Dynamics of core and occasional species in the marine plankton: tintinnid ciliates in the north-west Mediterranean Sea

Aim To assess short‐term variability in the community composition and community structure of tintinnid ciliates, herbivores of the microzooplankton. Location North‐west Mediterranean Sea. Methods We sampled on 18 dates over a 4‐week period in 2004 at an open‐water site. Species were classified as ‘core species’, found on every date, or ‘occasional species’, absent on one or more dates. Species abundance distributions of the entire community, and separately the core and occasional species, were compared with geometric, log‐series and log‐normal distributions. Core and occasional species were compared in terms of the shell or lorica oral diameter (LOD), analogous to gape size. Results We found 11 core and 49 occasional species. Diversity metrics were stable compared with shifts in abundances. Core species accounted for the majority of individuals in all samples. On each date, 9–22 occasional species, representing 10–15% of the population, were found. Species richness of the occasionals was positively related to population size. The identities of the occasional species found were unrelated to the time between sampling. The species abundance distribution of the occasional population was best fit by a log‐series distribution, while that of the core species was best fit by a log‐normal distribution. The species abundance distribution of the entire community was best fit by a log‐series distribution. Most of the occasional species had LODs distinct from that of a core species and occupied size classes left empty by the core population. However, the most abundant and frequent of the occasional species had a LOD similar to that of a core species. Main conclusions Among tintinnids, which are planktonic protists, occasional species have a species abundance distribution pattern distinct from that of core species. Occasional species appeared to be composed of two groups, one of relatively abundant species and similar to core species, and a second group of ephemeral species with morphologies distinct from core species. The existence of two categories of occasional or rare species may be common: (1) those similar to, and thus perhaps able to replace, dominant species in the absence of a change in the environment; and (2) those distinct from dominant species and requiring different conditions to prosper.     

Community Structure of Tintinnid Ciliates of the Microzooplankton in the South West Pacific Ocean: Comparison of a High Primary Productivity with a Typical Oligotrophic Site

Transient ‘hot spots’ of phytoplankton productivity occur in the generally oligotrophic Southern Pacific Ocean and we hypothesized that the population structure of tintinnid ciliates, planktonic grazers, would differ from that of a typical oligotrophic sites. Samples were collected over a 1‐wk period at each of two sites between Fiji and Tahiti: one of elevated chlorophyll a concentrations and primary productivity with an abundance of N‐fixing cyanobacteria Trichodesmium, and a distant oligotrophic site. Tintinnid abundance differed between the sites by a factor of 2. A single species (Favella sp.), absent from the oligotrophic site, highly dominated the ‘hot spot’ site. However, total species richness was identical (71 spp.) as well as short‐term temporal variability (2–4 d). At both sites, species abundance distributions most closely fit a log‐series or log‐normal distribution and the abundance distributions of ecological types, forms of distinct lorica oral diameter, were the typical geometric. Morphological diversity was only slightly lower at the high productivity site. We found that communities of these plankton grazers in ‘hot spots’ of phytoplankton productivity in oligotrophic systems, although harboring different species, differ little from surrounding oligotrophic areas in community structure.     

Relating species abundance distributions to species-area curves in two Mediterranean-type shrublands

Abstract. Based on both theoretical and empirical studies there is evidence that different species abundance distributions underlie different species‐area relationships. Here I show that Australian and Californian shrubland communities (at the scale from 1 to 1000 m²) exhibit different species‐area relationships and different species abundance patterns. The species‐area relationship in Australian heathlands best fits an exponential model and species abundance (based on both density and cover) follows a narrow log normal distribution. In contrast, the species‐area relationship in Californian shrublands is best fit with the power model and, although species abundance appears to fit a log normal distribution, the distribution is much broader than in Australian heathlands. I hypothesize that the primary driver of these differences is the abundance of small‐stature annual species in California and the lack of annuals in Australian heathlands. Species‐area is best fit by an exponential model in Australian heathlands because the bulk of the species are common and thus the species‐area curves initially rise rapidly between 1 and 100 m². Annuals in Californian shrublands generate very broad species abundance distributions with many uncommon or rare species. The power function is a better model in these communities because richness increases slowly from 1 to 100 m² but more rapidly between 100 and 1000 m² due to the abundance of rare or uncommon species that are more likely to be encountered at coarser spatial scales. The implications of this study are that both the exponential and power function models are legitimate representations of species‐area relationships in different plant communities. Also, structural differences in community organization, arising from different species abundance distributions, may lead to different species‐area curves, and this may be tied to patterns of life form distribution.     

