Environmental variability and acoustic signals: a multi-level approach in songbirds

Among songbirds, growing evidence suggests that acoustic adaptation of song traits occurs in response to habitat features. Despite extensive study, most research supporting acoustic adaptation has only considered acoustic traits averaged for species or populations, overlooking intraindividual variation of song traits, which may facilitate effective communication in heterogeneous and variable environments. Fewer studies have explicitly incorporated sexual selection, which, if strong, may favour variation across environments. Here, we evaluate the prevalence of acoustic adaptation among 44 species of songbirds by determining how environmental variability and sexual selection intensity are associated with song variability (intraindividual and intraspecific) and short-term song complexity. We show that variability in precipitation can explain short-term song complexity among taxonomically diverse songbirds, and that precipitation seasonality and the intensity of sexual selection are related to intraindividual song variation. Our results link song complexity to environmental variability, something previously found for mockingbirds (Family Mimidae). Perhaps more importantly, our results illustrate that individual variation in song traits may be shaped by both environmental variability and strength of sexual selection.     

The relationship between species richness and ecosystem variability is shaped by the mechanism of coexistence

Theory relating species richness to ecosystem variability typically ignores the potential for environmental variability to promote species coexistence. Failure to account for fluctuation‐dependent coexistence may explain deviations from the expected negative diversity–ecosystem variability relationship, and limits our ability to predict the consequences of increases in environmental variability. We use a consumer‐resource model to explore how coexistence via the temporal storage effect and relative nonlinearity affects ecosystem variability. We show that a positive, rather than negative, diversity–ecosystem variability relationship is possible when ecosystem function is sampled across a natural gradient in environmental variability and diversity. We also show how fluctuation‐dependent coexistence can buffer ecosystem functioning against increasing environmental variability by promoting species richness and portfolio effects. Our work provides a general explanation for variation in observed diversity–ecosystem variability relationships and highlights the importance of conserving regional species pools to help buffer ecosystems against predicted increases in environmental variability.     

Pathogen impacts on plant diversity in variable environments

Environmental variability can promote species diversity when species respond differently to environmental conditions, via the storage effect. Pathogens, predators and other shared consumers can also facilitate coexistence when they differentially limit common species. However, it remains unclear how environmental variation and consumers interact to determine species diversity. Here, I use a model based on California annual grassland plants to show that a generalist pathogen with no host specificity can enhance the positive effect of environmental variability on diversity. The model predicts that pathogens can promote diversity by increasing the covariance between the environment and competition, enhancing the storage effect. However, pathogen impacts depend on life history. Pathogens that infect germinating seeds or plants tend to increase the storage effect, whereas those that infect dormant seeds can undermine the storage effect by eliminating population buffering during unfavorable years. These results suggest that pathogens may mediate plant responses to environmental variability and change, and in doing so may maintain diversity.     

Do Epigeal Termite Mounds Increase the Diversity of Plant Habitats in a Tropical Rain Forest in Peninsular Malaysia?

The extent to which environmental heterogeneity can account for tree species coexistence in diverse ecosystems, such as tropical rainforests, is hotly debated, although the importance of spatial variability in contributing to species co-existence is well recognized. Termites contribute to the micro-topographical and nutrient spatial heterogeneity of tropical forests. We therefore investigated whether epigeal termite mounds could contribute to the coexistence of plant species within a 50 ha plot at Pasoh Forest Reserve, Malaysia. Overall, stem density was significantly higher on mounds than in their immediate surroundings, but tree species diversity was significantly lower. Canonical correspondence analysis showed that location on or off mounds significantly influenced species distribution when stems were characterized by basal area. Like studies of termite mounds in other ecosystems, our results suggest that epigeal termite mounds provide a specific microhabitat for the enhanced growth and survival of certain species in these species-rich tropical forests. However, the extent to which epigeal termite mounds facilitate species coexistence warrants further investigation.     

