Spatial variation in the magnitude and functional form of density‐dependent processes on the large skipper butterfly Ochlodes sylvanus

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Niche breadth explains the range size of European-centred butterflies,but dispersal ability does not

Aim

The breadth of ecological niches and dispersal abilities have long been discussed as important determinants of species' range sizes. However, studies directly comparing the relative effects of both factors are rare, taxonomically biased and revealed inconsistent results.

Location

Europe.

Time Period

Cenozoic.

Major Taxa

Butterflies, Lepidoptera.

Methods

We relate climate, diet and habitat niche breadth and two indicators of dispersal ability, wingspan and a dispersal tendency index, to the global range size of 369 European-centred butterfly species. The relative effects of these five predictors and their variation across the butterfly phylogeny were assessed by means of phylogenetic generalized least squares models and phylogenetically weighted regressions respectively.

Results

Climate niche breadth was the most important single predictor, followed by habitat and diet niche breadth, while dispersal tendency and wingspan showed no relation to species' range size. All predictors together explained 59% of the variation in butterfly range size. However, the effects of each predictor varied considerably across families and genera.

Main Conclusions

Range sizes of European-centred butterflies are strongly correlated with ecological niche breadth but apparently independent of dispersal ability. The magnitude of range size–niche breadth relationships is not stationary across the phylogeny and is often negatively correlated across the different dimensions of the ecological niche. This variation limits the generalizability of range size–trait relationships across broad taxonomic groups.     

Traits help explain species' performance away from their climate niche centre

Aim

Climate change impacts on biota are variable across sites, among species and throughout individual species' ranges. Niche theory predicts that population performance should decline as site climate becomes increasingly different from the species' climate niche centre, though studies find significant variation from these predictions. Here, we propose that predictions about climate responses can be improved by incorporating species' trait information.

Location

Europe.

Methods

We used observations of plant species abundance change over time to assess variation in climate difference sensitivity (CDS), defined as how species performance (colonization, extinction and abundance change) relates to the difference of site climate from the mean temperature and precipitation of each species' range. We then investigated if leaf economics, plant size and seed mass traits were associated with the species' CDS.

Results

Species that performed better (e.g. increased in abundance) towards sites progressively cooler than their niche centre were shorter and had more resource-acquisitive leaves (i.e. lower leaf dry matter content or LDMC) relative to species with zero or the opposite pattern of temperature difference sensitivity. This result supports the hypothesis that if sites cooler than niche centres are more stressful for a species, then shorter stature is advantageous compared with taller species. The LDMC result suggests the environment selects for more resource-acquisitive leaf strategies towards relatively cooler climates with shorter growing seasons, counter to expectations that conservative strategies would be favoured in such environments. We found few consistent relationships between precipitation difference sensitivities and traits.

Main Conclusions

The results supported key a priori foundations on how trait-based plant strategies dictate species responses to climate variation away from their niche centre. Furthermore, plant height emerged as the most consistent trait that varied with species CDS, suggesting height will be key for theory development around species response to climate change.     

Geographical barriers and climate influence demographic history in narrowleaf cottonwoods

Studies of genetic variation can clarify the role of geography and spatio-temporal variation of climate in shaping demography, particularly in temperate zone tree species with large latitudinal ranges. Here, we examined genetic variation in narrowleaf cottonwood, Populus angustifolia, a dominant riparian tree. Using multi-locus surveys of polymorphism in 363 individuals across the species'' 1800 km latitudinal range, we found that, first, P. angustifolia has stronger neutral genetic structure than many forest trees (simple sequence repeat (SSR) F_ST=0.21), with major genetic groups corresponding to large apparent geographical barriers to gene flow. Second, using SSRs and putatively neutral sequenced loci, coalescent simulations indicated that populations diverged before the last glacial maximum (LGM), suggesting the presence of population structure before the LGM. Third, the LGM and subsequent warming appear to have had different influences on each of these distinct populations, with effective population size reduction in the southern extent of the range but major expansion in the north. These results are consistent with the hypothesis that climate and geographic barriers have jointly affected the demographic history of P. angustifolia, and point the importance of both factors as being instrumental in shaping genetic variation and structure in widespread forest trees.     

