Comparing environmental and genetic variance as adaptive response to fluctuating selection

Phenotypic variation within populations has two sources: genetic variation and environmental variation. Here, we investigate the coevolution of these two components under fluctuating selection. Our analysis is based on the lottery model in which genetic polymorphism can be maintained by negative frequency-dependent selection, whereas environmental variation can be favored due to bet-hedging. In our model, phenotypes are characterized by a quantitative trait under stabilizing selection with the optimal phenotype fluctuating in time. Genotypes are characterized by their phenotypic offspring distribution, which is assumed to be Gaussian with heritable variation for its mean and variance. Polymorphism in the mean corresponds to genetic variance while the width of the offspring distribution corresponds to environmental variance. We show that increased environmental variance is favored whenever fluctuations in the selective optima are sufficiently strong. Given the environmental variance has evolved to its optimum, genetic polymorphism can still emerge if the distribution of selective optima is sufficiently asymmetric or leptokurtic. Polymorphism evolves in a diagonal direction in trait space: one type becomes a canalized specialist for the more common ecological conditions and the other type a de-canalized bet-hedger thriving on the less-common conditions. All results are based on analytical approximations, complemented by individual-based simulations.     

Seed dormancy in Rumex species in response to environmental factors

Abstract. Many Rumex species show similar seed dormancy characteristics but there is more information concerning R. crispus and R. obtusifolius than other species. These species respond positively to red or white light. Far-red light applied for short periods may promote or inhibit germination depending on the timing of the irradiation in relation to temperature change; but long periods of far-red inhibit germination. Seeds may also be stimulated to germinate in the dark by low-temperature stratification at 15°C or less providing the temperature of the seeds is subsequently raised to a minimum of about 15°C. Seeds can, however, germinate at lower temperatures providing they have received other appropriate stimulatory treatment. Seeds also respond to alternating temperatures. In a diurnal cycle the minimum upper temperature required is about 15°C and the maximum lower temperature is about 25°C. The optimum period spent at the upper temperature is about 8 h when it is 15–25°C but the optimum period decreases as the upper temperature is increased above this range so that at 45°C, for example, it is only about 30 min. The period spent at the lower temperature in a diurnal cycle is not critical. Providing these criteria are met, the percentage germination increases with the number and amplitude of the cycles. The warming part of the cycle is necessary for the response but so far there is no convincing evidence that cooling itself is important. Secondary dormancy is induced at constant temperatures at a rate dependent on temperature, but apparently only in the presence of oxygen. This feature affects the optimum timing of a temperature change or exposure to light. Strong positive interactions are shown between stimulatory temperature treatments and white or red light. Unlike many other weed species the seeds respond only slightly to nitrate ions. The implications of these responses are discussed in relation to field behaviour.     

Combining environmental suitability and habitat connectivity to map rare or Data Deficient species in the Tropics

The IUCN Red List is a widely accepted system for classifying species’ risk of extinction, based on quantitative criteria. Although IUCN discourages the liberal use of the category “Data Deficient” (DD), most assessed groups have a large number of their species assigned to this category, especially in the Tropics. Therefore, DD species can introduce considerable uncertainty into estimates of proportions of threatened species, and research focused on elucidating the true status of those species should be a priority. Here we propose a simple method to gather information on geographic distribution and guide the search for new populations of rare, small-ranged, forest species, using the literature, online data, and standard GIS procedures. The method involves: (i) creating a geographic distribution model; (ii) selecting the environmentally suitable sites from that model; (iii) removing sites that have lost natural vegetation; and (iv) removing habitat networks that are too small and/or isolated, based on thresholds established from known occurrence records and the literature for ecologically similar species. As a case study, we use Lonchophylla peracchii, a recently described forest-dependent bat endemic to southeastern Brazil. We found that environmentally suitable sites for L. peracchii are already heavily deforested, confirming habitat loss as a major threat. Importantly, we identified five priority sites to search for the species outside of its currently known distribution. From that, we discuss its likely status based on IUCN's Criterion B2 (Extent of Occurrence). This method could be useful for other poorly known forest species, especially in the Tropics where most of these species are, and funding for research and fieldwork is scarcest. Currently there are 1910 terrestrial vertebrates in tropical forest worldwide classified as DD that could be evaluated using this method, provided that they have at least 5–10 occurrence records.     

