Extinctions in competitive communities forced by coloured environmental variation

Understanding the relationships between environmental fluctuations, population dynamics and species interactions in natural communities is of vital theoretical and practical importance. This knowledge is essential in assessing extinction risks in communities that are, for example, pressed by changing environmental conditions and increasing exploitation. We developed a model of density dependent population renewal, in a Lotka–Volterra competitive community context, to explore the significance of interspecific interactions, demographic stochasticity, population growth rate and species abundance on extinction risk in populations under various autocorrelation (colour) regimes of environmental forcing. These factors were evaluated in two cases, where either a single species or the whole community was affected by the external forcing. Species' susceptibility to environmental noise with different autocorrelation structure depended markedly on population dynamics, species' position in the abundance hierarchy and how similarly community members responded to external forcing. We also found interactions between demographic stochasticity and environmental noise leading to a reversal in extinction probabilities from under- to overcompensatory dynamics. We compare our results with studies of single species populations and contrast possible mechanisms leading to extinctions. Our findings indicate that abundance rank, the form of population dynamics, and the colour of environmental variation interact in affecting species extinction risk. These interactions are further modified by interspecific interactions within competitive communities as the interactions filter and modulate the environmental noise.     

Variation and seasonal changes in hoverfly species: interactions between temperature, age and genotype

Eristalis tenax L. and E. intricarius L. are two hoverflies which vary considerably in colour pattern. Whilst much of the phenotypic variation in both species is due to genetic variation at major gene loci, there are interactions with pupal temperature and with age of adult. Low pupal temperatures produce, on average, darker abdominal patterns in E. tenax , although the effect is hard to pick out in natural populations. Changes of pattern with age convert a bimodal autumn population to a unimodal post-hibernation population in the following spring. Hair colour is also made darker by cold treatment. Pupal treatments have a strong effect on hair colouration in E. intricarius. High temperatures inhibit the production of black hairs on the thorax, but not all the genotypes are equally sensitive to temperature. Seasonal fluctuations in colour pattern frequencies are detectable in E. intricarius. In three Eristalis species that have been studied so far, the interactions between genotype, age and temperature operate in quite different ways. The temperature responses may be relevant to theories of thermal melanism, although the colour pattern polymorphisms are, more obviously, examples of Batesian mimicry.     

Nuclear and mitochondrial data reveal different evolutionary processes in the Lake Tanganyika cichlid genus <Emphasis Type="Italic">Tropheus</Emphasis>

Background  

Cichlid fishes are notorious for their wealth of intra- and interspecific colour pattern diversity. In Lake Tanganyika, the endemic genus Tropheus represents the most impressive example for geographic variation in the pattern and hue of integument colouration, but the taxonomy of the over 100 mostly allopatric colour morphs remains to a large degree unresolved. Previous studies of mitochondrial DNA sequence data revealed polyphyly of the six nominally described species and complex phylogeographic patterns influenced by lake level fluctuations and population admixture, and suggested the parallel evolution of similar colour patterns in divergent evolutionary lineages. A gene tree of a rapidly radiating group may be subject to incomplete and stochastic lineage sorting, and to overcome this problem we used multi-locus, nuclear AFLP data in comparison with mtDNA sequences to study diversification, migration and introgression in Tropheus colour morphs in Lake Tanganyika.     

Variable colour patterns indicate multidimensional,intraspecific trait variation and ecological generalization in moths

Animal colour patterns long have provided information about key processes that drive the ecological and evolutionary dynamics of biological diversity. Theory and empirical evidence indicate that variation in colour patterns and other traits among individuals generally improves the performance of populations and species, for example by reducing predation risk, increasing establishment success, improving resilience to environmental change, and decreasing risk of extinction. However, little is known about whether and how variation in colour pattern among species is associated with variation in other phenotypic dimensions. To address this issue, we analysed associations of colour pattern with morphological, behavioural and life-history traits on the basis of data for nearly 400 species of noctuid moths. We found that moths with more variable colour patterns had longer flight activity periods, more diverse habitats and a greater number of host plant species than species with less variable colour patterns. Variable coloration in adult noctuid moths therefore can be considered as indicative of broader niches and generalist diets. Colour pattern variability was not significantly associated with overwintering stage or body size (wing span), and it was independent of whether the colour pattern of the larvae was non-variable, variable or highly variable. Colour pattern variation during the larval stage tended to increase as the duration of the flight activity period increased, but was independent of the length of the larval period, diet breadth and habitat use. The realization that information on colour pattern variation in adult moths, and possibly other organisms, offers a proxy for niche breadth and dietary generalization can inform management and conservation biology.     