Increasing frequency of low summer precipitation synchronizes dynamics and compromises metapopulation stability in the Glanville fritillary butterfly

Climate change is known to shift species'' geographical ranges, phenologies and abundances, but less is known about other population dynamic consequences. Here, we analyse spatio-temporal dynamics of the Glanville fritillary butterfly (Melitaea cinxia) in a network of 4000 dry meadows during 21 years. The results demonstrate two strong, related patterns: the amplitude of year-to-year fluctuations in the size of the metapopulation as a whole has increased, though there is no long-term trend in average abundance; and there is a highly significant increase in the level of spatial synchrony in population dynamics. The increased synchrony cannot be explained by increasing within-year spatial correlation in precipitation, the key environmental driver of population change, or in per capita growth rate. On the other hand, the frequency of drought during a critical life-history stage (early larval instars) has increased over the years, which is sufficient to explain the increasing amplitude and the expanding spatial synchrony in metapopulation dynamics. Increased spatial synchrony has the general effect of reducing long-term metapopulation viability even if there is no change in average metapopulation size. This study demonstrates how temporal changes in weather conditions can lead to striking changes in spatio-temporal population dynamics.     

Predicting the sensitivity of butterfly phenology to temperature over the past century

Studies to date have documented substantial variation among species in the degree to which phenology responds to temperature and shifts over time, but we have a limited understanding of the causes of such variation. Here, we use a spatially and temporally extensive data set (ca. 48 000 observations from across Canada) to evaluate the utility of museum collection records in detecting broad‐scale phenology‐temperature relationships and to test for systematic differences in the sensitivity of phenology to temperature (days °C^?1) of Canadian butterfly species according to relevant ecological traits. We showed that the timing of flight season predictably responded to temperature both across space (variation in average temperature from site to site in Canada) and across time (variation from year to year within each individual site). This reveals that collection records, a vastly underexploited resource, can be applied to the quantification of broad‐scale relationships between species' phenology and temperature. The timing of the flight season of earlier fliers and less mobile species was more sensitive to temperature than later fliers and more mobile species, demonstrating that ecological traits can account for some of the interspecific variation in species' phenological sensitivity to temperature. Finally, we found that phenological sensitivity to temperature differed across time and space implying that both dimensions of temperature will be needed to translate species' phenological sensitivity to temperature into accurate predictions of species' future phenological shifts. Given the widespread temperature sensitivity of flight season timing, we can expect long‐term temporal shifts with increased warming [ca. 2.4 days °C^?1 (0.18 SE)] for many if not most butterfly species.     

Refining predictions of population decline at species' rear edges

According to broad‐scale application of biogeographical theory, widespread retractions of species' rear edges should be seen in response to ongoing climate change. This prediction rests on the assumption that rear edge populations are “marginal” since they occur at the limit of the species' ecological tolerance and are expected to decline in performance as climate warming pushes them to extirpation. However, conflicts between observations and predictions are increasingly accumulating and little progress has been made in explaining this disparity. We argue that a revision of the concept of marginality is necessary, together with explicit testing of population decline, which is increasingly possible as data availability improves. Such action should be based on taking the population perspective across a species' rear edge, encompassing the ecological, geographical and genetic dimensions of marginality. Refining our understanding of rear edge populations is essential to advance our ability to monitor, predict and plan for the impacts of environmental change on species range dynamics.     

Microclimate and resource quality determine resource use in a range-expanding herbivore

The consequences of climate change for biogeographic range dynamics depend on the spatial scales at which climate influences focal species directly and indirectly via biotic interactions. An overlooked question concerns the extent to which microclimates modify specialist biotic interactions, with emergent properties for communities and range dynamics. Here, we use an in-field experiment to assess egg-laying behaviour of a range-expanding herbivore across a range of natural microclimatic conditions. We show that variation in microclimate, resource condition and individual fecundity can generate differences in egg-laying rates of almost two orders of magnitude in an exemplar species, the brown argus butterfly (Aricia agestis). This within-site variation in fecundity dwarfs variation resulting from differences in average ambient temperatures among populations. Although higher temperatures did not reduce female selection for host plants in good condition, the thermal sensitivities of egg-laying behaviours have the potential to accelerate climate-driven range expansion by increasing egg-laying encounters with novel hosts in increasingly suitable microclimates. Understanding the sensitivity of specialist biotic interactions to microclimatic variation is, therefore, critical to predict the outcomes of climate change across species'' geographical ranges, and the resilience of ecological communities.     