南亚热带森林群落种-多度的对数正态分布模型研究

通过对地处南亚热带的广东省黑石顶自然保护区森林群落的定点研究结果的分析表明：当用１,２,３,…分组每种个体数ｒ时,５个不同类型的群落样地的种一多度分布的直方图都呈明显的倒Ｊ－形;经Ｐｒｅｓｔｏｎ“倍程（ｏｃｔａｖｅｓ）”法分组ｒ后,其种－多度都服从对数正态分布。由种－多度模型可以推出另一新的模型一个体一多度分布模型,即Ｉ（Ｒ）＝２R0ＳＯＥＸＰ［［１ｎ２）２／４ａ２〕ＥＸＰ｛－ａ２（Ｒ－（ＲＯ＋１ｎ２／２ａ２）］２｝,它也符合对数正态分布。另外,还运用积分方法推导出估计总体（整个群落）中总种数Ｓ*和总个体数Ｉ*理论值的公式,用此公式估计的结果较为合理。     

Ciliate diversity and distribution across an environmental and depth gradient in Long Island Sound,USA

The nature and extent of microbial biodiversity remain controversial with persistent debates over patterns of distributions (i.e. cosmopolitanism versus endemism) and the processes that structure these patterns (neutrality versus selection). We used culture‐independent approaches to address these issues focusing on two groups of ciliates, the Oligotrichia (Spirotrichea) and Choreotrichia (Spirotrichea) across an environmental gradient. We assessed SSU rDNA diversity in ciliate communities at six stations in Long Island Sound spanning the frontal region that separates the fresher Connecticut River outflow plume from the open Sound. As in previous studies, we find one abundant cosmopolitan species (Strombidium biarmatum), a few moderately abundant sequences, and a long list of rare sequences. Furthermore, neither ciliate diversity nor species composition showed any clear relationship to measured environmental parameters (temperature, salinity, accessory pigment composition and chorophyll). Overall, we observed that diversity decreased moving from nearshore to offshore. We also conducted analyses to detect clustering among the sampled communities using the software Unifrac. This approach revealed three significant clusters grouping samples from nearshore, surface and deep/well mixed stations. We find no strong fit of our communities to log series, geometric or log normal distributions, though one of the 3 clusters is most consistent with a log series distribution. However, when we remove the abundant cosmopolitan species S. biarmatum, all three clusters fit to a log series distribution. These analyses suggest that, with the exception of one cosmopolitan species, the oligotrich and choreotrich communities at these stations may be distributed in a neutral manner.     

中国暖温带若干灌丛群落多样性问题的研究

 暖温带灌丛的生物多样性的研究表明：1．灌丛群落的均一度指数同优势度指数的相关性较丰富度指数同变化度指数的相关性为高;在众多的多样性指数中,D4(Shannon信息度指数)和D6(Gini指数)因最大限度地包含了其它各类多样性指数的信息而被采用。2．灌丛群落的物种多样性指数值随着演替的进行,没有表现出降低的趋势。3．生物多样性较高的荆条灌丛和辽东栎萌丛的物种多度分布是对数分布,而生物多样性较低的鬼见愁灌丛和毛榛灌丛的物种多度分布是几何分布,处于中间的三桠绣线菊灌丛的物种多度分布是Broken-Stick分布;没有正态对数分布类型。     