Cranial postnatal growth of the rat relative to environmental variance

Neurocranial growth during the first 40 days of postnatal development was biometrically investigated relative to the specific parameters of calvarium length and width; basioccipital length and width; basisphenoid length; biparietal width; frontal bone length and cranial vault length. Specimens were obtained from rat litters subject to moderate nutritional suppression by raising the litter numbers above the norm via the cross-fostering technique and causing the overtaxing of the lactational capacity. Alterations in growth rate occurred throughout the pre- and postweaning periods with clearly demarcated phases of growth. Male and female growth in the preweaning period was uniform within each litter type but the phasic growth changes in the experimental litters differed from the control litters with regression line breakage emerging at different points and altering the phasic lengths. The preweaning phasic spectrum in both control and experimental litters was evidently linked to the maternal lactation capacity--relative to quantity and duration. Weaning was insidious in both litter types but occurred earlier in the experimental litters although the continuity of growth was maintained up to the end of the definitive preweaning period (day 20). The definitive postweaning period (day 21-40) showed considerable heterogeneity--unsatisfactory scatter--with differing regression line breakage points and phasic lengths. Analysis based on the total period (day 1-40) brought compensatory adjustments and indicated more acceptable phasic patterns in the postweaning period with confirmation of catch-up growth in the experimental litters. Allometric analysis of the parameters revealed varying rates of growth, line breakage points and differing phasic lengths indicating that each neurocranial parameter and the skeletal unit forming it had an individual characteristic response spectrum mostly attributable to the functional matrix with emphasis on the capsular rather than the periosteal.     

Current problems of environmental gradients and species response curves in relation to continuum theory

Abstract. Comparisons of the positions of species on Grimes'C-S-R triangular ordination model with their responses to individual environmental gradients indicates that the C-S-R model does not necessarily predict species ecological behaviour. The importance of the stress, productivity and disturbance gradients relative to other environmental gradients needs to be determined. In studies of species behaviour along a biomass/productivity gradient the collective vegetation property, biomass, has been confused with the environmental factor, fertility. Patterns of responses to biomass gradients e.g. Keddy's centrifugal model, should be examined in a two-dimensional environmental space to avoid such confounding effects. Assumptions regarding the shapes of species responses to environmental gradients remain untested. A recent model of species response functions to environmental gradients suggested that skewed responses curves show a pattern in the direction of the skew, always with the tail towards the presumed most mesic position on the gradient. Further evidence is presented to support this model for a temperature gradient in eucalypt forest in south-eastern Australia. 21 out of 24 species tested conform to the model.     

Benthic species of the Kerguelen Plateau show contrasting distribution shifts in response to environmental changes

Marine life of the Southern Ocean has been facing environmental changes and the direct impact of human activities during the past decades. Benthic communities have particularly been affected by such changes although we only slowly understand the effect of environmental changes on species physiology, biogeography, and distribution. Species distribution models (SDM) can help explore species geographic responses to main environmental changes. In this work, we modeled the distribution of four echinoid species with contrasting ecological niches. Models developed for [2005–2012] were projected to different time periods, and the magnitude of distribution range shifts was assessed for recent‐past conditions [1955–1974] and for the future, under scenario RCP 8.5 for [2050–2099]. Our results suggest that species distribution shifts are expected to be more important in a near future compared to the past. The geographic response of species may vary between poleward shift, latitudinal reduction, and local extinction. Species with broad ecological niches and not limited by biogeographic barriers would be the least affected by environmental changes, in contrast to endemic species, restricted to coastal areas, which are predicted to be more sensitive.     

揭示群落结构及其环境响应的联合物种分布模型的研究进展

物种分布模型通常用于基础生态和应用生态研究,用来确定影响生物分布和物种丰富度的因素,量化物种与非生物条件的关系,预测物种对土地利用和气候变化的反应,并确定潜在的保护区.在传统的物种分布模型中,生物的相互作用很少被纳入,而联合物种分布模型(JSDMs)作为近年提出的一种新的可行方法,可以同时考虑环境因素和生物交互作用,因而成为分析生物群落结构和种间相互作用过程的有力工具.JSDMs以物种分布模型(SDMs)为基础,通常采用广义线性回归模型建立物种对环境变量的多变量响应,以随机效应的形式获取物种间的关联,同时结合隐变量模型(LVMs),并基于Laplace近似和马尔科夫蒙脱卡罗模拟的最大似然估计或贝叶斯方法来估算模型参数.本文对JSDMs的产生及理论基础进行归纳总结,重点介绍了不同类型JSDMs的特点及其在现代生态学中的应用,阐述了JSDMs的应用前景、使用过程中存在的问题及发展方向.随着对环境因素与多物种种间关系研究的深入,JSDMs将是今后物种分布模型研究的重点.     