Global drivers of population density in terrestrial vertebrates

Aim

Although the effects of life history traits on population density have been investigated widely, how spatial environmental variation influences population density for a large range of organisms and at a broad spatial scale is poorly known. Filling this knowledge gap is crucial for global species management and conservation planning and to understand the potential impact of environmental changes on multiple species.

Location

Global.

Time period

Present.

Major taxa studied

Terrestrial amphibians, reptiles, birds and mammals.

Methods

We collected population density estimates for a range of terrestrial vertebrates, including 364 estimates for amphibians, 850 for reptiles, 5,667 for birds and 7,651 for mammals. We contrasted the importance of life history traits and environmental predictors using mixed models and tested different hypotheses to explain the variation in population density for the four groups. We assessed the predictive accuracy of models through cross‐validation and mapped the partial response of vertebrate population density to environmental variables globally.

Results

Amphibians were more abundant in wet areas with high productivity levels, whereas reptiles showed relatively higher densities in arid areas with low productivity and stable temperatures. The density of birds and mammals was typically high in temperate wet areas with intermediate levels of productivity. The models showed good predictive abilities, with pseudo‐R² ranging between 0.68 (birds) and 0.83 (reptiles).

Main conclusions

Traits determine most of the variation in population density across species, whereas environmental conditions explain the intraspecific variation across populations. Species traits, resource availability and climatic stability have a different influence on the population density of the four groups. These models can be used to predict the average species population density over large areas and be used to explore macroecological patterns and inform conservation analyses.     

Loss of adaptive variation during evolutionary responses to climate change

The changes in species' geographical distribution demanded by climate change are often critically limited by the availability of key interacting species. In such cases, species' persistence will depend on the rapid evolution of biotic interactions. Understanding evolutionary limits to such adaptation is therefore crucial for predicting biological responses to environmental change. The recent poleward range expansion of the UK brown argus butterfly has been associated with a shift in female preference from its main host plant, rockrose (Cistaceae), onto Geraniaceae host plants throughout its new distribution. Using reciprocal transplants onto natural host plants across the UK range, we demonstrate reduced fitness of females from recently colonised Geraniaceae‐dominated habitat when moved to ancestral rockrose habitats. By contrast, individuals from ancestral rockrose habitats show no reduction in fitness on Geraniaceae. Climate‐driven range expansion in this species is therefore associated with the rapid evolution of biotic interactions and a significant loss of adaptive variation.     

The influence of floral symmetry,dependence on pollinators and pollination generalization on flower size variation

Background and Aims

The pollinator-mediated stabilizing selection hypothesis suggests that the specialized pollination system of zygomorphic flowers might cause stabilizing selection, reducing their flower size variation compared with actinomorphic flowers. However, the degree of ecological generalization and of dependence on pollinators varies greatly among species of both flower symmetry types and this may also affect flower size variation.

Methods

Data on 43 species from two contrasting communities (one alpine and one lowland community) were used to test the relationships and interactions between flower size phenotypic variation, floral symmetry, ecological pollination generalization and species'' dependence on pollinators.

Key Results

Contrary to what was expected, higher flower size variation was found in zygomorphic than in actinomorphic species in the lowland community, and no difference in flower size variation was found between symmetry types in the alpine community. The relationship between floral symmetry and flower size variation depended on ecological generalization and species'' dependence on pollinators, although the influence of ecological generalization was only detected in the alpine community. Zygomorphic species that were highly dependent on pollinators and that were ecologically specialized were less variable in flower size than ecologically generalist and selfing zygomorphic species, supporting the pollinator-mediated stabilizing selection hypothesis. However, these relationships were not found in actinomorphic species, probably because they are not dependent on any particular pollinator for efficient pollination and therefore their flower size always shows moderate levels of variation.

Conclusions

The study suggests that the relationship between flower size variation and floral symmetry may be influenced by population-dependent factors, such as ecological generalization and species'' dependence on pollinators.     

