Which Models Are Appropriate for Six Subtropical Forests: Species-Area and Species-Abundance Models

The species-area relationship is one of the most important topic in the study of species diversity, conservation biology and landscape ecology. The species-area relationship curves describe the increase of species number with increasing area, and have been modeled by various equations. In this paper, we used detailed data from six 1-ha subtropical forest communities to fit three species-area relationship models. The coefficient of determination and F ratio of ANOVA showed all the three models fitted well to the species-area relationship data in the subtropical communities, with the logarithm model performing better than the other two models. We also used the three species-abundance distributions, namely the lognormal, logcauchy and logseries model, to fit them to the species-abundance data of six communities. In this case, the logcauchy model had the better fit based on the coefficient of determination. Our research reveals that the rare species always exist in the six communities, corroborating the neutral theory of Hubbell. Furthermore, we explained why all species-abundance figures appeared to be left-side truncated. This was due to subtropical forests have high diversity, and their large species number includes many rare species.     

闽北地区森林群落物种多样性的测定

采用8个物种多样性指数对闽北地区的8个森林群落类型物种多样性进行测定,并选用4个种一多度关系模型(对数正态分布、Weibull分布、对数级数分布、生态位优先占领模型)分析其物种-多度关系,把多样性指数与对数正态分布、Weibull分布和生态位优先占领模型的有关参数进行线性回归,以分析多度模型参数描述物种多样性的可行性。结果表明：(1)多数物种多样性指数对群落的测度是一致的,(2)对数正态分布、Weibull分布和生态位优先占领模型对8个群落的物种多度拟合效果很好;(3)对数正态分布、Weibull分布和生态位优先占领模型有关模型参数与部分物种多样性指数的线性关系达显著或极显著水平。通过闽北地区8个类型森林群落物种多样性指数测定,使生物多样性较准确地数量化,同时还说明采用改进单纯形法进行非线性分布函数的拟合是简单有效的,可推广应用。     

不同海拔高程梯度下鄱阳湖典型草洲植物群落物种-多度分布格局的模型拟合

基于恒湖农场茶叶港草洲4个样带、19个样地（5 m×5 m）沿海拔高程（高高程-湖边、中高程-洲滩中间、低高程-洲滩边缘）调查得到的植被数据,用4种生态位模型拟合研究该区域不同海拔高程段植物种-多度分布格局。结果表明：（1）随着海拔高程下降,群落偶见种不断减少,且平均盖度均较低;常见种数目保持不变,其中优势种灰化薹草（Carex cinerascens Kukenth）平均盖度、平均高度持续升高,在群落中保持较高的优势地位;（2）3个高程段内生态位优先占领模型对群落物种-多度分布格局有较高的拟合优度（P < 0.05,AIC值分别为-149.41、-61.11、-25.63）,断棍模型、重叠生态位模型、优势优先模型总体上拟合效果较差（P < 0.05,但AIC值均较大）。为研究区植物群落生物多样性的保护和构建机制的探索提供了一定参考。     

综防区和化防区稻田害虫-天敌群落特性的研究——种-多度关系及群落统计量的生态效应

对数级数模式是描述稻田害虫-天敌群落种-多度分布的理想模式。资料表明两个呈单一对数级数分布的群落经混合扩大后仍为对数级数分布.Shannon-weaver多样性指数比其他指数有较强的判别力。但在大样本中,对群落组成的变化亦不敏感,采用对数级数分布的参数(?)作为多样性统计量较为合适。     

When and why do non-neutral metacommunities appear neutral?

Hubbell's neutral theory assumes that all species in a community have the same per capita fitness. Despite the overwhelming evidence against this assumption in most communities the neutral theory has often been, though not always, successful at predicting patterns of diversity in nature. I analyze a non-neutral model in order to suggest conditions under which observed species-abundance distributions (SADs) could be expected to resemble neutral distributions. The non-neutral model consists of two guilds of species such that (1) individuals between guilds do not interact, (2) dynamics within guilds follow Hubbell's model and (3) neutral parameters between guilds differ. This two-guild model generates SADs that appear neutral in some cases and clearly non-neutral in other cases. This result suggests that SADs may be more informative about niche structure than previously thought. The two-guild model could be tested in communities composed of fairly well-defined guilds or functional groups.     