塔里木荒漠河岸林不同生境群落物种多度分布格局

为解释塔里木荒漠河岸林群落构建和物种多度分布格局形成的机理, 本文以塔里木荒漠河岸林2个不同生境(沙地、河漫滩) 4 ha固定监测样地为研究对象, 基于两样地物种调查数据, 采用统计模型(对数级数模型、对数正态模型、泊松对数正态分布模型、Weibull分布模型)、生态位模型(生态位优先占领模型、断棍模型)和中性理论模型(复合群落零和多项式模型、Volkov模型)拟合荒漠河岸林群落物种多度分布, 并用K-S检验与赤池信息准则(AIC)筛选最优拟合模型。结果表明: (1)随生境恶化(土壤水分降低), 植物物种多度分布曲线变化减小, 群落物种多样性、多度和群落盖度降低, 常见种数减少。(2)选用的3类模型均可拟合荒漠河岸林不同生境群落物种多度分布格局, 统计模型和中性理论模型拟合效果均优于生态位模型。复合群落零和多项式模型对远离河岸的干旱沙地生境拟合效果最好; 对数正态模型和泊松对数正态模型对洪水漫溢的河漫滩生境拟合效果最优; 中性理论模型与统计模型无显著差异。初步推断中性过程在荒漠河岸林群落构建中发挥着主导作用, 但模型拟合结果只能作为推断群落构建过程的必要非充分条件, 不能排除生态位过程的潜在作用。     

Limits to the estimation of species richness: The use of relative abundance distributions

The present study demonstrates the possibility of estimating species numbers of animal or plant communities from samples using relative abundance distributions. We use log‐abundance–species‐rank order plots and derive two new estimators that are based on log‐series and lognormal distributions. At small to moderate sample sizes these estimators appear to be more precise than previous parametric and nonparametric estimators. We test our estimators using samples from 171 published medium‐sized to large animal and plant communities taken from the literature. By this we show that our new estimators define also limits of precision.     

Edaphic Distribution of Some Species of the Fern Genus Adiantum in Western Amazonia1

The distribution patterns of Adiantum species were inventoried at twelve noninundated rain forest (tierra firme) sites in Amazonian Peru and Ecuador. Six species belonging to the genus were found, and four of these were abundant at some of the sites. At six of the sites none of the species was particularly abundant. Both the occurrence and the abundance of the species varied among sites with regard to soil texture and concentration of exchangeable bases and within sites with regard to topography. Adiantum tomentosum) Klotzsch was restricted to loamy soils with intermediate to low content of extractable bases and was locally most abundant in the well drained parts of slopes and ridge tops. Adiantum terminatum Kunze ex Miq. and A. humile Kunze occurred at sites with intermediate to high content of extractable bases, but their relative abundances differed among and within sites. Adiantum pulverulentum L. was restricted to soils with relatively high base concentration. Ecological differences among the species obviously explained many of the differences in their distribution patterns at the local and regional scales, and may be important for the understanding of such patterns at biogeographical scales as well.     

Use of the abundance spectrum and relative-abundance distributions to analyze assemblage change in massively altered landscapes

Fragmentation of natural landscapes is a pervasive process in the world. Common models predict coherent change in assemblages, with less numerous species becoming locally extinct first, then species of intermediate abundance, and so forth. Relative-abundance distributions should change systematically in landscapes characterized by greater change. Such a predictable sequence of change is not evident in the avifaunas of landscapes of central Victoria, Australia, where relative-abundance patterns in more affected landscapes bear little resemblance to reference distributions. I provide two sets of analyses of relative-abundance distributions: (1) analyses that do not depend on the identity of individual species and (2) abundance spectra, which use ordered lists of species ranked by species' commonness in reference systems. While abundance spectra change dramatically in smaller remnants, relative-abundance distributions change little, suggesting that the "reorganization" of abundances occurs over ecological time frames. The dispersal-limited multinomial is a flexible distribution that may fit many data sets yet be unrelated to assumptions (species neutrality) and processes (fixed total numbers of individuals) of the unified neutral theory. A more complete understanding of human impacts at landscape scales must include capacities to predict those species that will be advantaged by change, as well as those that will be disadvantaged.     