Leaf hydraulic vulnerability to drought is linked to site water availability across a broad range of species and climates

Background and Aims

Vulnerability of the leaf hydraulic pathway to water-stress-induced dysfunction is a key component of drought tolerance in plants and may be important in defining species'' climatic range. However, the generality of the association between leaf hydraulic vulnerability and climate across species and sites remains to be tested.

Methods

Leaf hydraulic vulnerability to drought (P50_leaf, the water potential inducing 50 % loss in hydraulic function) was measured in a diverse group of 92 woody, mostly evergreen angiosperms from sites across a wide range of habitats. These new data together with some previously published were tested against key climate indices related to water availability. Differences in within-site variability in P50_leaf between sites were also examined.

Key Results

Values of hydraulic vulnerability to drought in leaves decreased strongly (i.e. became more negative) with decreasing annual rainfall and increasing aridity across sites. The standard deviation in P50_leaf values recorded within each site was positively correlated with increasing aridity. P50_leaf was also a good indicator of the climatic envelope across each species'' distributional range as well as their dry-end distributional limits within Australia, although this relationship was not consistently detectable within sites.

Conclusions

The findings indicate that species sorting processes have influenced distributional patterns of P50_leaf across the rainfall spectrum, but alternative strategies for dealing with water deficit exist within sites. The strong link to aridity suggests leaf hydraulic vulnerability may influence plant distributions under future climates.     

Geographical structuring of genetic diversity across the whole distribution range of Narcissus longispathus, a habitat-specialist, Mediterranean narrow endemic

Background and Aims

High mountain ranges of the Mediterranean Basin harbour a large number of narrowly endemic plants. In this study an investigation is made of the levels and partitioning of genetic diversity in Narcissus longispathus, a narrow endemic of south-eastern Spanish mountains characterized by a naturally fragmented distribution due to extreme specialization on a rare habitat type. By using dense sampling of populations across the species'' whole geographical range, genetic structuring at different geographical scales is also examined.

Methods

Using horizontal starch-gel electrophoresis, allozyme variability was screened at 19 loci for a total of 858 individuals from 27 populations. The data were analysed by means of standard statistical approaches in order to estimate gene diversity and the genetic structure of the populations.

Key Results

Narcissus longispathus displayed high levels of genetic diversity and extensive diversification among populations. At the species level, the percentage of polymorphic loci was 68 %, with average values of 2·1, 0·11 and 0·14 for the number of alleles per locus, observed heterozygosity and expected heterozygosity, respectively. Southern and more isolated populations tended to have less genetic variability than northern and less-isolated populations. A strong spatial patterning of genetic diversity was found at the various spatial scales. Gene flow/drift equilibrium occurred over distances <4 km. Beyond that distance divergence was relatively more influenced by drift. The populations studied seem to derive from three panmictic units or ‘gene pools’, with levels of admixture being greatest in the central and south-eastern portions of the species'' range.

Conclusions

In addition to documenting a case of high genetic diversity in a narrow endemic plant with naturally fragmented populations, the results emphasize the need for dense population sampling and examination of different geographical scales for understanding population genetic structure in habitat specialists restricted to ecological islands.Key words: Allozymes, genetic diversity, geographical scale, habitat isolation, Narcissus longispathus, Mediterranean endemism, mountain range, natural fragmented distribution     

Joint effects of species traits and environmental preferences on range edge shifts of British birds

Aim

Despite the strong evidence of species range shifts as a response to environmental change, attempts to identify species traits that modulate those shifts have been equivocal. We investigate the role of species traits and environmental preferences on birds' range shifts in Great Britain, an island where dispersal is limited by the English Channel and the North Sea.

Location

Great Britain (England, Scotland and Wales).

Taxa

Birds (Aves).

Time Period

1968–2011.

Methods

Using 404,949 occurrence records from two time periods, we investigated the potential drivers of leading and rear range edge shifts of breeding birds using phylogenetic linear mixed models. We hypothesized that shifts are influenced by species' trophic and morphological traits, dispersal abilities and environmental preferences, but also by the geographical boundaries of Great Britain.