Inter‐individual variation in colour patterns in noctuid moths characterizes long‐distance dispersers and agricultural pests

A high capacity for long‐distance dispersal is a key to success for species confronted with environmental heterogeneity, habitat modification, fragmentation and loss. However, dispersal capacity is difficult to quantify and therefore poorly known in most taxa. Here, we report on a test for an association of variation in dispersal capacity with variable colouration of noctuid moths. First, using data from 12 experienced lepidopterologists, we showed that despite variation among experts in average assessments, different species are consistently classified as having non‐variable, variable or highly variable colour patterns when assessed by different experts. We then compared the incidence of non‐resident species with high inter‐individual variation in colour patterns recorded on the isolated island Utklippan (n = 47), with that in a species pool of potential long‐distance dispersers from the nearest mainland (n = 295). Species with high inter‐individual colour pattern variation were over‐represented on the island compared with species having non‐variable colouration. This finding constitutes rare evidence from the wild of long‐distance dispersal, measured on a spatial scale relevant for moths when tracking habitats in fragmented and changing landscapes or when keeping pace with environmental challenges associated with climate change. Finally, we showed that Swedish noctuid moths classified as agricultural pests (n = 28) had more variable colour patterns compared with non‐pests (n = 368). The majority of agricultural pests were also recorded on the isolated island, an outcome that is indicative of pest species having high dispersal capacity. Data on colour pattern variation may thus offer a simple and cost‐effective proxy to estimate dispersal capacity and can also help identify potential pest species. Our findings are potentially useful when modelling and predicting population and range dynamics of species in spatiotemporally heterogeneous environments, with direct implications for conservation biology and pest management.     

Self-organized instability in complex ecosystems

Why are some ecosystems so rich, yet contain so many rare species? High species diversity, together with rarity, is a general trend in neotropical forests and coral reefs. However, the origin of such diversity and the consequences of food web complexity in both species abundances and temporal fluctuations are not well understood. Several regularities are observed in complex, multispecies ecosystems that suggest that these ecologies might be organized close to points of instability. We explore, in greater depth, a recent stochastic model of population dynamics that is shown to reproduce: (i) the scaling law linking species number and connectivity; (ii) the observed distributions of species abundance reported from field studies (showing long tails and thus a predominance of rare species); (iii) the complex fluctuations displayed by natural communities (including chaotic dynamics); and (iv) the species-area relations displayed by rainforest plots. It is conjectured that the conflict between the natural tendency towards higher diversity due to immigration, and the ecosystem level constraints derived from an increasing number of links, leaves the system poised at a critical boundary separating stable from unstable communities, where large fluctuations are expected to occur. We suggest that the patterns displayed by species-rich communities, including rarity, would result from such a spontaneous tendency towards instability.     

Commonly Rare and Rarely Common: Comparing Population Abundance of Invasive and Native Aquatic Species

Invasive species are leading drivers of environmental change. Their impacts are often linked to their population size, but surprisingly little is known about how frequently they achieve high abundances. A nearly universal pattern in ecology is that species are rare in most locations and abundant in a few, generating right-skewed abundance distributions. Here, we use abundance data from over 24,000 populations of 17 invasive and 104 native aquatic species to test whether invasive species differ from native counterparts in statistical patterns of abundance across multiple sites. Invasive species on average reached significantly higher densities than native species and exhibited significantly higher variance. However, invasive and native species did not differ in terms of coefficient of variation, skewness, or kurtosis. Abundance distributions of all species were highly right skewed (skewness>0), meaning both invasive and native species occurred at low densities in most locations where they were present. The average abundance of invasive and native species was 6% and 2%, respectively, of the maximum abundance observed within a taxonomic group. The biological significance of the differences between invasive and native species depends on species-specific relationships between abundance and impact. Recognition of cross-site heterogeneity in population densities brings a new dimension to invasive species management, and may help to refine optimal prevention, containment, control, and eradication strategies.     

