Long-term demographic surveys reveal a consistent relationship between average occupancy and abundance within local populations of a butterfly metapopulation

Species distribution models are the tool of choice for large-scale population monitoring, environmental association studies and predictions of range shifts under future environmental conditions. Available data and familiarity of the tools rather than the underlying population dynamics often dictate the choice of specific method – especially for the case of presence–absence data. Yet, for predictive purposes, the relationship between occupancy and abundance embodied in the models should reflect the actual population dynamics of the modelled species. To understand the relationship of occupancy and abundance in a heterogeneous landscape at the scale of local populations, we built a spatio-temporal regression model of populations of the Glanville fritillary butterfly Melitaea cinxia in a Baltic Sea archipelago. Our data comprised nineteen years of habitat surveys and snapshot data of land use in the region. We used variance partitioning to quantify relative contributions of land use, habitat quality and metapopulation covariates. The model revealed a consistent and positive, but noisy relationship between average occupancy and mean abundance in local populations. Patterns of abundance were highly variable across years, with large uncorrelated random variation and strong local population stochasticity. In contrast, the spatio-temporal random effect, habitat quality, population connectivity and patch size explained variation in occupancy, vindicating metapopulation theory as the basis for modelling occupancy patterns in fragmented landscapes. Previous abundance was an important predictor in the occupancy model, which points to a spillover of abundance into occupancy dynamics. While occupancy models can successfully model large-scale population structure and average occupancy, extinction probability estimates for local populations derived from occupancy-only models are overconfident, as extinction risk is dependent on actual, not average, abundance.     

The wealth of species: ecological communities, complex systems and the legacy of Frank Preston

General statistical patterns in community ecology have attracted considerable recent debate. Difficulties in discriminating among mathematical models and the ecological mechanisms underlying them are likely related to a phenomenon first described by Frank Preston. He noted that the frequency distribution of abundances among species was uncannily similar to the Boltzmann distribution of kinetic energies among gas molecules and the Pareto distribution of incomes among wage earners. We provide additional examples to show that four different 'distributions of wealth' (species abundance distributions, species–area and species–time relations, and distance decay of compositional similarity) are not unique to ecology, but have analogues in other physical, geological, economic and cultural systems. Because these appear to be general statistical patterns characteristic of many complex dynamical systems they are likely not generated by uniquely ecological mechanistic processes.     

Biogeography of parasite abundance: latitudinal gradient and distance decay of similarity in the abundance of fleas and mites,parasitic on small mammals in the Palearctic,at three spatial scales

We tested whether biogeographic patterns characteristic for biological communities can also apply to populations and investigated geographic patterns of variation in abundance of ectoparasites (fleas and mites) collected from bodies of their small mammalian hosts (rodents and shrews) in the Palearctic at continental, regional and local scales. We asked whether (i) there is a relationship between latitude and abundance and (ii) similarity in abundance follows a distance decay pattern or it is better explained by variation in extrinsic biotic and abiotic factors. We analysed the effect of latitude on mean intraspecific abundance using general linear models including proportional abundance of its principal host as an additional predictor variable. Then, we examined the relative effect of geographic distance, biotic and abiotic dissimilarities among regions, subregions or localities on the intraspecific dissimilarity in abundance among regions, subregions or localities using Generalized Dissimilarity Modelling. We found no relationship between latitude and intraspecific flea or mite abundance. In both taxa, environmental dissimilarity explained the largest part of the deviance of spatial variation in abundance, whereas the effect of the dissimilarity in the principal host abundance was of secondary importance and the effect of geographic distance was minor. These patterns were generally consistent across the three spatial scales, although environmental variation and dissimilarity in principal host abundance were equally important at the local scale in fleas but not in mites. We conclude that biogeographic patterns related to latitude and geographic distance do not apply to spatial variation of ectoparasite abundance. Instead, the geographic distribution of abundance in arthropod ectoparasites depends on their responses, mainly to the off-host environment and to a lesser extent the abundance of their principal hosts.     

