Ecological and geographic modes of species divergence in wild tomatoes

Understanding the role of geography and ecology in species divergence is central to the study of evolutionary diversification. We used climatic, geographic, and biological data from nine wild Andean tomato species to describe each species' ecological niche and to evaluate the likely ecological and geographical modes of speciation in this clade. Using data from >1000 wild accessions and publicly available data derived from geographic information systems for various environmental variables, we found most species pairs were significantly differentiated for one or more environmental variables. By comparing species' predicted niches generated by species distribution modeling (SDM), we found significant niche differentiation among three of four sister-species pairs, suggesting ecological divergence is consistently associated with recent divergence. In comparison, based on age-range correlation (ARC) analysis, there was no evidence for a predominant geographical (allopatric vs. sympatric) context for speciation in this group. Overall, our results suggest an important role for environmentally mediated differentiation, rather than simply geographical isolation, in species divergence.     

Use of direct gradient analysis to study the climate-vegetation relationships in Galicia,Spain

This paper reports a bioclimatic analysis of plant species in Galicia, NW Spain. A set of floristic data obtained from 150 plots located at euclimatopes (sites with monitored climate) was analysed using direct gradient analysis and clustering with respect to the 8 climatic variables thought to play a major role in regulating the distribution of the species considered in the study area. Principal component analysis (PCA) and hierarchical clustering were based on a matrix of species by climatic variables. Indicator taxa for the variables were identified on the basis of their Indicator values (Brisse & Grandjouan 1978) and grouped by cluster analysis. The groups produced were compatible with the results of principal component analysis and the frequential analysis of the species, which identified their phytoclimatic nature. The groups were then characterized by determining their climatic positions and indicator values with respect to the chief climatic variables. The first three PCA axes, which were associated with Oceanity, Mean minimum temperatures and the temperature range in the coldest month, together accounted for 97.2% of the variance of the data.     

Evaluating the ecological realism of plant species distribution models with ecological indicator values

Species distribution models (SDMs) are routinely applied to assess current as well as future species distributions, for example to assess impacts of future environmental change on biodiversity or to underpin conservation planning. It has been repeatedly emphasized that SDMs should be evaluated based not only on their goodness of fit to the data, but also on the realism of the modeled ecological responses. However, possibilities for the latter are hampered by limited knowledge on the true responses as well as a lack of quantitative evaluation methods. Here we compared modeled niche optima obtained from European-scale SDMs of 1476 terrestrial vascular plant species with empirical ecological indicator values indicating the preferences of plant species for key environmental conditions. For each plant species we first fitted an ensemble SDM including three modeling techniques (GLM, GAM and BRT) and extracted niche optima for climate, soil, land use and nitrogen deposition variables with a large explanatory power for the occurrence of that species. We then compared these SDM-derived niche optima with the ecological indicator values by means of bivariate correlation analysis. We found weak to moderate correlations in the expected direction between the SDM-derived niche optima and ecological indicator values. The strongest correlation occurred between the modeled optima for growing degree days and the ecological indicator values for temperature. Correlations were weaker for SDM-derived niche optima with a more distal relationship to ecological indicator values (notably precipitation and soil moisture). Further, correlations were consistently highest for BRT, followed by GLM and GAM. Our method gives insight into the ecological realism of modeled niche optima and projected core habitats and can be used to improve SDMs by making a more informed selection of environmental variables and modeling techniques.     