Concordant population dynamics of Lepidoptera herbivores in a forest ecosystem

It is frequently assumed that population fluctuations are largely independent within a community of trophically‐similar species, but this need not be so. If population fluctuations are partly synchronized or concordant, this will produce interannual variability in the community's aggregate abundance and generate temporal variance in ecosystem structure. We studied the community of Lepidoptera inhabiting northern hardwood forests in New Hampshire, USA, to evaluate the hypothesis that fluctuations in consumer communities can arise from concordant dynamics of constituent populations. Interannual comparisons of moth abundances for >75 species sampled at three sites over four years revealed that concordant dynamics contribute strongly to interannual variability in the abundance of consumers. A conspicuous decline in community abundance from 2004 to 2005 was the result of predominantly negative population growth rates of the component species, while an increase in community abundance from 2006 to 2007 was the result of predominantly positive population growth rates. Population dynamics most strongly linked species that feed in the early season (perhaps due to shared responses to climatic effects), but not species that might share natural enemies or host plants. The observed concordant dynamics introduced conspicuous temporal variation in the abundance of primary consumers relative to plants and secondary consumers, thereby altering the forest's trophic structure. Such variance in the aggregate abundance of forest primary consumers could generate time‐lagged fluctuations in abundances of secondary consumers and will generally have important consequences for ecosystem properties and processes that are nonlinear functions of consumer abundance, such as plant community structure and nutrient cycling.     

Low Predictability of Colour Polymorphism in Introduced Guppy (Poecilia reticulata) Populations in Panama

Colour polymorphism is a recurrent feature of natural populations, and its maintenance has been studied in a range of taxa in their native ranges. However, less is known about whether (and how) colour polymorphism is maintained when populations are removed from their native environments, as in the case of introduced species. We here address this issue by analyzing variation in colour patterns in recently-discovered introduced populations of the guppy (Poecilia reticulata) in Panama. Specifically, we use classic colour analysis to estimate variation in the number and the relative area of different colour spots across low predation sites in the introduced Panamanian range of the species. We then compare this variation to that found in the native range of the species under low- and high predation regimes. We found aspects of the colour pattern that were both consistent and inconsistent with the classical paradigm of colour evolution in guppies. On one hand, the same colours that dominated in native populations (orange, iridescent and black) were also the most dominant in the introduced populations in Panama. On the other, there were no clear differences between either introduced-low and native low- and high predation populations. Our results are therefore only partially consistent with the traditional role of female preference in the absence of predators, and suggest that additional factors could influence colour patterns when populations are removed from their native environments. Future research on the interaction between female preference and environmental variability (e.g. multifarious selection), could help understand adaptive variation in this widely-introduced species, and the contexts under which variation in adaptive traits parallels (or not) variation in the native range.     

Prevalence of intraspecific relationships between range size and abundance in Danish birds

In this study, we investigate patterns in the prevalence of dynamic range–abundance relationships of the Danish avifauna, using breeding bird atlases from 1971 to 1974 and from 1993 to 1996. We focus on differences between common and rare species by dividing the assemblage into range-size quartiles. The trend in total population size was determined using an index. Range was determined as grid cell occupancy and standardized to facilitate comparisons between common and rare species. While narrow-ranging species showed strong and consistent range–abundance relationships, the relationships for widespread species were weak and exhibited considerable variation. This may be due to differences in patterns of resource use, since widespread species generally have wider niches, and so may be less affected by resource-based factors linking range and abundance. Since a tight and dynamic relationship is upheld for rare species, monitoring strategies based on range size surveys seem viable.     

Population fluctuations affect inference in ecological networks of multi‐species interactions

Local abundance and population fluctuations are key factors affecting the realized interaction frequencies in biotic interactions, but they are commonly ignored when network metrics are calculated over aggregated sets of observations. Here we studied how abundance fluctuations (i.e. demographic and stochastic population dynamics in one of the trophic levels) may affect derived network‐level inferences in bipartite ecological networks. Variation at both the species and network level in network indices (d’, Dependence, Fisher's alpha diversity for both levels, H₂’, weighted NODF) were strongly correlated with the extent of abundance fluctuations, with a strong effect of environmental stochasticity on all indices except NODF; this was the only index for which considerable variation was caused by varying carrying capacities among species. Binary connectance, in turn, does not take interaction frequency (and thus abundance) into account and was only influenced by abundance fluctuations at low population sizes if this led to non‐occurrence of ‘true’ interactions. Overall, abundance and population dynamics are likely to play an important role in determining what is commonly observed and summarized into ecological networks. We suggest that ecological network inference should account for underlying population dynamics and uncertainty in what is observed as interaction frequencies, modelling mechanisms at operative organisational levels below the network rather than using aggregated data of observations. Modelling population dynamics may be a valuable tool in this field to conceptualize and tease apart different sources of variation and express uncertainty in our inference from small samples.     