Intraspecific and interspecific trait variability in tadpole meta‐communities from the Brazilian Atlantic rainforest

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Variations in species and functional plant diversity along climatic and grazing gradients

Different components of biodiversity may vary independently of each other along environmental gradients giving insights into the mechanisms that regulate species coexistence. In particular, the functional diversity (FD) or the presence of rare or endemic species in natural assemblages do not necessarily increase with species diversity. We studied if different components of plant species diversity (species richness, Simpson diversity, evenness) varied similarly to FD (measured as a generalization of the Simpson index) and rarity along grazing intensity and climatic gradients. Plots under different sheep grazing regimes (high and low intensity, abandonment) were surveyed in five locations along a climatic gradient in north-eastern Spain, from semi-arid lowland to moist upland locations. Variation in species diversity, functional diversity and rarity followed different patterns. Species diversity was lowest in water-stressed environments (arid locations and southern aspects) and increased with grazing more makedly in humid locations. The FD was comparable between the most species-poor and species-rich locations and decreased with grazing in the moistest location, i.e. where species diversity markedly increased. The FD did not show a strong correlation with species richness nor with the Simpson index and less specious communities could show the highest functional diversity. The rarest species in the region were more frequently found in the abandoned areas, which held the lowest species diversity. Consequently, the mechanisms that enhance the diversity of species do not necessarily support a functional differentiation among those species or the maintenance of rare species in a region. We hypothesize that the degree of dependence of functional diversity on species diversity might be mostly related to the amplitude of the species' traits pool and on how species partition the niche space available.     

Disparate relatives: Life histories vary more in genera occupying intermediate environments

Species within clades are commonly assumed to share similar life history traits, but within a given region some clades show much greater variability in traits than others. Are variable clades older, allowing more time for trait diversification? Or do they occupy particular environments, providing a wider range of abiotic or biotic opportunities for the establishment and maintenance of diverse trait attributes? Does environmental opportunity increase trait variability across all species, or is it specific to species belonging to the same clade, increasing only within-clade trait variability? We studied the variability of six life-history traits (initiation of flowering, duration of flowering, plant life span, seed mass, stress tolerance, type of reproduction) within 383 angiosperm genera from Central Europe distributed along six abiotic gradients. We compared patterns of within-genus variability to those present in the entire dataset, independent of genus membership. We found that trait variability differed strongly between genera, but did not depend on their age. Trait variability was higher within genera occupying intermediate positions along regional abiotic environmental gradients, compared with patterns across the entire dataset (and unbiased by geographical sampling, family membership or species richness). Increasing trait variability within genera reflected increasing independence of traits from the abiotic environment. We conclude that intermediate abiotic environments play an important role in maintaining and possibly generating the striking diversity of life history traits present within certain clades. They may do so by relaxing the abiotic constraints on the evolution and maintenance of species traits within clades.     

Intraspecific variability improves environmental matching,but does not increase ecological breadth along a wet‐to‐dry ecotone

It is widely assumed that higher levels of intraspecific variability in one or more traits should allow species to persist under a wider range of environmental conditions. However, few studies have examined whether species that exhibit high variability are found in a wider range of environmental conditions, and whether variability increases the ability of a species to adapt to prevailing ecological gradients. We used four plant functional traits, specific leaf area (SLA), leaf dry matter content (LDMC), leaf carbon to nitrogen ratio (C:N) and maximum plant height in 49 species across a strong environmental gradient to answer three questions: 1) is there evidence for ‘high‐variability’ species (that is, species which show high variability in multiple traits, simultaneously)? 2) are species with more variable traits present across a wider range of environmental conditions than less variable species? And 3) whether more variable species show better trait–environment matching to the prevailing abiotic (soil moisture) gradient at the site? We found little evidence for a ‘high‐variability’ species. Variability was correlated for two leaf traits, SLA and LDMC, while variability in leaf traits and plant height were not correlated. We found little evidence that more variable species were present in more diverse conditions: only variation in SLA was correlated with a wider ecological niche breadth. For plant traits along the soil‐moisture gradient, higher variability led to better trait–environment matching in half of measured traits. Overall, we found little support for the existence of ‘high‐variability’ species, but that variability in SLA is correlated with a wider ecological breadth. We also found evidence that variation in traits can improve trait–environment matching, a relationship which may facilitate our understanding ecological breadth along prevailing gradients, and community assembly on the basis of traits.     