Tracking of climatic niche boundaries under recent climate change

1.?Global climate has changed significantly during the past 30?years and especially in northern temperate regions which have experienced poleward shifts in temperature regimes. While there is evidence that some species have responded by moving their distributions to higher latitudes, the efficiency of this response in tracking species' climatic niche boundaries over time has yet to be addressed. 2.?Here, we provide a continental assessment of the temporal structure of species responses to recent spatial shifts in climatic conditions. We examined geographic associations with minimum winter temperature for 59 species of winter avifauna at 476 Christmas Bird Count circles in North America from 1975 to 2009 under three sampling schemes that account for spatial and temporal sampling effects. 3.?Minimum winter temperature associated with species occurrences showed an overall increase with a weakening trend after 1998. Species displayed highly variable responses that, on average and across sampling schemes, contained a strong lag effect that weakened in strength over time. In general, the conservation of minimum winter temperature was relevant when all species were considered together but only after an initial lag period (c. 35?years) was overcome. The delayed niche tracking observed at the combined species level was likely supported by the post1998 lull in the warming trend. 4.?There are limited geographic and ecological explanations for the observed variability, suggesting that the efficiency of species' responses under climate change is likely to be highly idiosyncratic and difficult to predict. This outcome is likely to be even more pronounced and time lags more persistent for less vagile taxa, particularly during the periods of consistent or accelerating warming. Current modelling efforts and conservation strategies need to better appreciate the variation, strength and duration of lag effects and their association with climatic variability. Conservation strategies in particular will benefit through identifying and maintaining dispersal corridors that accommodate diverging dispersal strategies and timetables.     

Environmental data sets matter in ecological niche modelling: an example with Solenopsis invicta and Solenopsis richteri

Aim  In response to a recent paper suggesting the failure of ecological niche models to predict between native and introduced distributional areas of fire ants ( Solenopsis invicta ), we sought to assess methodological causes of this failure.
Location  Ecological niche models were developed on the species' native distributional area in South America, and projected globally.
Methods  We developed ecological niche models based on six different environmental data sets, and compared their respective abilities to anticipate the North American invasive distributional area of the species.
Results  We show that models based on the 'bioclimatic variables' of the WorldClim data set indeed fail to predict the full invasive potential of the species, but that models based on four other data sets could predict this potential correctly.
Main conclusions  The difference in predictive abilities appears to centre on the complexity of the environmental variables involved. These results emphasize important influences of environmental data sets on the generality and ability of ecological niche models to anticipate novel phenomena, and offer a simpler explanation for the lack of predictive ability among native and invaded distributional areas than that of niche shifts.     

Weighted averaging,logistic regression and the Gaussian response model

The indicator value and ecological amplitude of a species with respect to a quantitative environmental variable can be estimated from data on species occurrence and environment. A simple weighted averaging (WA) method for estimating these parameters is compared by simulation with the more elaborate method of Gaussian logistic regression (GLR), a form of the generalized linear model which fits a Gaussian-like species response curve to presence-absence data. The indicator value and the ecological amplitude are expressed by two parameters of this curve, termed the optimum and the tolerance, respectively. When a species is rare and has a narrow ecological amplitude — or when the distribution of quadrats along the environmental variable is reasonably even over the species' range, and the number of quadrats is small — then WA is shown to approach GLR in efficiency. Otherwise WA may give misleading results. GLR is therefore preferred as a practical method for summarizing species' distributions along environmental gradients. Formulas are given to calculate species optima and tolerances (with their standard errors), and a confidence interval for the optimum from the GLR output of standard statistical packages.Nomenclature follows Heukels-van der Meijden (1983).We would like to thank Drs I. C. Prentice, N. J. M. Gremmen and J. A. Hoekstra for comments on the paper. We are grateful to Ir. Th. A. de Boer (CABO, Wageningen) for permission to use the data of the first example.     

全国栽培太子参生态适宜性区划分析

采用Arc GIS地理信息系统软件的空间分析方法和SPSS统计软件的统计分析方法,分析了全国太子参潜在适宜种植地理分布、适宜生境及化学成分与环境因子的相关性。结果显示:采用聚类分析从46个样地中筛选出34个多糖含量较高的样地。所建模型经ROC曲线验证,预测效果非常好(AUC0.9)。最干月降水量(贡献率68.4%)、土壤类型(9.7%)、最暖季平均温(6.0%)、等温性(4.0%)是影响太子参种植分布最重要的环境因子,其最适宜生境范围为:最干月降水量20—60mm;最暖季平均温21.5—30.5℃;等温性18%—35%;土壤类型为不饱和薄层土、饱和粘磬土中的黄棕壤、黄壤、黄红壤。相关性分析显示,多糖和生境适宜度均与最湿季降水量、最暖季降水量、最暖季平均温、最干月降水量及年平均气温无显著相关性。太子参最适宜区主要分布在长江中下游区域,主要有贵州中部,重庆与湖南、湖北接壤处,河南南部,安徽西部,江苏中部,福建东北部以及浙江北部和东南部区域。因此,明确全国太子参潜在的适宜种植区可为太子参种植基地的选取和合理布局提供参考。     