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。     

Diurnal lighting patterns and habitat alter opsin expression and colour preferences in a killifish

Spatial variation in lighting environments frequently leads to population variation in colour patterns, colour preferences and visual systems. Yet lighting conditions also vary diurnally, and many aspects of visual systems and behaviour vary over this time scale. Here, we use the bluefin killifish (Lucania goodei) to compare how diurnal variation and habitat variation (clear versus tannin-stained water) affect opsin expression and the preference to peck at different-coloured objects. Opsin expression was generally lowest at midnight and dawn, and highest at midday and dusk, and this diurnal variation was many times greater than variation between habitats. Pecking preference was affected by both diurnal and habitat variation but did not correlate with opsin expression. Rather, pecking preference matched lighting conditions, with higher preferences for blue at noon and for red at dawn/dusk, when these wavelengths are comparatively scarce. Similarly, blue pecking preference was higher in tannin-stained water where blue wavelengths are reduced. In conclusion, L. goodei exhibits strong diurnal cycles of opsin expression, but these are not tightly correlated with light intensity or colour. Temporally variable pecking preferences probably result from lighting environment rather than from opsin production. These results may have implications for the colour pattern diversity observed in these fish.     

Spatio-temporal dynamics in waterbirds during the non-breeding season: Effects of local movements,migration and weather are monthly,not yearly

Predicting population sizes and dynamics of mobile organisms is important for trend estimation, but this is difficult when the origin of individuals cannot be determined, i.e. residents and migrants are indistinguishable in the field.Here we examine fluctuations in populations during the non-breeding season (autumn to spring) of seven waterbird species, which co-occur on 122 neighbouring water bodies in south-west Germany. We asked whether site-level abundance patterns can be predicted by weather variables, and whether these effects vary over months and years. We used a spatially explicit hidden Markov model to estimate the effects of local and regional movement on population fluctuations.Although the species varied in their dynamics, with different amounts of movement estimated between sites, several patterns were common across species: density effects were highly month-specific, but with little variation in the strength of effects over years. The abundance of most species was positively related to temperature, especially in winter.The common teal Anas crecca and common pochard Aythya ferina were the most site-fidelic species once they were present in the study area, but also exhibited the strongest regional migration. The mallard Anas platyrhynchos, tufted duck Aythya fuligula and great crested grebe Podiceps cristatus each behaved more like a single population, as individuals more frequently moved between sites and abundance fluctuations at sites were not explained by migratory movements alone.Our study shows that the strength of population parameters and environmental forces can be decomposed into monthly and yearly effects. Estimating the unknown origin and movement of individuals may show that commensurate populations of mobile species may have different underlying dynamics, while responding similarly to environmental factors.     

Larval dispersal dampens population fluctuation and shapes the interspecific spatial distribution patterns of rocky intertidal gastropods

Many marine benthic invertebrates pass through a planktonic larval stage whereas others spend their entire lifetimes in benthic habitats. Recent studies indicate that non‐planktonic species show relatively greater fine‐scale patchiness than do planktonic species, but the underlying mechanisms remain unknown. One hypothesis for such a difference is that larval dispersal enhances the connectivity of populations and buffers population fluctuations and reduces local extinction risk, consequently increasing patch occupancy rate and decreasing spatial patchiness. If this mechanism does indeed play a significant role, then the distribution of non‐planktonic species should be more aggregated – both temporally and spatially – than the distribution of species with a planktonic larval stage. To test this prediction, we compared 1) both the spatial and the temporal abundance–occupancy relationships and 2) both the spatial and the temporal mean–variance relationships of population size across species of rocky intertidal gastropods with differing dispersive traits from the Pacific coast of Japan. We found that, compared to planktonic species, non‐planktonic species exhibited 1) a smaller occupancy rate for any given level of mean population size and 2) greater variations in population size, both spatially and temporally. This suggests that the macroecological patterns observed in this study (i.e. the abundance–occupancy relationships and mean–variance relationships of population size across species) were shaped by the effect of larval dispersal dampening population fluctuation, which works over both space and time. While it has been widely assumed that larval dispersal enhances population fluctuations, larval dispersal may in fact enhance the connectively of populations and buffer population fluctuations and reduce local extinction risks.     

