium hybridum) in experimental grasslands varying in species richness (1, 2, 4, 8, 16 and 60) and composition. We identified O. viciifolia and T. hybridum as species that reached higher biomass production in mixtures than expected from monoculture yields, while L. corniculatus and M. lupulina mostly had lower yields than expected. Variation of morphological traits across the species-richness gradient was lowest in the highly competitive O. viciifolia, but increased in the smaller species. The tall-growing O. viciifolia achieved higher biomass production by both, a higher number of plant individuals and an increase in mean mass per individual. Mean shoot number per individual remained constant, but individuals produced heavier shoots. The medium-sized T. hybridum also increased the number of plant individuals, but mean mass per individual did not respond to community species richness. The average mass per shoot was increased in mixtures, but the species developed less shoots per individual. Shoot length and stem weight ratio of T. hybridum increased with community species richness. Morphological changes in the less successful L. corniculatus and M. lupulina with a smaller growth stature were similarly directed as those of T. hybridum. The observed morphological changes are known as typical shade-avoidance mechanisms in dense vegetation. Our study shows that stress responses to changes in resource availability may be a mechanism to enforce higher aboveground biomass production of individual species in mixtures, but it depends on species identity whether trait plasticity is large enough to exceed stress-induced growth limitations.     

Roles of succession,light, nutrients and disturbance on population vigor and maintenance of the rare plant Solidago shortii (Asteraceae)

Results of field and glasshouse experiments on Solidago shortii, and our observations on this species over many years, were used to construct a conceptual model of the roles of succession, light, soil nutrients and disturbance on population vigor and maintenance of this federal-endangered species. As cover of woody vegetation increased at a population site between 1986 and 1992, number of flowering ramets of S. shortii significantly decreased but number of vegetative ramets remained nearly constant. Adult plants transplanted into a redcedar thicket and those shaded in a glasshouse produced many fewer flowering ramets and capitula per flowering ramet and less biomass and had higher mortality than those in the open. Seedlings/juveniles shaded in a glasshouse had significantly less dry biomass; lower RGR, NAR, leaf area and root/shoot ratio and higher LAR, SLA and LWR than nonshaded ones. In a field site and glasshouse, fertilized plants (NPK) consistently had more flowering ramets and capitula per flowering ramet than nonfertilized ones. Hierarchy of dry weight of plants grown in a glasshouse in soils derived from five types of bedrock was phosphatic limestone > calcareous shale > sandstone > black shale = dolomite. Flowering and biomass production in the field-fertilizer and soil-type experiments were associated closely with levels of P. Number of flowering ramets significantly increased in plants transferred from shaded to nonshaded glasshouse conditions, but no such increase occurred after opening the canopy above plants in a thicket. Both high light and high nutrient levels apparently are necessary to maintain high vigor of S. shortii. In areas subject to invasion by woody plants, periodic high intensity disturbance may be required to prevent population extirpation.     

Effects of additional illumination and fertilization on seasonal changes in fine-scale grassland community structure

Abstract. Fine-scale structure of a species-rich grassland was examined for seasonal changes caused by manipulated changes in the availability of above and below-ground resources (additional illumination with the help of mirrors and fertilization) in a field experiment. If changes induced by fertilization — which are expected to lead to a reduction in small-scale diversity —are due to intensified light competition, they should be compensated for by additional light input. Permanent plots of 40 cm × 40 cm were sampled by the point quadrat method at three angles (60°, 90° and 120° from the horizontal North-South direction), using a laser beam to position the quadrats, in early July and early September. The applied treatments did not cause apparent changes in plant leaf orientation. The degree of spatial aggregation of biomass increased seasonally in fertilized, non-illuminated plots: greater productivity at a constant light supply led to a faster growth rate of potentially dominant species, as compared to the subordinate ones. Additional illumination mitigated this effect of fertilization, indicating that the observed changes in biomass aggregation were due to increased light competition. There was a considerable seasonal decrease of variance ratio (ratio of observed variance of richness at a point and variance expected at random) in fertilized only and in illuminated only plots. In fertilized plots this was due to the positive relationship between biomass aggregation and expected variance of richness. Biomass constancy occurs to be inversely related to deficit in variance of richness. In illuminated plots, in contrast, only the observed variance of richness decreased seasonally, indicating a more uniform use of space by different species. Evidently, a deficit in variance of richness can be caused by drastically different processes, showing that the variance ratio statistic may not have a significant explanatory value in fine-scale community studies.     

Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness

Species richness is a fundamental measurement of community and regional diversity, and it underlies many ecological models and conservation strategies. In spite of its importance, ecologists have not always appreciated the effects of abundance and sampling effort on richness measures and comparisons. We survey a series of common pitfalls in quantifying and comparing taxon richness. These pitfalls can be largely avoided by using accumulation and rarefaction curves, which may be based on either individuals or samples. These taxon sampling curves contain the basic information for valid richness comparisons, including category–subcategory ratios (species-to-genus and species-to-individual ratios). Rarefaction methods – both sample-based and individual-based – allow for meaningful standardization and comparison of datasets. Standardizing data sets by area or sampling effort may produce very different results compared to standardizing by number of individuals collected, and it is not always clear which measure of diversity is more appropriate. Asymptotic richness estimators provide lower-bound estimates for taxon-rich groups such as tropical arthropods, in which observed richness rarely reaches an asymptote, despite intensive sampling. Recent examples of diversity studies of tropical trees, stream invertebrates, and herbaceous plants emphasize the importance of carefully quantifying species richness using taxon sampling curves.