Multiple modes in a coral species abundance distribution

Species abundance distributions are an important measure of biodiversity and community structure. These distributions are affected by sampling, and alternative species-abundance models often make similar predictions for small sample sizes. Very large samples reveal the relative abundances of rare species, and thus provide information about species relative abundances that small samples cannot. Here, we present the species-abundance distribution for a sample of > 40,000 coral colonies at a single site, exceeding existing samples of coral local assemblages by over an order of magnitude. This abundance distribution is multimodal when examined on a logarithmic scale. Four different model selection procedures all indicate that the underlying community abundance distribution has at least three modes. We show that the multiple modes are not caused by mixtures of species with different habitat preferences. However, spatial aggregation partially explains our results. We inspect published work on species abundance distributions, and suggest that multimodality may be a common feature of large samples.     

长白山不同演替阶段针阔混交林群落物种多度分布格局

为解释长白山温带森林群落构建和物种多度格局的形成过程, 该文以不同演替阶段的针阔混交林监测样地数据为基础, 采用中性理论模型、生物统计模型(对数正态分布模型)和生态位模型(Zifp模型、分割线段模型、生态位优先模型)拟合森林群落物种多度分布, 并用χ ²检验、Kolmogorov-Smirnov (K-S)检验和赤池信息准则(AIC)选择最佳拟合模型。结果显示: 中性模型能很好地预测长白山温带森林不同演替阶段植物群落的物种多度分布。在10 m × 10 m尺度上, 5种模型均可被χ ²检验和K-S检验接受, 但中性模型拟合效果不如对数正态分布模型、Zifp模型、分割线段模型和生态位优先模型, 表明小尺度上中性过程和生态位过程均能解释群落物种多度分布, 但生态位过程的解释能力相对较大。而在中大尺度上(30 m × 30 m、60 m × 60 m和90 m × 90 m), 中性模型为最优拟合模型, 并且随着研究尺度增加, 生态位模型和生物统计模型逐渐被χ ²检验拒绝, 表明中性过程在长白山针阔混交林群落物种多度分布格局形成中的作用随着研究尺度增加而逐渐增大。该文证实了中性过程在长白山温带针阔混交林群落结构形成中具有重要作用, 但未否认生态位机制在群落构建中的贡献。因此, 温带森林群落构建过程中中性理论和生态位理论并非相互矛盾, 而是相互融合的。在研究森林群落物种多度分布时, 应重视取样尺度和演替阶段的影响, 并采用多种模型进行拟合。     

The stochastic community and the logistic-J distribution

A new formal model called the multispecies logistical (MSL) system produces species/abundance distributions that are compared statistically with those found in natural communities. The system, which is able to handle thousands of individuals from hundreds of species, iteratively selects random pairs of individuals and transfers a unit of biomass (or energy) between the respective species. Several elaborations of the model, including those with trophic compartments, appear to produce the same distribution. The theoretical distribution underlying the MSL system is a hyperbolic section, here called the logistic-J distribution. In the study reported here, the logistic-J distribution has been fitted to the species-abundance histograms of 125 randomly selected taxocoenoses. Since the overall chi square score of the logistic-J achieved near-optimality in this study, it cannot be distinguished statistically from the J-curves observed by field biologists. For comparison, the log-series distribution was given the same test and scored significantly higher (more poorly) than the mean logistic-J score. If there is a single, major distribution underlying natural communities, it is not the log-series distribution. Nor, owing to a mathematical error in its formulation, can it be the lognormal distribution. In the MSL system each species follows a “stochastic orbit” about the mean abundance producing, in consequence, the logistic-J distribution. Such orbits are produced by any system in which the probabilities of reproduction and death are approximately equal. Accordingly, the “stochastic communities hypothesis” is proposed here as the overall mechanism governing abundances in all natural communities. It is not a single mechanism, per se, but the net effect of all environmental influences.     