Synchrony of woodland bird populations: the effect of landscape structure

The influence of environmental stochasticity and dispersal in producing patterns in population synchrony was examined for 53 woods censused annually from 1990 to 1999 for nine resident bird species (wren Troglodytes troglodytes , dunnock Prunella modularis , robin Erithacus rubecula , blackbird Turdus merula , song thrush Turdus philomelos , long-tailed tit Aegithalos caudatus , blue tit Parus caeruleus , great tit Parus major , and chaffinch Fringilla coelebs ) and four migrant bird species (garden warbler Sylvia borin , blackcap Sylvia atricapilla , chiffchaff Phylloscopus collybita and willow warbler Phylloscopus trochilus ). Twelve species showed global synchrony of population counts due to regional population trends and widespread annual population fluctuations. There was a clear link between population fluctuations and winter weather for wren, and three other species showed their largest population declines after the coldest winters. Eight species showed a decline in synchrony with distance between woods, and there was evidence for dispersal causing this pattern in three species. Landscape structure affected patterns of synchrony in several species, with lower synchrony in landscapes with less woodland. For three species, this difference in synchrony across a landscape gradient of decreasing woodland cover accounted for the decline in synchrony over distance. Three species showed greater synchrony between woods with similar amounts of hedgerow in the surrounding landscape, suggesting that the surroundings of a wood influence the population dynamics of some species breeding in the wood. Habitat fragmentation can alter the processes contributing to population synchrony. Loss of woodland reduces the relative abundance of woodland bird species. The remaining patches of habitat are smaller, more isolated and are set in a more hostile landscape, all of which may disrupt dispersal between patches and alter the population dynamics within woods.     

Latitudinal variation in genetic divergence of populations and the potential for future speciation

The increase in biological diversity with decreasing latitude is widely appreciated but the cause of the pattern is unknown. This pattern reflects latitudinal variation in both the origin of new species (cladogenesis) and the number of species that coexist. Here we address latitudinal variation in species origination, by examining population genetic processes that influence speciation. Previous data suggest a greater number of speciation events at lower latitudes. If speciation events occur more frequently at lower latitudes, we predicted that genetic divergence among populations within species, an important component of cladogenesis, should be greater among lower latitude populations. We tested this prediction using within-species patterns of mtDNA variation across 60 vertebrate species that collectively spanned six continents, two oceans, and 119 degrees latitude. We found greater genetic divergence of populations, controlling for geographic distance, at lower latitudes within species. This pattern remained statistically significant after removing populations that occur in localities previously covered by continental glaciers during the last glaciation. Results suggest that lower latitude populations within species exhibit greater evolutionary independence, increasing the likelihood that mutation, recombination, selection, and/or drift will lead to divergence of traits important for reproductive isolation and speciation. Results are consistent with a greater influence of seasonality, reduced energy, and/or glacial (Milankovitch) cycles acting on higher latitude populations, and represent one of the few tests of predictions of latitudinal variation in speciation rates using population genetic data.     

The contribution of common and rare species to plant species richness patterns: the effect of habitat type and size of sampling unit

Understanding how overall patterns of spatial variation in species richness are affected by distributional patterns of species has been an area of growing concern. In the present study, we investigated the relative importance of common and rare species as contributors in overall plant species richness. We further examined if the effects of common or rare species in richness patterns are affected by the size of the sampling units and if the observed patterns hold at different habitats. We used a dataset of 5,148 higher plant species distributed across 16,114 sampling plots located in 240 sites of the NATURA 2000 network of Greece. We ranked all species based on the number of sites they occupied and we developed a common to rare and a rare to common sequence. We correlated those sequences with cumulative species distributions. We performed this analysis in nine different sizes of sampling units and in three different datasets referring to (a) all habitat types together, (b) coniferous habitats only and (c) alpine habitats only. Our analysis showed that despite the proportionally higher numbers of restricted species, widespread species make a greater contribution to overall richness patterns and that this observed pattern does not depend on the size of the sampling units. Moreover, the observed pattern stands for different habitat types. Our findings support the generality of this pattern and highlight the importance of widespread species as adequate indicators of biodiversity patterns at various habitat types. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Spatial synchrony in red grouse population dynamics

Red grouse Lagopus lagopus scoticus populations exhibit unstable dynamics that are often characterised by regular periodic fluctuations in abundance. Time-series' of grouse harvesting records collected from 287 management units (moors) across Scotland, England and Wales were analysed to investigate the broad scale patterns of synchrony in these fluctuations. Estimation of the spatial autocorrelation of grouse population dynamics across moors indicates relatively high levels of synchrony between populations on adjacent moors, but that this synchrony declines sharply with increasing inter-moor distance. At distances of greater than 100  km, grouse population time-series exhibit only weakly positive cross-correlation coefficients. Twenty-eight geographical, environmental and other candidate variables were examined to construct a general linear model to explain variation in local synchrony. Grouse moor productivity (average size of shooting bag), distance from the Atlantic coast moving in a north-easterly direction, April and June temperatures, and June rainfall significantly increased the explanatory power of this model. An understanding of the processes underlying synchrony in red grouse population dynamics is a prerequisite to anticipating the effects of large-scale environmental change on regional patterns of grouse distribution and abundance.