How do ecologists select and use indicator species to monitor ecological change? Insights from 14 years of publication in Ecological Indicators

Indicator species (IS) are used to monitor environmental changes, assess the efficacy of management, and provide warning signals for impending ecological shifts. Though widely adopted in recent years by ecologists, conservation biologists, and environmental practitioners, the use of IS has been criticized for several reasons, notably the lack of justification behind the choice of any given indicator. In this review, we assess how ecologists have selected, used, and evaluated the performance of the indicator species. We reviewed all articles published in Ecological Indicators (EI) between January 2001 and December 2014, focusing on the number of indicators used (one or more); common taxa employed; terminology, application, and rationale behind selection criteria; and performance assessment methods. Over the last 14 years, 1914 scientific papers were published in EI, describing studies conducted in 53 countries on six continents; of these, 817 (43%) used biological organisms as indicators. Terms used to describe organisms in IS research included “ecological index”, “environmental index”, “indicator species”, “bioindicator”, and “biomonitor,” but these and other terms often were not clearly defined. Twenty percent of IS publications used only a single species as an indicator; the remainder used groups of species as indicators. Nearly 50% of the taxa used as indicators were animals, 70% of which were invertebrates. The most common applications behind the use of IS were to: monitor ecosystem or environmental health and integrity (42%); assess habitat restoration (18%); and assess effects of pollution and contamination (18%). Indicators were chosen most frequently based on previously cited research (40%), local abundance (5%), ecological significance and/or conservation status (13%), or a combination of two or more of these reasons (25%). Surprisingly, 17% of the reviewed papers cited no clear justification for their choice of indicator. The vast majority (99%) of publications used statistical methods to assess the performance of the selected indicators. This review not only improves our understanding of the current uses and applications of IS, but will also inform practitioners about how to better select and evaluate ecological indicators when conducting future IS research.     

Climatic mean diagrams: A technique for assessing the climatic differentiation of species

 A new method for comparisons of the ecogeographical and climatic constitutions of taxa is presented. The method will be important for understanding the evolution of species in time and space because ecogeographical characters of the species become directly comparable. On a global analytical level, distribution ranges of species may be considered as a function of the ecological constitution of the species, i.e. their endogenous hardiness to environmental forces, and climate. Frequency diagrams of the species' occurrences are calculated for the monthly climatic means of temperature and precipitation using high-resolution maps of distribution and climate. These frequency diagrams are used to construct monthly temperature-precipitation diagrams (TPD) that show the monthly climate spaces of the species'. For the first time, within the TPDs the climate spaces of the plants become directly comparable. However, comparability is somewhat restricted because of the necessity to consider the TPDs of all 12 months. A higher degree of abstraction is obtained using the climatic centre of a species as inferred from the TPD. The twelve monthly climatic means are transferred to the climatic mean diagram (CMD). In the CMD the climate spaces of the species become directly comparable in the course of the year. Despite the high degree of abstraction each diagram type reflects particular ecogeographical characteristics of the species even at a regional geographical level. Especially the CMD offers ways for understanding evolutionary shifts in the ecogeographical constitution of closely related species. The reasons for the shifts are composed of a hydric and thermal component. They may be addressed in molecular and physiological studies concerning evolutionary changes of these ecogeographical traits. Received August 7, 2000 Accepted November 5, 2001     

Experimental test for adaptive differentiation of ginseng populations reveals complex response to temperature

Background and Aims Local climatic adaptation can influence species' response to climate change. If populations within a species are adapted to local climate, directional change away from mean climatic conditions may negatively affect fitness of populations throughout the species' range. Methods Adaptive differentiation to temperature was tested for in American ginseng (Panax quinquefolius) by reciprocally transplanting individuals from two populations, originating at different elevations, among temperature treatments in a controlled growth chamber environment. Fitness-related traits were measured in order to test for a population × temperature treatment interaction, and key physiological and phenological traits were measured to explain population differences in response to temperature. Key Results Response to temperature treatments differed between populations, suggesting genetic differentiation of populations. However, the pattern of response of fitness-related variables generally did not suggest 'home temperature' advantage, as would be expected if populations were locally adapted to temperature alone. Conclusions Failure consistently to detect a 'home temperature' advantage response suggests that adaptation to temperature is complex, and environmental and biotic factors that naturally covary with temperature in the field may be critical to understanding the nature of adaptation to temperature.     