太白山锐齿栎林物种-多度分布格局

以太白山1.5 hm²的锐齿栎原始林和次生林样地中环境因子和胸径≥1 cm的木本植物调查数据为基础,采用统计模型(对数正态模型)、生态位模型(Zipf模型、断棍模型、生态位优先模型)和中性模型,拟合了锐齿栎群落的物种多度分布。结果表明: 太白山锐齿栎林物种多度分布格局受到生境异质性的影响。其中,地形因子对原始林物种分布影响较大,在凹凸度较大的生境中,物种分布同时受到中性过程和生态位过程的影响,但中性过程发挥的作用较小;而在凹凸度较小的生境中,中性模型被拒绝,物种的多度分布符合生态位理论的假设。在群落坡度大的区域,群落中生态位过程和中性过程同等重要;而在坡度较小的平缓区域,生态位分化对群落物种分布的影响较大。在次生林中,影响物种分布的因素主要是土壤养分。在次生林土壤速效磷含量高的生境中,生态位过程是影响群落物种分布的主要生态学过程;而在土壤速效磷含量低的生境中,中性过程和生态位过程在群落物种分布中同时存在。太白山锐齿栎林物种多度分布格局存在明显的尺度效应。原始林在20 m×20 m尺度上,生态位模型和中性模型都能预测物种多度分布,而在40 m×40 m和70 m×70 m尺度上,生态位过程可解释物种多度分布格局。在次生林样地20 m×20 m、40 m×40 m、70 m×70 m尺度上,生态位过程和中性过程共同作用于物种的分布,但是生态位过程更为重要。可见,除了尺度和生境异质性外,原始林与受干扰的次生林中的物种多度分布也存在明显的差异。     

A statistical theory for sampling species abundances

The pattern of species abundances is central to ecology. But direct measurements of species abundances at ecologically relevant scales are typically unfeasible. This limitation has motivated a long-standing interest in the relationship between the abundance distribution in a large, regional community and the distribution observed in a small sample from the community. Here, we develop a statistical sampling theory to describe how observed patterns of species abundances are influenced by the spatial distributions of populations. For a wide range of regional-scale abundance distributions we derive exact expressions for the sampled abundance distributions, as a function of sample size and the degree of conspecific spatial aggregation. We show that if populations are randomly distributed in space then the sampled and regional-scale species-abundance distribution typically have the same functional form: sampling can be expressed by a simple scaling relationship. In the case of aggregated spatial distributions, however, the shape of a sampled species-abundance distribution diverges from the regional-scale distribution. Conspecific aggregation results in sampled distributions that are skewed towards both rare and common species. We discuss our findings in light of recent results from neutral community theory, and in the context of estimating biodiversity.     

Toward ecologically explicit null models of nestedness

A community is "nested" when species assemblages in less rich sites form nonrandom subsets of those at richer sites. Conventional null models used to test for statistically nonrandom nestedness are under- or over-restrictive because they do not sufficiently isolate ecological processes of interest, which hinders ecological inference. We propose a class of null models that are ecologically explicit and interpretable. Expected values of species richness and incidence, rather than observed values, are used to create random presence-absence matrices for hypothesis testing. In our examples, based on six datasets, expected values were derived either by using an individually based random placement model or by fitting empirical models to richness data as a function of environmental covariates. We describe an algorithm for constructing unbiased null matrices, which permitted valid testing of our null models. Our approach avoids the problem of building too much structure into the null model, and enabled us to explicitly test whether observed communities were more nested than would be expected for a system structured solely by species-abundance and species-area or similar relationships. We argue that this test or similar tests are better determinants of whether a system is truly nested; a nested system should contain unique pattern not already predicted by more fundamental ecological principles such as species-area relationships. Most species assemblages we studied were not nested under these null models. Our results suggest that nestedness, beyond that which is explained by passive sampling processes, may not be as widespread as currently believed. These findings may help to improve the utility of nestedness as an ecological concept and conservation tool.     