Predictor species: Improving assessments of rare species occurrence by modeling environmental co‐responses

Designing an effective conservation strategy requires understanding where rare species are located. Because rare species can be difficult to find, ecologists often identify other species called conservation surrogates that can help inform the distribution of rare species. Species distribution models typically rely on environmental data when predicting the occurrence of species, neglecting the effect of species' co‐occurrences and biotic interactions. Here, we present a new approach that uses Bayesian networks to improve predictions by modeling environmental co‐responses among species. For species from a European peat bog community, our approach consistently performs better than single‐species models and better than conventional multi‐species approaches that include the presence of nontarget species as additional independent variables in regression models. Our approach performs particularly well with rare species and when calibration data are limited. Furthermore, we identify a group of “predictor species” that are relatively common, insensitive to the presence of other species, and can be used to improve occurrence predictions of rare species. Predictor species are distinct from other categories of conservation surrogates such as umbrella or indicator species, which motivates focused data collection of predictor species to enhance conservation practices.     

Spatiotemporal variance of environmental conditions in Australian artesian springs affects the distribution and abundance of six endemic snail species

Artesian springs in arid Australia house endemic species with very small geographic distributions (most <50 km²). These species have limited dispersal capabilities, but little is known about environmental variance within and across these springs and how it, too, may limit their distribution and abundance. At the Pelican Creek springs complex, the full diversity of endemic gastropod fauna is found only in springs with deep pools, an area thought to provide greater environmental stability. This implies that the distributions of most snail species at this site may be restricted by their narrow environmental requirements and limits. This study monitored spatiotemporal environmental variance in a subset of the Pelican Creek springs (within Edgbaston Reserve) across one year to assess whether pool areas differ from tail areas, and how patterns of abundance of six snail species from three different families correspond to this variance. Springs fluctuated considerably in size, depth, water chemistry and temperature at daily and seasonal scales. Patterns of environmental variance differed across areas; pools were spatiotemporally stable, and tails were ephemeral and environmentally variable. The snail species occupied these areas in different ways. Species restricted to deep springs generally had significantly higher abundance in pool areas, and most had narrow environmental limits. In contrast, species found in a greater number of springs, including those with no pool, occupied pool and tail areas and generally had broader environmental limits. Environmental variance within and across springs affects the distribution of snails in a species-specific fashion. This has important implications for how we study springs and reveals that whilst the vast majority of species are restricted to areas of environmental stability, some can persist in the most environmentally variable areas.     

Sex ratio variance and the maintenance of environmental sex determination

Although variation in population sex ratios is predicted to increase the extinction rate of clades with environmental sex determination (ESD), ESD is still seen in a wide array of natural systems. It is unclear how this common sex-determining system has persisted despite this inherent disadvantage associated with ESD. We use simulation modelling to examine the effect of the sex ratio variance caused by ESD on population colonization and establishment. We find that an accelerating function of establishment success on initial population sex ratio favours a system that produces variance in sex ratios over one that consistently produces even sex ratios. This sex ratio variance causes ESD to be favoured over genetic sex determination, even when the mean global sex ratio under both sex-determining systems is the same. Data from ESD populations suggest that the increase in population establishment can more than offset the increased risk of extinction associated with temporal fluctuations in the sex ratio. These findings demonstrate that selection in natural systems can favour increased variance in a trait, irrespective of the mean trait value. Our results indicate that sex ratio variation may provide an advantage to species with ESD, and may help explain the widespread existence of this sex-determining system.     

BioMove – an integrated platform simulating the dynamic response of species to environmental change

BioMove simulates plant species' geographic range shifts in response to climate, habitat structure and disturbance, at annual time steps. This spatially explicit approach integrates species' bioclimatic suitability and population‐level demographic rates with simulation of landscape‐level processes (dispersal, disturbance, species' response to dynamic dominant vegetation structure). Species population dynamics are simulated through matrix modelling that includes scaling demographic rates by climatic suitability. Dispersal functions simulate population spread. User‐specified plant functional types (PFTs) provide vegetation structure that determines resource competition and disturbance. PFTs respond annually through dispersal, inter‐PFT competition and demographic shifts. BioMove provides a rich framework for dynamic range simulations.     

干热河谷草本植物生物量分配及其对环境因子的响应

以干热河谷6种草本植物为对象,研究了水分、养分、刈割对生物量在根、茎、叶的分配及异速生长关系的影响.结果表明:刈割处理叶生物量质量分数从25.1%显著增加到31.2%,茎生物量质量分数从43.7%显著降低到34.2%;养分添加处理根生物量质量分数从34.0%显著降低到30.8%;水分处理对生物量分配没有显著影响.物种对根、茎、叶生物量分配有显著影响,适应贫瘠土壤的物种将更多生物量分配给叶和根,对茎生物量的分配相对较低.物种与环境因子存在显著的互作效应,表明环境因子对不同物种的生物量分配影响不同.适应贫瘠土壤的物种叶-茎标度指数和异速生长常数大于其他物种,而茎-根标度指数和异速生长常数小于其他物种.养分显著增加了叶-茎和叶-根的异速生长常数,刈割显著降低了茎-根的标度指数,水分处理则没有显著效应.环境因素对器官间异速生长关系的影响存在种间差异.生物量分配的种间差异及其对环境因素的响应特征可能对植物适应环境变化产生重要影响.     