Testing a biological mechanism of the insurance hypothesis in experimental aquatic communities

1.  The insurance hypothesis predicts a stabilizing effect of increasing species richness on community and ecosystem properties. Difference among species' responses to environmental fluctuations provides a general mechanism for the hypothesis. Previous experimental investigations of the insurance hypothesis have not examined this mechanism directly.
2.  First, responses to temperature of four protist species were measured in laboratory microcosms. For each species, we measured the response of intrinsic rate of increase ( r ) and carrying capacity ( K ) to temperature.
3.  Next, communities containing pairs of species were exposed to temperature fluctuations. Community biomass varied less when correlation in K between species (but not r ) was more negative, and this resulted from more negative covariances in population sizes, as predicted. Results were contingent on species identity, with findings differing between analyses including or not including communities containing one particular species.
4.  These findings provide the clearest support to date for this mechanism of the insurance hypothesis. Biodiversity, in terms of differences in species' responses to environmental fluctuations (i.e. functional response diversity) stabilizes community dynamics.     

Relating habitat and climatic niches in birds

Predicting species' responses to the combined effects of habitat and climate changes has become a major challenge in ecology and conservation biology. However, the effects of climatic and habitat gradients on species distributions have generally been considered separately. Here, we explore the relationships between the habitat and thermal dimensions of the ecological niche in European common birds. Using data from the French Breeding Bird Survey, a large-scale bird monitoring program, we correlated the habitat and thermal positions and breadths of 74 bird species, controlling for life history traits and phylogeny. We found that cold climate species tend to have niche positions in closed habitats, as expected by the conjunction of the biogeographic history of birds' habitats, and their current continent-scale gradients. We also report a positive correlation between thermal and habitat niche breadths, a pattern consistent with macroecological predictions concerning the processes shaping species' distributions. Our results suggest that the relationships between the climatic and habitat components of the niche have to be taken into account to understand and predict changes in species' distributions.     

Humpback whale diets respond to variance in ocean climate and ecosystem conditions in the California Current

Large, migratory predators are often cited as sentinel species for ecosystem processes and climate‐related changes, but their utility as indicators is dependent upon an understanding of their response to environmental variability. Documentation of the links between climate variability, ecosystem change and predator dynamics is absent for most top predators. Identifying species that may be useful indicators and elucidating these mechanistic links provides insight into current ecological dynamics and may inform predictions of future ecosystem responses to climatic change. We examine humpback whale response to environmental variability through stable isotope analysis of diet over a dynamic 20‐year period (1993–2012) in the California Current System (CCS). Humpback whale diets captured two major shifts in oceanographic and ecological conditions in the CCS. Isotopic signatures reflect a diet dominated by krill during periods characterized by positive phases of the North Pacific Gyre Oscillation (NPGO), cool sea surface temperature (SST), strong upwelling and high krill biomass. In contrast, humpback whale diets are dominated by schooling fish when the NPGO is negative, SST is warmer, seasonal upwelling is delayed and anchovy and sardine populations display increased biomass and range expansion. These findings demonstrate that humpback whales trophically respond to ecosystem shifts, and as a result, their foraging behavior is a synoptic indicator of oceanographic and ecological conditions across the CCS. Multi‐decadal examination of these sentinel species thus provides insight into biological consequences of interannual climate fluctuations, fundamental to advancing ecosystem predictions related to global climate change.     