Results

Geographical boundaries—the distance from the northern or southern boundaries of Britain—accounted for most of the variability in range edge shifts. Species traits and environmental preferences emerged as relevant drivers of range shifts only for northern and Passeriform species. Northern habitat specialist, those with more predators and those sensitive to precipitation were more likely to shift their rear edge poleward. For Passeriformes, habitat generalists, species with smaller dispersal capabilities, under higher predatory pressure or associated with forest and grassland were more likely to shift their rear edge poleward.

Main Conclusions

While geographical boundaries impose constraints on range shifts in British birds, the subtle effects of species traits and environmental preferences emerge as relevant predictors for Northern and passeriform species' rear edge shifts. This highlights the importance of accounting for geographical boundaries when predicting species responses to global change. Differential range shifts of species across different trophic levels could result in the reorganization of biotic interactions, with consequences for ecosystem structure and stability.     

Heat resistance throughout ontogeny: body size constrains thermal tolerance

Heat tolerance is a trait of paramount ecological importance and may determine a species' ability to cope with ongoing climate change. Although critical thermal limits have consequently received substantial attention in recent years, their potential variation throughout ontogeny remained largely neglected. We investigate whether such neglect may bias conclusions regarding a species' sensitivity to climate change. Using a tropical butterfly, we found that developmental stages clearly differed in heat tolerance. It was highest in pupae followed by larvae, adults and finally eggs and hatchlings. Strikingly, most of the variation found in thermal tolerance was explained by differences in body mass, which may thus impose a severe constraint on adaptive variation in stress tolerance. Furthermore, temperature acclimation was beneficial by increasing heat knock‐down time and therefore immediate survival under heat stress, but it affected reproduction negatively. Extreme temperatures strongly reduced survival and subsequent reproductive success even in our highly plastic model organism, exemplifying the potentially dramatic impact of extreme weather events on biodiversity. We argue that predictions regarding a species' fate under changing environmental conditions should consider variation in thermal tolerance throughout ontogeny, variation in body mass and acclimation responses as important predictors of stress tolerance.     

Range expansion through fragmented landscapes under a variable climate

Ecological responses to climate change may depend on complex patterns of variability in weather and local microclimate that overlay global increases in mean temperature. Here, we show that high‐resolution temporal and spatial variability in temperature drives the dynamics of range expansion for an exemplar species, the butterfly Hesperia comma. Using fine‐resolution (5 m) models of vegetation surface microclimate, we estimate the thermal suitability of 906 habitat patches at the species' range margin for 27 years. Population and metapopulation models that incorporate this dynamic microclimate surface improve predictions of observed annual changes to population density and patch occupancy dynamics during the species' range expansion from 1982 to 2009. Our findings reveal how fine‐scale, short‐term environmental variability drives rates and patterns of range expansion through spatially localised, intermittent episodes of expansion and contraction. Incorporating dynamic microclimates can thus improve models of species range shifts at spatial and temporal scales relevant to conservation interventions.     

Breeding ground temperature rises,more than habitat change,are associated with spatially variable population trends in two species of migratory bird

Habitat loss and climate change are key drivers of global biodiversity declines but their relative importance has rarely been examined. We attempted to attribute spatially divergent population trends of two Afro-Palaearctic migrant warbler species, Willow Warbler Phylloscopus trochilus and Common Chiffchaff Phylloscopus collybita, to changes in breeding grounds climate or habitat. We used bird counts from over 4000 sites across the UK between 1994 and 2017, monitored as part of the BTO/JNCC/RSPB Breeding Bird Survey. We modelled Willow Warbler and Common Chiffchaff population size and growth in relation to habitat, climate and weather. We then used the abundance model coefficients and observed environmental changes to determine the extent to which spatially varying population trends in England and Scotland were consistent with attribution to climate and habitat changes. Both species' population size and growth correlated with habitat, climate and weather on their breeding grounds. Changes in habitat, in particular woodland expansion, could be linked to small population increases for both species in England and Scotland. Both species' populations correlated more strongly with climate than weather, and both had an optimum breeding season temperature: 11°C for Willow Warbler and around 13.5°C for Common Chiffchaff (with marginally different predictions from population size and growth models). Breeding ground temperature increases, therefore, had the potential to have caused some of the observed Willow Warbler declines in England (where the mean breeding season temperature was 12.7°C) and increases in Scotland (mean breeding season temperature was 10.2°C), and some of the differential rates of increase for Common Chiffchaff. However, much of the variation in species' population abundance and trends were not well predicted by our models and could be due to other factors, such as species interactions, habitat and climate change in their wintering grounds and on migration. This study provides evidence that the effect of climate change on a species may vary spatially and may switch from being beneficial to being detrimental if a temperature threshold is exceeded.     