Moving from pattern to process: coexistence mechanisms under intermediate disturbance regimes

Coexistence mechanisms that require environmental variation to operate contribute importantly to the maintenance of biodiversity. One famous hypothesis of diversity maintenance under disturbance is the intermediate disturbance hypothesis (IDH). The IDH proposes patterns of peaked diversity under intermediate disturbance regimes, based on a tension between competitively superior species and species which can rapidly colonize following disturbance. We review the literature, and describe recent research that suggests that more than one underlying mechanism can generate this unimodal diversity pattern in disturbed environments. Several exciting emerging research areas are identified, including interactions between disturbance types, operation of the IDH in multi‐trophic systems, and changes in disturbance regimes. However, empirical work is still focussed on describing the IDH pattern, with little emphasis on identifying its mechanistic basis. We discuss how to extend methods for identifying different coexistence mechanisms, developed in the theoretical literature, to experimental research. In an attempt to operationalize these various ideas we outline a hypothetical IDH research programme. A solid understanding of the life history attributes of the component species and their responses to disturbance will facilitate identification of the coexistence mechanism(s) underlying the IDH pattern, and provide a framework by which empirical and theoretical results can be more fully integrated.     

Emergence dynamics and bet hedging in a desert bee,Perdita portalis

Bees (series Apiformes, superfamily Apoidea) are most diverse in arid regions of the world. Arid regions (deserts and semi-deserts) are characterized by discrete rainy seasons and extreme temporal variability in rainfall. This paper documents several novel mechanisms by which one desert bee species (Perdita portalis) copes with harsh and unpredictable conditions in xeric habitats. These same mechanisms are likely to be present in diverse bee families. First, diapausing P. portalis follow a bet-hedging emergence pattern, such that only approximately half of the larvae pupate under optimal conditions. Second, emergence is condition dependent such that larvae with a low average body weight are significantly more likely to emerge than are larvae with a high average body weight under similar conditions. Finally, larval emergence is induced by exposure to high humidity (rainfall). The parallels between bee larvae and angiosperm seeds in arid environments are striking. In both cases there is clear evidence of bet hedging, emergence (or germination) is dependent on larval (or seed) condition and rainfall triggers emergence (or germination). These patterns of emergence are significant for understanding bee species diversity in arid regions.     

Environmental gradients determine the potential for ecosystem engineering effects

Understanding processes that determine biodiversity is a fundamental challenge in ecology. At the landscape scale, physical alteration of ecosystems by organisms, called ecosystem engineering, enhances biodiversity worldwide by increasing heterogeneity in resource conditions and enhancing species coexistence across engineered and non‐engineered habitats. Engineering–diversity relationships can vary along environmental gradients due to changes in the amount of physical structuring created by ecosystem engineering, but it is unclear how this variation is influenced by the responsiveness of non‐structural abiotic properties to engineering. Here we show that environmental gradients determine the capacity for engineering to alter resource availability and species diversity, independent of the magnitude of structural change produced by engineering. We created an experimental rainfall gradient in an arid grassland where rodents restructure soils by constructing large, long‐lasting burrows. We found that greater rainfall increased water availability and productivity in both burrow and inter‐burrow habitats, causing a decline in local (alpha) plant diversity within both of these habitats. However, increased rainfall also resulted in greater differences in soil resources between burrow and inter‐burrow habitats, which increased species turnover (beta diversity) across habitats and stabilized landscape‐level (gamma) diversity. These responses occurred regardless of rodent presence and without changes in the extent of physical alteration of soils by rodents. Our results suggest that environmental gradients can influence the effects of ecosystem engineering in maintaining biodiversity via resource heterogeneity and species turnover. In an era of rapid environmental change, accounting for this interaction may be critical to conservation and management.     