Shotgun mitogenomics across body size classes in a local assemblage of tropical Diptera: Phylogeny,species diversity and mitochondrial abundance spectrum

Mitochondrial genomes can be assembled readily from shotgun‐sequenced DNA mixtures of mass‐trapped arthropods (“mitochondrial metagenomics”), speeding up the taxonomic characterization. Bulk sequencing was conducted on some 800 individuals of Diptera obtained by canopy fogging of a single tree in Borneo dominated by small (<1.5 mm) individuals. Specimens were split into five body size classes for DNA extraction, to equalize read numbers across specimens and to study how body size, a key ecological trait, interacts with species and phylogenetic diversity. Genome assembly produced 304 orthologous mitochondrial contigs presumed to each represent a different species. The small‐bodied fraction was the by far most species‐rich (187 contigs). Identification of contigs was through phylogenetic analysis together with 56 reference mitogenomes, which placed most of the Bornean community into seven clades of small‐bodied species, indicating phylogenetic conservation of body size. Mapping of shotgun reads against the mitogenomes showed wide ranges of read abundances within each size class. Ranked read abundance plots were largely log‐linear, indicating a uniformly filled abundance spectrum, especially for small‐bodied species. Small‐bodied species differed greatly from other size classes in neutral metacommunity parameters, exhibiting greater levels of immigration, besides greater total community size. We suggest that the established uses of mitochondrial metagenomics for analysis of species and phylogenetic diversity can be extended to parameterize recent theories of community ecology and biodiversity, and by focusing on the number mitochondria, rather than individuals, a new theoretical framework for analysis of mitochondrial abundance spectra can be developed that incorporates metabolic activity approximated by the count of mitochondria.     

Assessing latitudinal gradients in speciation rates and biodiversity at the global scale

The mechanisms responsible for latitudinal biodiversity gradients have fascinated and perplexed biologists since the time of Darwin. Ecological theory has yielded two general classes of mechanisms to account for variation in biodiversity: dispersal–assembly mechanisms that invoke differences in stochastic rates of speciation, extinction and dispersal; and niche–assembly mechanisms that invoke species differences, species interactions and environmental heterogeneity. Distinguishing between these two classes of mechanisms requires explicit consideration of macroevolutionary dynamics. Here, we assess the importance of dispersal–assembly mechanisms in the origin and maintenance of biodiversity using fossil data that encompass 30 million years of macroevolution for three distinct groups of ocean plankton: foraminifera, nannoplankton and radiolaria. Applying new methods of analysis to these fossil data, we show here for the first time that latitudinal biodiversity gradients exhibit strong positive correlations with speciation rates even after explicitly controlling for variation in sampling effort and for increases in habitat area towards the equator. These findings provide compelling evidence that geographical variation in macroevolutionary dynamics is a primary determinant of contemporary biodiversity gradients, as predicted by dispersal–assembly theory.     

生态学代谢理论: 基于个体新陈代谢过程解释物种多样性的地理格局

新陈代谢是生物的基本生理过程。生态学代谢理论(metabolic theory of ecology)基于生物个体大小和环境温度对个体新陈代谢过程的影响, 使用尺度推移(scaling)的方法来解释多种生态学格局和过程。James Borwn等将这一理论用于解释物种多样性的大尺度格局, 并从机制上解释了物种多样性与环境温度的关系。这一理论提出了两个明确的预测: (1)物种多样性的对数与绝对温度的倒数之间呈线性关系; (2) 这一线性关系的斜率为–0.6至–0.7。这一理论自提出以来, 饱受争议, 经过了正反两方面经验数据的检验, 目前仍未形成一致的结论。虽然这一理论仍面临着一些有待解决的问题, 但它以崭新的思路和方法有别于以往的基于统计学方法的研究。人们过去对该理论的实证检验忽略了两个重要的约束条件, 即除温度以外的环境条件处于理想状态和群落处于平衡状态, 而这两个约束条件是理解该理论的基础。本文对生态学代谢理论的理论框架、预测和含义, 以及以往的检验结果进行阐述, 在此基础上提出了作者对该理论的若干认识和未来研究中应考虑的若干问题。     