Reduced variability in range‐edge butterfly populations over three decades of climate warming

Populations at the high latitude edge of species’ geographical ranges are thought to show larger interannual population fluctuations, with subsequent higher local extinction risk, than those within the ‘core’ climatic range. As climate envelopes shift northward under climate warming, however, we would expect populations to show dampened variability. We test this hypothesis using annual abundance indices from 19 butterfly species across 79 British monitoring sites between 1976 and 2009, a period of climatic warming. We found that populations in the latter (warmer) half of the recording period show reduced interannual population variability. Species with more southerly European distributions showed the greatest dampening in population variability over time. Our results suggest that increases in population variability occur towards climatic range boundaries. British sites, previously existing at the margins of suitable climate space, now appear to fall closer to the core climatic range for many butterfly species.     

Geographic differences in Blainville's beaked whale (Mesoplodon densirostris) echolocation clicks

Aim

Understanding cetacean species' distributions and population structure over space and time is necessary for effective conservation and management. Geographic differences in acoustic signals may provide a line of evidence for population-level discrimination in some cetacean species. We use acoustic recordings collected over broad spatial and temporal scales to investigate whether global variability in echolocation click peak frequency could elucidate population structure in Blainville's beaked whale (Mesoplodon densirostris), a cryptic species well-studied acoustically.

Location

North Pacific, Western North Atlantic and Gulf of Mexico.

Time period

2004–2021.

Major taxa studied

Blainville's beaked whale.

Methods

Passive acoustic data were collected at 76 sites and 150 cumulative years of data were analysed to extract beaked whale echolocation clicks. Using an automated detector and subsequent weighted network clustering on spectral content and interclick interval of clicks, we determined the properties of a primary cluster of clicks with similar characteristics per site. These were compared within regions and across ocean basins and evaluated for suitability as population-level indicators.

Results

Spectral averages obtained from primary clusters of echolocation clicks identified at each site were similar in overall shape but varied in peak frequency by up to 8 kHz. We identified a latitudinal cline, with higher peak frequencies occurring in lower latitudes.

Main conclusions

It may be possible to acoustically delineate populations of Blainville's beaked whales. The documented negative correlation between signal peak frequency and latitude could relate to body size. Body size has been shown to influence signal frequency, with lower frequencies produced by larger animals, which are subsequently more common in higher latitudes for some species, although data are lacking to adequately investigate this for beaked whales. Prey size and depth may shape frequency content of echolocation signals, and larger prey items may occur in higher latitudes, resulting in lower signal frequencies of their predators.     

Broad-scale latitudinal variation in female reproductive success contributes to the maintenance of a geographic range boundary in bagworms (Lepidoptera: Psychidae)

Background

Geographic range limits and the factors structuring them are of great interest to biologists, in part because of concerns about how global change may shift range boundaries. However, scientists lack strong mechanistic understanding of the factors that set geographic range limits in empirical systems, especially in animals.

Methodology/Principal Findings

Across dozens of populations spread over six degrees of latitude in the American Midwest, female mating success of the evergreen bagworm Thyridopteryx ephemeraeformis (Lepidoptera: Psychidae) declines from ∼100% to ∼0% near the edge of the species range. When coupled with additional latitudinal declines in fecundity and in egg and pupal survivorship, a spatial gradient of bagworm reproductive success emerges. This gradient is associated with a progressive decline in local abundance and an increased risk of local population extinction, up to a latitudinal threshold where extremely low female fitness meshes spatially with the species'' geographic range boundary.

Conclusions/Significance

The reduction in fitness of female bagworms near the geographic range limit, which concords with the abundant centre hypothesis from biogeography, provides a concrete, empirical example of how an Allee effect (increased pre-reproductive mortality of females in sparsely populated areas) may interact with other demographic factors to induce a geographic range limit.