The role of climate,water and biotic interactions in shaping biodiversity patterns in arid environments across spatial scales

Aim

Desert ecosystems, with their harsh environmental conditions, hold the key to understanding the responses of biodiversity to climate change. As desert community structure is influenced by processes acting at different spatial scales, studies combining multiple scales are essential for understanding the conservation requirements of desert biota. We investigated the role of environmental variables and biotic interactions in shaping broad and fine‐scale patterns of diversity and distribution of bats in arid environments to understand how the expansion of nondesert species can affect the long‐term conservation of desert biodiversity.

Location

Levant, Eastern Mediterranean.

Methods

We combine species distribution modelling and niche overlap statistics with a statistical model selection approach to integrate interspecific interactions into broadscale distribution models and fine‐scale analysis of ecological requirements. We focus on competition between desert bats and mesic species that recently expanded their distribution into arid environment following anthropogenic land‐use changes.

Results

We show that both climate and water availability limit bat distributions and diversity across spatial scales. The broadscale distribution of bats was determined by proximity to water and high temperatures, although the latter did not affect the distribution of mesic species. At the fine‐scale, high levels of bat activity and diversity were associated with increased water availability and warmer periods. Desert species were strongly associated with warmer and drier desert types. Range and niche overlap were high among potential competitors, but coexistence was facilitated through fine‐scale spatial partitioning of water resources.

Main conclusions

Adaptations to drier and warmer conditions allow desert‐obligate species to prevail in more arid environments. However, this competitive advantage may disappear as anthropogenic activities encroach further into desert habitats. We conclude that reduced water availability in arid environments under future climate change projections pose a major threat to desert wildlife because it can affect survival and reproductive success and may increase competition over remaining water resources.     

Species richness, environmental fluctuations, and temporal change in total community biomass

Theory and empirical results suggest that high biodiversity should often cause lower temporal variability in aggregate community properties such as total community biomass. We assembled microbial communities containing 2 to 8 species of competitors in aquatic microcosms and found that the temporal change in total community biomass was positively but insignificantly associated with diversity in a constant temperature environment. There was no evidence of any trend in variable temperature environments. Three non-exclusive mechanisms might explain the lack of a net stabilising effect of species richness on temporal change. (1) A direct destabilising effect of diversity on population level variances caused some populations to vary more when embedded in more diverse communities. (2) Similar responses of the different species to environmental variability might have limited any insurance effect of increased species richness. (3) Large differences in the population level variability of different species (i.e., unevenness) could weaken the relation between species richness and community level stability. These three mechanisms may outweigh the stabilising effects of increases in total community biomass with diversity, statistical averaging, and slightly more negative covariance in more diverse communities. Our experiment and analyses advocate for further experimental investigations of diversity-variability relations.     

Changes in traits of shrub canopies across an aridity gradient in northern Patagonia, Argentina