里海西北部Caspian地区草原扩大与啮齿类群落结构的变化

于1997-1999年检测了俄联邦卡尔梅克(Kalmyki)共和国ChernieZemly半干旱草原由10个物种组成的啮齿类群落。该区域的植被特点是家畜大量减少后形成的恢复植被。我们的任务之一就是了解不同啮齿动物的生境需求,以预测不同物种对环境变化的特异性反应。我们也检测了群落空间结构、生态位参数和物种多样性。逐步回归分析表明,各种啮齿动物的空间分布仅部分地决定于已知环境因子(2-6个变量)。虽然方程式具有强显著性,但决定系数R2很低,不超过18%。对于大多数种类,这种结果可由最近草原扩大而来的生境异质性降低解释。物种生境选择条件和资源的明显低水平可能是由物种对环境变化的惯性反应造成的。判别函数分析和主分量分析结果表明,群落结构以及单个物种的生态位参数在不同年间并不稳定。空间分层结构在种间中等和低重叠的高物种多样性年度表现明显。多样性、生态位组成和生态位宽度分析结果表明,不同物种对环境条件和资源变化的反应具有不同的个性化方式。同时,物种对环境动态的明显个性化反应成为物种多样性定向变化的原因。限定空间和相对单一条件下的结果表明,啮齿类α多样性与生境结构复杂性呈正相关,而与生境生产力特征呈负相关。观察表明,卡尔梅克国草原扩大伴随着初级生产力的升高和生境异质性的降低。随着时间的推移会导致大多数草原和半干旱区域啮齿类多样性的降低,并形成动物区系核心.     

Epigeic bryophytes do not improve bioindication by Ellenberg values in mountain forests

Based on a stratified random sample of 93 vegetation plots and coincident measurements of ecological conditions in mountain forests of the Bavarian Alps, the degree to which species composition and Ellenberg indicator values derived thereof were related to measured environmental variables was assessed for vascular understorey plants and epigeic bryophytes. According to Mantel tests vascular composition contained ca. 30% more ecological information than bryophyte composition. When expressed as average Ellenberg or Düll values, vascular plant-based values reflected 60% more of measured variables than bryophyte-based values. The differences remained after rarefaction of the vascular matrix to the gamma diversity of bryophytes, showing that indication is not a function of indicator richness. Analysing vascular plants and bryophytes combined yielded very similar, or even slightly less stringent relationships with the environment than using vascular plants only.Bivariate relationships of indicator values with corresponding ecological measurements confirmed the specific potential of the values to estimate ecological factors from both plant groups, but vascular plants performed better for all factors. Bryophyte indication was particularly poor for light, temperature and base saturation. Bryophyte-based indicator values did not significantly predict the residuals of measured ecological variables against vascular plant-based Ellenberg values.For the study region, it is concluded that indicator values of vascular forest understorey should be used without consideration of Düll's indicator values for epigeic bryopyhtes. There appears to be potential to improve bioindication by recalibrating indicator values of epigeic bryophytes based on ecological measurements and vascular plant indicator values.     

气候变化对杉木适生区和生态位的影响

杉木（Cunninghamia lanceolata）是我国主要的造林树种之一,具有重要的药用、经济和生态服务功能。在全球变暖趋势持续的背景下,气候成为制约物种生存和发展的重要因素,其中空间分布和生态位变化对生态过程的解释至关重要。基于211个杉木分布点和20个环境变量,利用MaxEnt模型和R语言的ecospat软件包对杉木适生区和生态位的变化进行研究,分析杉木对未来气候持续变暖的响应。结果表明杉木当前潜在适生区219.67万km²,约占国土总面积的22.88％,主要分布在我国800 mm等降水量线以东的地区,年均温、昼夜温差日均值和最干月降雨量是影响杉木分布的最主要环境变量。未来杉木适生区分布将沿着经纬度方向迁移,高度和低度适生区面积均不断减少。主成分分析（PCA）表明杉木气候生态位在不同时段不同代表性浓度路径下发生转移和扩展,气候生态位中心将向年均温和最暖季降雨量移动。生态位重叠指数均呈现不断下降的趋势,且RCP8.5情景下的生态位重叠率下降最为明显。结果表明全球气候变暖会改变物种的空间分布区域,并将对现存的生态系统产生不同程度的影响。杉木与气候变化关系的研究拓展了人们对气候变化与植物物种生态特征的认识,为杉木乃至乔木树种的保护和利用提供了理论依据。     