A THEORY OF FAUNAL BUILDUP FOR COMPETITION COMMUNITIES

Invasion-structured communities have more species than do coevolution-structured communities assembled using the same resource distribution. Species in invasion-structured communities are tightly packed, occupying the upper portion of the resource axis; species in coevolution-structured communities are more widely spaced, and most are located in the lower portion of the resource axis. As a consequence, coevolution-structured communities tend to be more stable than comparable invasion-structured communities, but more open to invasion. Both invasion-structured and coevolution-structured communities have niche separations that are significantly different than would be expected if species were assorted at random. Two-species communities formed by the invasion-only algorithm under asymmetric competition had the majority of their niche separations in the range 0–0.5. All other communities had niche separations that were greater than expected. The most common separations were in the range 1.0–3.5. Thus, while not a common feature of many communities, nicheseparation patterns similar to those described by Hutchinson (1959) appear as an “ensemble” property of many communities. The faunal-buildup graphs formed by the coevolutionary algorithm differ from those formed by the invasion-only algorithm, showing community cycling whenever asymmetric competition is present. Through this cycling behavior the coevolutionary faunal-buildup algorithm provides both a theoretical basis for Wilson's (1959) taxon cycle and a hypothesis explaining the distribution of Anolis lizards in the Lesser Antilles.     

Geographical gradients of stability in European land bird communities

Summary European data from 15 long-term censuses of breeding land bird communities were analysed in order to measure community stability. Stability was here operationally defined as year-to-year persistence of community structure (total density, number of species, diversity, evenness of the species-abundance distribution, species list, and frequencies of species). Central Europe and southern Scandinavia formed a zone of fairly stable communities, but more northern communities were fairly unstable. This result contrasts with several recent arguments, but accords well with the classic dogma that instability of biological communities increases northwards in the northern hemisphere.Three theoretical explanations for the difference in stability between northern and southern communities were explored, and causal mechanisms underlying the hypotheses were specified and tested. Considerable evidence supported the hypothesis that environmental (climatic) unpredictability increases northwards and mainly causes the phenomenon observed. Gradients in productivity may, perhaps in interaction with unpredictability, also contribute to geographical differences in stability, but tests suggested that low diversity is not an important cause of northern instability.     

群落结构特性的分析方法探讨-以高寒草甸和高寒灌丛为例

 从种—多度关系和物种多样性对高寒草甸和高寒灌丛在不同放牧强度下群落结构特性的分析表明,种在群落中的相对重要性发生了变化,种相对多度的分布模式符合对数正态分布,并随放牧强度的变化显示出有规律的改变;同时,表示群落结构特性的指数(Simpson多样性指数λ、Shannon信息多样性指数H′、McIntosh多样性指数Dmc、均匀性指数E和物种丰富度指数R)值在各放牧强度群落间却未表现出统计学上的显著差异(P>0.05)。这表明在有些情况下单纯对物种多样性等指数值的比较并不一定能准确地反映出群落结构上的变化,同时也证实了West关于多样性指数值保持不变或近似但群落结构可能发生变化的推断。     

大熊猫栖息地地震干扰后恢复群落的中性理论验证

为了验证中性理论在干扰后恢复群落中的适用性,本文利用不同地震干扰强度下大熊猫栖息地恢复群落的野外调查数据进行了中性理论的验证。预测发现,在干扰较重及未受干扰的原始林群落中,中性理论模型能够较好地预测物种多度分布,但在干扰相对较轻的恢复群落中中性理论模型预测物种多度分布效果较差。原因分析表明,生境的均一性程度是影响中性理论模型预测物种多度分布效果的重要影响因子。因此可认为,中性理论不适用于生境均一性差的群落。     

EVOLUTION,PHYLOGENY, SEXUAL DIMORPHISM AND MATING SYSTEM IN THE GRACKLES (QUISCALUS SPP.: ICTERINAE)

According to theory, two consequences of sexual selection are sexual dimorphism in size and secondary sexual characteristics, due to either intra- or intersexual selection. In this paper I suggest three criteria for the test of an evolutionary hypothesis involving quantitative morphological characters. First, the postulated change must be shown to have occurred in evolutionary time. Second, this change must be positively correlated with a change in the proposed selective agent. Third, given two taxa with different degrees of sexual size dimorphism and different mating system, the possible influence of drift must be rejected. If the hypothesis is not rejected by these three criteria, then we still have no proof of causality, but we can at least be more confident about its plausibility. This is applied to the particular hypothesis that sexual dimorphism in the Boat-tailed and Great-tailed grackles (Quiscalus spp; Icterinae; Aves) is caused by the highly polygynous mating system in these species. In relation to an outgroup, both species have increased disproportionately in male tarsus and tail size, creating an increased sexual dimorphism. This has cooccurred with the evolution of their particular mating system. However, the variance among species in male tarsus size can be accounted for by drift, and need not be a result of selection for increased size. In contrast, the variance among species in male tail size was much larger than expected under a null model of drift, indicating directional selection for long tails. The variance in female tail size was not larger than expected by drift, whereas the variance in female tarsus size was in fact lower than expected by drift, indicating stabilizing selection. The data are consistent with the hypothesis with regard to tail size, but not with regard to body size.     