Low and highly variable precipitation pulses exert a strong selective pressure on plant traits and this might provide axes of ecological differentiation among plant species in arid ecosystems. We asked whether aridity contributes to maintain high diversity of species and morphotypes in shrub canopies. We selected eleven study sites evenly distributed across a 400-km transect in northern Patagonia, Argentina. Precipitation is low and highly variable within and between years but almost homogeneous across the transect (125–150 mm). Mean annual temperature varied, however, ranging from 8 °C (west) to 13.5 °C (east) creating a west–east gradient of aridity (aridity index from 3.7 to 7.3, respectively). Sheep grazing commenced in the early 1900s at a similar intensity across the transect. We recorded the richness and cover of shrubs by species and by morphotypes (drought deciduous tall shrubs, evergreen tall shrubs, medium shrubs, and dwarf shrubs), and further calculated the species and morphotype Shannon diversity index at each site. We assessed the presence of spiny leaves, leaf pubescence, thorny stems, and photosynthetic stems in shrub species of all morphotypes and collected green leaves of the dominant shrub species (more than 80% of the total shrub cover) to assess the leaf area, leaf mass per unit area, N-, lignin- and soluble phenolic-concentrations per species at each site. Richness and diversity of shrub species and morphotypes were positively associated with aridity. The richness and diversity of shrub species with pubescent leaves and thorny stems, and nitrogen concentration in green leaves of dominant shrubs increased with increasing aridity. We conclude that our findings on increased diversification in life history traits, species and morhotypes in shrub canopies with increasing aridity support the hypothesis that variability in aridity provides axes of ecological differentiation among shrub species facilitating their coexistence.     

Levels of genetic polymorphism: marker loci versus quantitative traits.

Species are the units used to measure ecological diversity and alleles are the units of genetic diversity. Genetic variation within and among species has been documented most extensively using allozyme electrophoresis. This reveals wide differences in genetic variability within, and genetic distances among, species, demonstrating that species are not equivalent units of diversity. The extent to which the pattern observed for allozymes can be used to infer patterns of genetic variation in quantitative traits depends on the forces generating and maintaining variability. Allozyme variation is probably not strictly neutral but, nevertheless, heterozygosity is expected to be influenced by population size and genetic distance will be affected by time since divergence. The same is true for quantitative traits influenced by many genes and under weak stabilizing selection. However, the limited data available suggest that allozyme variability is a poor predictor of genetic variation in quantitative traits within populations. It is a better predictor of general phenotypic divergence and of postzygotic isolation between populations or species, but is only weakly correlated with prezygotic isolation. Studies of grasshopper and planthopper mating signal variation and assortative mating illustrate how these characters evolve independently of general genetic and morphological variation. The role of such traits in prezygotic isolation, and hence speciation, means that they will contribute significantly to the diversity of levels of genetic variation within and among species.     

基于系统发育及功能性状的不同坡向荒漠植物群落构建研究——以博湖县沙化封禁保护区为例

荒漠植物群落构建机制的研究有助于荒漠生态系统植物资源的保护及系统平衡稳定的维持。基于系统发育与功能性状相结合的方法,以博湖县沙化封禁保护区植物群落为研究对象,在研究区内纵向沙垄的不同坡向上(丘间、阴坡、阳坡)设置样方,进行植物群落物种功能性状和土壤因子的调查与测定,通过检验植物群落物种功能性状的系统发育信号,分析不同坡向植物群落物种系统发育结构和功能结构的表现模式,利用主成分分析(PCA)和线性回归模型(Linear regression model)筛选出影响物种共存的环境因子,进而揭示研究区干旱荒漠生态系统植物群落物种共存的驱动因素。结果表明：(1)研究区植被主要以耐旱的灌木和藜科草本植物为优势种;不同坡向土壤因子具有异质性,丘间、阴坡土壤养分、水分更为丰富。(2)研究区样地植物群落物种10个功能性状指标的系统发育信号K值均小于1,说明物种功能性状受系统进化影响较小,物种功能性状未表现出系统发育保守性。(3)不同坡向系统发育结构均趋于发散,限制相似性在植物分布中占主导作用;丘间和阴坡上较丰富的土壤肥力是物种功能结构发散的主要原因,阳坡物种功能结构表现为聚集效应,生境过滤为其驱动因素...     