面积、温度及分布区限制对物种丰富度海拔格局的影响: 以秦岭太白山为例

 物种丰富度的分布格局及其形成机制是生态学研究的热点。以往的研究主要描述丰富度的格局, 而对其形成机制研究较少, 且主要集中于探讨单个因子或过程的影响。物种丰富度同时受到多个因子和过程的综合作用, 面积、温度及物种分布区限制被认为是控制山地物种丰富度海拔格局的主要因素, 三者同时沿海拔梯度而变化, 同时作用于丰富度的海拔格局。幂函数种-面积关系(SAR)、生态学代谢理论(MTE)及中域效应假说(MDE)分别基于以上3个因素, 从机制上解释了物种丰富度 的海拔格局。探讨这些假说的相对影响对研究物种丰富度的大尺度格局及其形成机制具有重要意义。方差分离方法有利于分解不同因素的影响, 为此, 该文以秦岭太白山的植物物种丰富度为例, 采用方差分离和逐步回归方法, 分析了SAR、MTE及MDE对物种丰富度海拔格局的影响。结果表明, 太白山的植物物种丰富度沿海拔梯度呈单峰分布格局, 但丰富度峰值存在类群差异; 对太白山所有植物物种丰富度的垂直格局而言, SAR、MTE及MDE分别解释了其物种丰富度随海拔变化的66.4%、19.8%和37.9%, 共同解释了84.6%, 在消除其他因素的影响后, SAR和MTE的独立影响较高(分别为25.5%和17.7%), 而MDE的独立影响不显著; 分类群研究则发现, 苔藓植物丰富度的海拔格局主要受MDE的影响, 蕨类植物丰富度的海拔格局同时受到SAR、MTE以及MDE的影响, 而种子植物物种丰富度的海拔格局主要受SAR和MTE影响。     

Small area and low connectivity constrain the diversity of plant life strategies in temporary ponds

Aim

(i) To determine whether area and connectivity of temporary ponds can predict plant species diversity, and the diversity and abundance of different plant life histories; (ii) To explore whether pond connectivity with the river prior to river regulation predicts better plant diversity patterns than current pond connectivity, suggestive of possible effects of connectivity loss.

Location

Eastern Carpathian Mountains, Romania, Europe.

Methods

We fitted linear and generalized linear models (LM and GLM) to examine whether pond area and current distance from the Olt River predict plant species richness, Shannon diversity and relative cover of different social behaviour types and overall plant species richness and Shannon diversity. Using historical maps, we measured pond distance from the river ca. 60 years before the Olt River was regulated, and we refitted the LM and GLM models using pond area and past distance from the river as independent variables.

Results

Total plant species richness increased with pond area, and it decreased with the distance from the river, but total plant Shannon diversity index was affected, positively, only by pond area. The strength of responses to pond area and connectivity of species richness, Shannon diversity and relative cover varied across the different social behaviour types. Past and current distances between ponds and riverbeds had similar effects on plant diversity, with some evidence for stronger effect of the present connectivity on specialist species Shannon diversity and a weaker effect on disturbance tolerants, generalists and competitors.

Main Conclusions

Pond area and connectivity with the landscape are important predictors of the diversity of plant life history strategies, and therefore, useful tools in pond conservation. Consistent species richness and Shannon diversity responses of wetland specialists to pond area and connectivity make this life history type well suited for monitoring pond condition.     

A general framework for neutral models of community dynamics

Neutral models of community dynamics are a powerful tool for ecological research, but their applications are currently limited to unrealistically simple types of dynamics and ignore much of the complexity that characterize natural ecosystems. Here, we present a new analytical framework for neutral models that unifies existing models of neutral communities and extends the applicability of existing models to a much wider spectrum of ecological phenomena. The new framework extends the concept of neutrality to fitness equivalence and in spite of its simplicity explains a wide spectrum of empirical patterns of species diversity including positive, negative and unimodal productivity–diversity relationships; gradual and highly delayed declines in species diversity with habitat loss; and positive and negative responses of species diversity to habitat heterogeneity. Surprisingly, the abundance distribution in all of these cases is given by the dispersal limited multinomial (DLM), the abundance distribution in Hubbell's zero-sum model, showing DLM's robustness and demonstrating that it cannot be used to infer the underlying community dynamics. These results support the hypothesis that ecological communities are regulated by a limited set of fundamental mechanisms much simpler than could be expected from their immense complexity.     

Global species–energy relationship in forest plots: role of abundance,temperature and species climatic tolerances