外来入侵植物对环境梯度和天敌逃逸的适应进化

进化假说认为, 在入侵地外来种能发生遗传变化, 以适应新的环境, 最终成功定殖和扩散。种内或种间杂交、遗传漂变、新环境带来的新的选择压力等是促使外来种发生进化的重要原因。在入侵地, 响应来自非生物和生物因素的选择压力是外来种发生适应进化的主要原因。本文主要介绍外来植物如何通过进化适应入侵地的纬度、海拔等非生物环境和天敌逃逸等生物环境。关于外来入侵植物对纬度和海拔等环境梯度的适应进化, 本文在强调表型进化研究应与分子标记研究相结合的基础上, 介绍了一些同质种植园实验和交互移植实验。关于外来入侵植物对天敌逃逸的适应进化, 本文主要介绍增强竞争能力的进化假说和修正的增强竞争能力的进化假说, 及其在理论上和验证方法上存在的问题。最后, 本文介绍了氮分配的进化假说, 该假说认为天敌逃逸可使外来入侵植物降低叶氮向防御的分配, 同时增加氮向光合的分配。     

甘南亚高寒草甸稀有种对物种多样性和物种多度分布格局的贡献

稀有种不仅影响群落的物种多度分布格局, 同时也是α多样性的重要贡献者。本研究主要通过加性分配和Fortran软件的RAD程序包拟合的方法, 研究了甘南亚高寒草甸不同坡向物种多样性及多度分布格局的变化, 分析了物种多度分布格局及其α多样性的变化特征, 确定了稀有种在物种多度分布格局中的相对贡献。结果表明: (1)在南坡到北坡的变化中, 环境因子差异比较明显, 其中, 土壤全磷、有机碳、速效磷、碳氮比及含水量呈递增趋势; 土壤氮磷比和pH值呈递减趋势; 土壤全氮在西坡显著低于其他坡向, 而速效氮在所有坡向上差异不显著。(2)稀有种对群落物种多样性的影响在南-北坡向梯度上依次增大, 去除稀有种的影响在各坡向均高于去除非稀有种, 可见, 稀有种在甘南亚高寒草甸物种多样性中的相对贡献高于非稀有种。(3)各坡向的稀有种资源获取模式以随机分配占领模式(random fraction模型)为主, 而非稀有种则以生态位优先占领模式(geometric series模型)为主。由于稀有种有较大的扩散率, 在物种多样性较高的生态系统中, 物种之间的生态位重叠会更加明显, 从而抑制物种多样性的增加, 因此能达到维持原有物种多样性的目的。     

The response of fish size and species diversity to environmental gradients in two Neotropical coastal streams

Hydrobiologia - This study investigated the effects of environmental variation on fish diversity patterns in two coastal streams in northwestern Ecuador. Specifically, we examined the role of...     

Within and between species variation in response to environmental gradients in Polygonum pensylvanicum and Polygonum virginianum

Summary The responses of Polygonum pensylvanicum L., an early successional annual, and of Polygonum virginianum L., a late successional perennial, were examined along discontinuous gradients of soil moisture, light and nutrient availability. In the case of P. virginianum both individuals grown from seed and individuals grown from rhizomes were examined. The results show that variation in the response of individuals of a species of different age to environmental variation is as great as that found between the two congeneric species of different successional habitats. In general, individuals of the two species were more similar to one another in response to the resource gradients when both were started from seed, than were individuals of P. virginianum grown from seed and from rhizomes. Potential differences in stored reserves (starting capital) between rhizomes and seeds appeared to have little effect upon responses to resource availability. A number of plant characters were found to vary along the gradients in ways that were unique to the character, the gradient, and the age of the individual. These characters included aspects of leaf size, shape, and orientation, as well as whole plant architecture. Many if not all of these characters are likely to have significant effects upon the functioning of plants. The origin of this difference in response to the gradients of individuals of P. virginianum of differing age may be ontogenetic or may reflect differences in genetic composition created by recombination.     

Near-periodic substitution and the genetic variance induced by environmental change

We investigate a model that describes the evolution of a diploid sexual population in a changing environment. Individuals have discrete generations and are subject to selection on the phenotypic value of a quantitative trait, which is controlled by a finite number of bialleic loci. Environmental change is taken to lead to a uniformly changing optimal phenotypic value. The population continually adapts to the changing environment, by allelic substitution, at the loci controlling the trait. We investigate the detailed interrelation between the process of allelic substitution and the adaptation and variation of the population, via infinite population calculations and finite population simulations. We find a simple relation between the substitution rate and the rate of change of the optimal phenotypic value.