Ant Diversity and Abundance along an Elevational Gradient in the Philippines1

Ant communities were surveyed along an elevational gradient in the Philippines extending from lowland dipterocarp forest (250 m elevation) to mossy forest (1750 m). Standardized pitfall trapping in arboreal and terrestrial microhabitats at seven sites yielded 51 species. Collecting by hand at five of the sites yielded 48 species. The two methods produced substantially different assemblages, with only 22 species (29%) taken in common. Only a fraction of the total ant community appeared to be sampled at most of the sites. Measures of species richness and relative abundance peaked at mid-elevations and declined sharply with increasing elevation. Ants were extremely rare above 1500 m elevation. Arboreal ants were trapped much less frequently than terrestrial ants at all sites. Ant species that were abundant had broader elevational distributions than those that were less common, but most species were rare and occurred at only one or two sites. The elevational patterns for ants are largely the inverse of those documented for Philippine small mammals which reach their greatest diversity and abundance at high elevations where ants are rare. This suggests that the two groups may interact competitively. Some of the patterns observed or inferred from this study may apply to tropical ant communities in general, and are presented as series of testable hypotheses as a guide and stimulus for future research.     

A simple method to fit geometric series and broken stick models in community ecology and island biogeography

Species abundance distributions are widely used in explaining natural communities, their natural evolution and the impacts of environmental disturbance. A commonly used approach is that of rank-abundance distributions. Favored, biologically founded models are the geometric series (GS) and the broken stick (BS) model. Comparing observed abundance distributions with those predicted by models is an extremely time-consuming task. Also, using goodness-of-fit tests for frequency distributions (like Chi-square or Kolmogorov–Smirnov tests) to compare observed with expected frequencies is problematic because the best way to calculate expected frequencies may be controversial. More important, the Chi-square test may prove if an observed distribution statistically differs from a model, but does not allow the investigator to choose among competing models from which the observed distribution does not differ. Both models can be easily tested by regression analysis. In GS, if a log scale is used for abundance, the species exactly fall along a straight line. The BS distribution shows up as nearly linear when a log scale is used for the rank axis. Regression analysis is proposed here as a simpler and more efficient method to fit the GS and BS models. Also, regression analysis (1) does not suffer from assumptions related to Chi-square tests; (2) obviates the need to establish expected frequencies, and (3) offers the possibility to choose the best fit among competing models. A possible extension of abundance-rank analysis to species richness on islands is also proposed as a method to discriminate between relict and equilibrial models. Examples of application to field data are also presented.     

A simple method to fit geometric series and broken stick models in community ecology and island biogeography

Species abundance distributions are widely used in explaining natural communities, their natural evolution and the impacts of environmental disturbance. A commonly used approach is that of rank-abundance distributions. Favored, biologically founded models are the geometric series (GS) and the broken stick (BS) model. Comparing observed abundance distributions with those predicted by models is an extremely time-consuming task. Also, using goodness-of-fit tests for frequency distributions (like Chi-square or Kolmogorov–Smirnov tests) to compare observed with expected frequencies is problematic because the best way to calculate expected frequencies may be controversial. More important, the Chi-square test may prove if an observed distribution statistically differs from a model, but does not allow the investigator to choose among competing models from which the observed distribution does not differ. Both models can be easily tested by regression analysis. In GS, if a log scale is used for abundance, the species exactly fall along a straight line. The BS distribution shows up as nearly linear when a log scale is used for the rank axis. Regression analysis is proposed here as a simpler and more efficient method to fit the GS and BS models. Also, regression analysis (1) does not suffer from assumptions related to Chi-square tests; (2) obviates the need to establish expected frequencies, and (3) offers the possibility to choose the best fit among competing models. A possible extension of abundance-rank analysis to species richness on islands is also proposed as a method to discriminate between relict and equilibrial models. Examples of application to field data are also presented.     