Testing the standard neutral model of biodiversity in lake communities

Hubbell's (2001) neutral model describes how local communities are structured if population dynamics are statistically identical among species in a constant, possibly patchy, environment with random speciation. Tests of this model have been restricted largely to terrestrial communities. Here we tested the fit of this neutral model to fish, zooplankton and phytoplankton species–abundance distributions from 30 well-studied lake communities varying widely in lake size and productivity. We measured the fit of the communities to the neutral model in three ways. All but two zooplankton (7 of 9) and all but three fish (9 of 12) communities were consistent with all three measures of fit. However, all nine phytoplankton communities did not fit the neutral model by at least one measure. This result for phytoplankton communities represents to date the most consistent failure of the standard neutral model to predict the shape of species-abundance distributions.     

A comparative analysis of alternative approaches to fitting species-abundance models

Aims Fits of species-abundance distributions to empirical data are increasingly used to evaluate models of diversity maintenance and community structure and to infer properties of communities, such as species richness. Two distributions predicted by several models are the Poisson lognormal (PLN) and the negative binomial (NB) distribution; however, at least three different ways to parameterize the PLN have been proposed, which differ in whether unobserved species contribute to the likelihood and in whether the likelihood is conditional upon the total number of individuals in the sample. Each of these has an analogue for the NB. Here, we propose a new formulation of the PLN and NB that includes the number of unobserved species as one of the estimated parameters. We investigate the performance of parameter estimates obtained from this reformulation, as well as the existing alternatives, for drawing inferences about the shape of species abundance distributions and estimation of species richness.Methods We simulate the random sampling of a fixed number of individuals from lognormal and gamma community relative abundance distributions, using a previously developed 'individual-based' bootstrap algorithm. We use a range of sample sizes, community species richness levels and shape parameters for the species abundance distributions that span much of the realistic range for empirical data, generating 1?000 simulated data sets for each parameter combination. We then fit each of the alternative likelihoods to each of the simulated data sets, and we assess the bias, sampling variance and estimation error for each method.Important findings Parameter estimates behave reasonably well for most parameter values, exhibiting modest levels of median error. However, for the NB, median error becomes extremely large as the NB approaches either of two limiting cases. For both the NB and PLN,>90% of the variation in the error in model parameters across parameter sets is explained by three quantities that corresponded to the proportion of species not observed in the sample, the expected number of species observed in the sample and the discrepancy between the true NB or PLN distribution and a Poisson distribution with the same mean. There are relatively few systematic differences between the four alternative likelihoods. In particular, failing to condition the likelihood on the total sample sizes does not appear to systematically increase the bias in parameter estimates. Indeed, overall, the classical likelihood performs slightly better than the alternatives. However, our reparameterized likelihood, for which species richness is a fitted parameter, has important advantages over existing approaches for estimating species richness from fitted species-abundance models.     

Gradient analysis of spider communities in a streamside forest

Summary Species composition and diversity of a guild of wandering spiders was studied by pitfall trapping over an elevational gradient in an Illinois streamside forest. Differences in flooding frequency and their effect on the litter habitat (removal and/or compression) account for a majority of the variation in the number of species between elevations. Changes in spider communities with elevation over the flooding gradient are indicative of a transition from a harsh to a moderate environment: (1) increased abundance and species diversity; (2) decreased dominance of flood tolerant species accompanied by increased dominance of species with specialized microhabitats found in complex litter; (3) greater similarity in species composition between sites; and, (4) a change in species-abundance curves from a geometric series to a lognormal distribution. The influence of the flooding regime in regulating community structure of spiders is discussed. A multiple regression equation including flood frequency and litter depth as variables was used to predict the impact of altered flooding regimes (due to reservoir construction downstream) on spider diversity.