Phenotypic population divergence in terrestrial vertebrates at macro scales

Phenotypic divergence between populations, i.e. how much phenotypes within a species vary geographically, is critical to many aspects of ecology and evolution, including eco-geographical trends, speciation and coexistence. Yet, the variation of divergence across species with different ecologies and distributions and the relative role of adaptive causes remains little understood. We predict that genetic control vs. phenotypic plasticity of traits, geographical distance and (assuming adaptation) environmental differences should explain much of the phenotypic variability between populations. We tested these predictions with body sizes of 1447 populations in 98 terrestrial vertebrate species. Population phenotypic variability differs strongly across species, and divergence increases with increasing levels of clade-typical phenotypic plasticity, the area covered by populations and body size. Geographical distance and environmental dissimilarity are similarly important predictors of divergence within species, highlighting a potential role for biotic and environmental conditions. Increased availability of phylogeographical and ecological data should facilitate further understanding of population divergence drivers at broad scales.     

Thermophysiology,microclimates, and species distributions of lizards in the mountains of the Brazilian Atlantic Forest

Thermophysiological traits, particularly thermal tolerances and sensitivity, are key to understanding how organisms are affected by environmental conditions. In the face of ongoing climate change, determining how physiological traits structure species’ ranges is especially important in tropical montane systems. In this study, we ask whether thermal sensitivity in physiological performance restricts montane lizards to high elevations and excludes them from the warmer environments reported at low elevations. For three montane lizard species in the Brazilian Atlantic Forest, we collect thermophysiological data from lizards in the highest elevation site of each species’ distribution, and ask how well the individuals exhibiting those traits would perform across the Atlantic Forest. We use microclimatic and organism‐specific models to directly relate environmental conditions to an organism's body temperature and physiological traits, and estimate measures of thermophysiological performance. Our findings demonstrate that thermophysiological constraints do not restrict montane lizards to high elevations in this system, and thus likely do not determine the warm boundaries of these montane species’ distributions. Results also suggest that competition may be important in limiting the warm boundaries of the species’ ranges for two of the focal species. These experimental results suggest that caution should be used when claiming that physiology drives patterns of diversity and endemism within montane environments. They also highlight the importance of interdisciplinary experimental studies that bridge the fields of evolution and ecology to improve predictions of biological responses to future environmental shifts.     

Competition and coexistence in plant communities

Few ecologists today doubt that competition is an important structuring factor in plant communities, but researchers disagree on the circumstances where it is most intense, and on which traits can be considered to contribute to competitive ability in different species. The distinction between a species' effect on resources and its response to reduced resource levels might help to solve these questions. Whereas classical competition theory predicts competitive exclusion of species with similar requirements, recent ideas stress that species diversity may be explained by a multitude of processes acting at different scales, and that similarities in competitive abilities often may facilitate coexistence.     

Environmental correlates of physiological variables in marsupials

We analyzed body temperature (T(b)), basal metabolic rate (BMR), wet thermal conductance (C(wet)), and evaporative water loss (EWL) of marsupials by conventional and phylogenetically corrected regression. Allometric effects were substantial for BMR, C(wet), and EWL but not T(b). There was a strong phylogenetic signal for mass and all physiological traits. A significant phylogenetic signal remained for BMR, C(wet), and EWL even after accounting for the highly significant phylogenetic signal of mass. T(b), BMR, C(wet), and EWL allometric residuals were correlated with some diet, distribution, and climatic variables before and after correction for phylogeny. T(b) residuals were higher for marsupials from arid environments (high T(a) and more variable rainfall). The fossorial marsupial mole had a lower-than-expected T(b) residual. The allometric slope for BMR was 0.72-0.75. Residuals were consistently related to distribution aridity and rainfall variability, with species from arid and variable rainfall habitats having a low BMR, presumably to conserve energy in a low-productivity environment. The nectarivorous honey possum had a higher-than-expected BMR. For C(wet), the allometric slope was 0.55-0.62; residuals were related to diet, with folivores having low and insectivores high C(wet) residuals. The allometric slope for EWL was 0.68-0.73. EWL residuals were consistently correlated with rainfall variability, presumably facilitating maintenance of water balance during dry periods.