Breathing some air into the single-species vacuum: multi-species responses to environmental change

Studies of ecological responses to climate change have often analysed species independently of each other, yet interactions between species are fundamental aspects of ecology. Mutshinda, O'Hara & Woiwod (2011) used light-trapping data for Lepidoptera (moths) to examine population responses to intraspecific effects and effects of winter rainfall and temperature. They show how Bayesian hierarchical models can analyse residual correlations among species' responses, illustrating an approach to account for and measure dependencies that are not fully explained by the candidate explanatory variables. A key result is that the responses of the different moth species did not appear to have strong residual correlation (Mutshinda, O'Hara & Woiwod 2011). These analyses provide an approach for synthesising across species and can better inform ecological responses to environmental change.     

The ‘abundant centre’ distribution: to what extent is it a biogeographical rule?

Several ecological and evolutionary hypotheses are based on the assumption that species reach their highest abundance in the centre of their range and decline in abundance toward the range edges. We reviewed empirical tests of this assumption, which we call the 'abundant centre' hypothesis. We found that of 145 separate tests conducted as part of 22 direct empirical studies, only 56 (39%) support the abundant centre hypothesis. More problematic than the percentage of studies that support the hypothesis is the finding that most studies inadequately sampled the species' ranges. Only two of the studies analysed data that were collected throughout the species' range. The remaining studies relied on data from a small number of points in their analysis, meaning that the range edges were severely under-sampled. Patterns of abundance across the entire range must be known to draw testable hypotheses about the consequences of species' geographical abundance distributions. Indirect tests of the abundant centre hypothesis, in which ecological or evolutionary expectations of abundant centre distributions were examined, did not support or reject the abundant centre hypothesis overall. We conclude that more exploration of species' abundance distributions is necessary and we suggest methods to use in future studies.     

Lutzomyia vectors for cutaneous leishmaniasis in Southern Brazil: ecological niche models,predicted geographic distributions,and climate change effects

Geographic and ecological distributions of three Lutzomyia sand flies that are cutaneous leishmaniasis vectors in South America were analysed using ecological niche modelling. This new tool provides a large-scale perspective on species' geographic distributions, ecological and historical factors determining them, and their potential for change with expected environmental changes. As a first step, the ability of this technique to predict geographic distributions of the three species was tested statistically using two subsampling techniques: a random-selection technique that simulates 50% data density, and a quadrant-based technique that challenges the method to predict into broad unsampled regions. Predictivity under both test schemes was highly statistically significant. Visualisation of ecological niches provided insights into the ecological basis for distributional differences among species. Projections of potential geographic distributions across scenarios of global climate change suggested that only Lutzomyia whitmani is likely to be experiencing dramatic improvements in conditions in south-eastern Brazil, where cutaneous leishmaniasis appears to be re-emerging; Lutzomyia intermedia and Lutzomyia migonei may be seeing more subtle improvements in climatic conditions, but the implications are not straightforward. More generally, this technique offers the possibility of new views into the distributional ecology of disease, vector, and reservoir species.     

植物种分布的模拟研究进展

从植物种水平研究植被与气候的关系一直是生态学的热点之一。该文综述了植物种与气候关系的早期研究历史和国内外近期研究进展,尤其是20世纪80年代以来,随着全球变化研究的不断发展和深入,植物种地理分布与气候因子关系研究的最新发展,汇总了最近20年来国际上模拟预测植物种潜在地理分布的模型,比较了不同模型的优缺点。统计模型主要包括以生物气候分室模型或气候分室模型为代表的相关模型、以广义线性模型和广义加性模型为代表的回归模型、以分类和回归树分析及人工神经网络为代表的基于规则的模型、以及生态位模型、气候响应面模型等。机理模型主要介绍了基于BIOME1生物地理模型和FORSKA林窗模型的STASH模型、基于过程的物候模型PHENOFIT,以及一种基于水分平衡、温度和植物物候现象的模型。总结不同模型模拟预测的不同地区植物种未来分布的格局,并介绍中国植物种潜在分布区及未来变化的模拟预测工作,从而为更加准确地模拟预测植物种在未来全球变化情景下的变化趋势提供背景知识。