Inferring local competition intensity from patch size distributions: a test using biological soil crusts

Dryland vegetation is inherently patchy. This patchiness goes on to impact ecology, hydrology, and biogeochemistry. Recently, researchers have proposed that dryland vegetation patch sizes follow a power law which is due to local plant facilitation. It is unknown what patch size distribution prevails when competition predominates over facilitation, or if such a pattern could be used to detect competition. We investigated this question in an alternative vegetation type, mosses and lichens of biological soil crusts, which exhibit a smaller scale patch‐interpatch configuration. This micro‐vegetation is characterized by competition for space. We proposed that multiplicative effects of genetics, environment and competition should result in a log‐normal patch size distribution. When testing the prevalence of log‐normal versus power law patch size distributions, we found that the log‐normal was the better distribution in 53% of cases and a reasonable fit in 83%. In contrast, the power law was better in 39% of cases, and in 8% of instances both distributions fit equally well. We further hypothesized that the log‐normal distribution parameters would be predictably influenced by competition strength. There was qualitative agreement between one of the distribution's parameters (μ) and a novel intransitive (lacking a ‘best’ competitor) competition index, suggesting that as intransitivity increases, patch sizes decrease. The correlation of μ with other competition indicators based on spatial segregation of species (the C‐score) depended on aridity. In less arid sites, μ was negatively correlated with the C‐score (suggesting smaller patches under stronger competition), while positive correlations (suggesting larger patches under stronger competition) were observed at more arid sites. We propose that this is due to an increasing prevalence of competition transitivity as aridity increases. These findings broaden the emerging theory surrounding dryland patch size distributions and, with refinement, may help us infer cryptic ecological processes from easily observed spatial patterns in the field.     

秦岭及陕北黄土区辽东栎林群落物种多样性特征

应用５种物种多样性测度指标及４种物种—多度分布模型对秦岭和黄土区辽东栎林种子植物物种多样性及木本植物种—多度分布进行了对比研究,结果表明：秦岭辽东栎林无论是群落物种多样性还是各层次的物种多样性均高于黄土区辽东栎林。秦岭辽东栎林木本植物种—多度分布符合对数正态分布而黄土区辽东栎林则符合对数级数分布,分布区气候差异是造成两类辽东栎林物种多样性差异的主要原因。物种—多度分布分析与物种多样性分析结果是一致的。     

Spatial patterns in benthic invertebrate community structure: products of habitat stability or are they habitat specific?

1. The effects of habitat stability on benthic invertebrate community structure were examined at eleven sites (ten streams and a wind-swept lake shore) with similar physicochemical characteristics but differing stability. 2. Habitat characteristics were assessed to place the study sites within the framework of the disturbance-productivity-diversity model, the intermediate disturbance hypothesis and a variant of the habitat templet model to examine their predictions with respect to community structure. 3. Many of the most common invertebrate species were present at all the study site, although their relative abundance and density differed markedly between sites. Thus, while stability did not appear to affect colonization of the study sites by these taxa, it did affect their relative success. 4. Communities at unstable sites were very similar and shared a number of taxa such as Deleatidium, Austrosimulium and several species of chironomid, presumably well-adapted to surviving and recolonizing after flood events. 5. Communities at the stable sites differed markedly, both from each other and the group of unstable sites. The characteristic fauna at each of the stable sites seemed to be a result of the site's intrinsic character and possibly biotic interactions. 6. Although stability was a pervading influence on community structure, acting as a bottleneck to the development of a site-specific suite of taxa, none of the above models could adequately explain the observed patterns.     

Frequent and occasional species and the shape of relative-abundance distributions

Recently, three different models have been proposed to explain the distribution of abundances in natural communities: the self‐similarity model; the zero‐sum ecological drift model; and the occasional–frequent species model of Magurran and Henderson. Here we study patterns of relative abundance in a large community of forest Hymenoptera and show that it is indeed possible to divide the community into a group of frequent species and a group of occasional species. In accordance with the third model, frequent species followed a lognormal distribution. Relative abundances of the occasional species could be described by the self‐similarity model, but did not follow a log‐series as proposed by the occasional–frequent model. The zero‐sum ecological drift model makes no explicit predictions about frequent and occasional species but the abundance distributions of the hymenopteran species did not show the excess of rare species predicted by this model. Separate fits of this model to the frequent and to the occasional species were worse than the respective fits of the lognormal and the self‐similarity model.     