Aim To evaluate the strength of evidence for hypotheses explaining the relationship between climate and species richness in forest plots. We focused on the effect of energy availability which has been hypothesized to influence species richness: (1) via the effect of productivity on the total number of individuals (the more individuals hypothesis, MIH); (2) through the effect of temperature on metabolic rate (metabolic theory of biodiversity, MTB); or (3) by imposing climatic limits on species distributions. Location Global. Methods We utilized a unique ‘Gentry‐style’ 370 forest plots data set comprising tree counts and individual stem measurements, covering tropical and temperate forests across all six forested continents. We analysed variation in plot species richness and species richness controlled for the number of individuals by using rarefaction. Ordinary least squares (OLS) regression and spatial regressions were used to explore the relative performance of different sets of environmental variables. Results Species richness patterns do not differ whether we use raw number of species or number of species controlled for number of individuals, indicating that number of individuals is not the proximate driver of species richness. Productivity‐related variables (actual evapotranspiration, net primary productivity, normalized difference vegetation index) perform relatively poorly as correlates of tree species richness. The best predictors of species richness consistently include the minimum temperature and precipitation values together with the annual means of these variables. Main conclusion Across the world's forests there is no evidence to support the MIH, and a very limited evidence for a prominent role of productivity as a driver of species richness patterns. The role of temperature is much more important, although this effect is more complex than originally assumed by the MTB. Variation in forest plot diversity appears to be mostly affected by variation in the minimum climatic values. This is consistent with the ‘climatic tolerance hypothesis’ that climatic extremes have acted as a strong constraint on species distribution and diversity.     

Latitude,tree species diversity and the metabolic theory of ecology

The latitudinal diversity gradient (LDG) has been known for over a century, but its origin remains poorly understood. Because both latitude and species richness are broadly related to temperature, environmental temperature has been proposed as a driver of the LDG. Recently, Wang et al. (2009, Proceedings of the National Academy of Sciences USA, 106 ,13388–13392) used datasets compiled from tree distributions in eastern Asia and North America to compare the species richness?temperature relationship between the two regions at several spatial scales and framed their analyses in the context of the metabolic theory of ecology. Here, we show that their datasets lack comparability between eastern Asia and North America and that some aspects of their analyses probably biased their results, casting doubt on some of their conclusions.     

Disentangling the effects of geographic distance and environmental dissimilarity on global patterns of species turnover

Aim To distinguish the effects of geographic distance and environmental dissimilarity on global patterns of species turnover in four classes of terrestrial vertebrates (mammals, birds, reptiles and amphibians). Location Six hundred and sixty terrestrial ecoregions across the globe. Methods We calculated species turnover between each pair of ecoregions, using the Jaccard index (J). We selected seven variables to quantify environment in each ecoregion, and subjected the environmental values to a principal components analysis. For each realm, we applied multiple regression analysis relating the natural logarithm of the Jaccard index (lnJ) to geographic distance alone and in combination with differences in the environment variables measured as principal components (PC). We used partial correlations to partition variance in lnJ between unique contributions of distance and environmental PC scores, the covariation between distance and environment, and unexplained variance. To examine the latitude and species turnover relationship, we regressed lnJ on latitude with distance between ecoregions being included as a covariate. Results The natural logarithm of the Jaccard index (lnJ) decreased significantly with increasing geographic distance for all vertebrate classes in each zoogeographic realm, and the slopes of the relationships per 1000 km ranged from ?0.251 to ?1.043. With environmental differences included in the analysis, both geographic distance and environmental differences were substantial predictors of lnJ for every combination of taxon and realm. On average, the unique contribution of geographic distance to variation in species turnover between ecoregions was about 1.4 times that of the environmental differences between ecoregions. Species turnover generally decreased with increasing latitude when controlling for geographic distance. The value of lnJ for each vertebrate class was highly and positively correlated with those of the other vertebrate classes. Main conclusions Our analyses suggest that both dispersal‐based and niche‐based processes have played important roles in determining faunal similarities among vertebrate assemblages at the spatial scale examined. Furthermore, reptiles and amphibians exhibited greater distance‐independent faunal heterogeneity among ecoregions and greater turnover among ecoregions with respect to geographic and environmental distance than birds and mammals.     

Life expectancy,maximum longevity and lifetime reproductive success in female Thornicroft's giraffe in Zambia

Gestation and longevity scale with body mass across taxa, yet within size dimorphic taxa, males tend to have reduced lifespans compared with females. Testing life history models, and accounting for sex differences in longevity, requires obtaining accurate longitudinal data from wild populations. We provide the first report describing key life history parameters from a long‐term study of giraffes in Africa. We followed a population of Thornicroft's giraffe (Giraffa camelopardalis thornicrofti) in Zambia for over 40 years. Maximum longevity among females was approximately 28 years, with lifespan accounting for 81% of the variance in lifetime reproductive success. Average adult female life expectancy was no different than average adult male life expectancy. However, the breeding lifespan of males was about half that of females, while maximum lifespan of males was 75% that of females. Our findings support the suggestion that sex differences in maximum lifespan arise from stronger selection for lengthy lives in females than in males. Among females, longer lives are associated with greater reproductive output.