Geographic distribution and community structure of bumblebees in the northern Iberian peninsula

Summary Relationships between species distribution and abundance, the influence of proboscis length on species-packing, and species associations within the local assemblages were studied in local communities of bumblebees in northern Spain along an altitudinal gradient. Local species abundance and altitudinal range occupied accounted for much of the variation in species distribution. Altitudinal range occupied by species was related to species distribution, but the most important variable accounting for species distribution was the local percentage abundance. Despite this, there was no evidence for age abundance. Despite this, there was no evidence for bimodality in the distributions of species incidence. A general trends for mean proboscis length in each locality to be greater in lowland localities exists, but this variable was not related to species distribution or abundance. Proboscis length spacings were studied among species in local assemblages and in most of the cases observed spacing did not differ from random expectations. The same patterns were demonstrated calculating spacings for core species in each local assemblage. Furthermore, species showed little tendency towards associations, so it may be concluded that bumblebee assemblages were irregularly structured and no clear patterns emerged from the present study.     

The distribution patterns of dragonflies (Insecta: Odonata) along the Kiewa River,Australia, and their relevance in conservation assessment

Sampling of larval and adult Odonata from 16 sites along the Kiewa River, Victoria, yielded 34 species: 10 Zygoptera, 24 Anisoptera. Patterns of larval and adult incidence were appraised, and showed that most species were restricted in incidence to several consecutive sites along the river, and that there is clear distinction also between the faunas of the potamon, rhithron and eucrenon regions. Different species of some genera of Anisoptera displayed different zonational distributions, and patterns of incidence and relative abundance of larvae and adults confirmed zonational occupancy. For larvae, these distribution patterns transcended the mode of collection, although many species were found most abundantly in one microhabitat or by one of several sampling methods employed at each site. Sampling of the two stages separately showed considerable concurrence of distributional patterns, so that either stage alone may provide data of value in faunal and conservation assessment. This revised version was published online in July 2006 with corrections to the Cover Date.     

Relationships between local population persistence, local abundance and regional occupancy of species: distribution patterns of diatoms in boreal streams

Aims We have two aims: (1) to examine the relationship between local population persistence, local abundance and regional occupancy of stream diatoms and (2) to characterize the form of the species–occupancy frequency distribution of stream diatoms. Location Boreal streams in Finland. There were three spatial extents: (1) across ecoregions in Finland, (2) within ecoregions in Finland, and (3) within a single drainage system in southern Finland. Methods Diatoms were sampled from stones (epilithon), sediment (epipelon) and aquatic plants (epiphyton) in streams using standardized sampling methods. To assess population persistence, diatom sampling was conducted monthly at four stream sites from June to October. The relationships between local population persistence, local abundance and regional occupancy were examined using correlation analyses. Results There was a significant positive relationship between local persistence and abundance of diatoms in epilithon, epipelon and epiphyton. Furthermore, local abundance and regional occupancy showed a significant positive relationship at multiple spatial extents; that is, across ecoregions, within ecoregions and within a drainage system. The relationships between occupancy and abundance did not differ appreciably among impacted and near pristine‐reference sites. The occupancy–frequency distribution was characterized by a large number of satellite species which occurred at only a few sites, whereas core species that occurred at most sites were virtually absent. Main conclusions The positive relationship between local population persistence and abundance suggested that a high local abundance may prevent local extinction or that high persistence is facilitated by a high local cell density. High local persistence and local abundance may also positively affect the degree of regional occupancy in stream diatoms. The results further showed that anthropogenic effects were probably too weak to bias the relationship between occupancy and abundance, or that the effects have already modified the distribution patterns of stream diatoms. The small number of core species in the species–occupancy frequency distribution suggested that the regional distribution patterns of stream diatoms, or perhaps unicellular microbial organisms in general, may not be fundamentally different from those described previously for multicellular organisms, mainly in terrestrial environments, although average global range sizes may differ sharply between these two broad groups of organisms.