Adjacent woodlands rather than habitat connectivity influence grassland plant,carabid and bird assemblages in farmland landscapes

One response to biodiversity decline is the definition of ecological networks that extend beyond protected areas and promote connectivity in human-dominated landscapes. In farmland, landscape ecological research has focused more on wooded than open habitat networks. In our study, we assessed the influence of permanent grassland connectivity, described by grassland amount and spatial configuration, on grassland biodiversity. We selected permanent grasslands in livestock farming areas of north-western France, which were sampled for plants, carabids and birds. At two spatial scales we tested the effects of amount and configuration of grasslands, wooded habitats and crops on richness and abundance of total assemblages and species ecological groups. Grassland connectivity had no significant effects on total richness or abundance of any taxonomic group, regardless of habitat affinity or dispersal ability. The amount of wooded habitat and length of wooded edges at the 200 m scale positively influenced forest and generalist animal groups as well as grassland plant species, in particular animal-dispersed species. However, for animal groups such as open habitat carabids or farmland bird specialists, the same wooded habitats negatively influenced richness and abundance at the 500 m scale. The scale and direction of biodiversity responses to landscape context were therefore similar among taxonomic groups, but opposite for habitat affinity groups. We conclude that while grassland connectivity is unlikely to contribute positively to biodiversity, increasing or maintaining wooded elements near grasslands would be a worthwhile conservation goal. However, the requirements of open farmland animal species groups must be considered, for which such action may be deleterious.     

A long‐term habitat fragmentation experiment leads to morphological change in a species of carabid beetle

1. Habitat fragmentation and transformation are key drivers of species declines in landscapes. Most of the current understanding of species' responses to environmental change originates from studies of populations and communities. However, phenotypic variation offers another key aspect of species responses and could provide additional insights into the functional drivers of population change. 2. The goal of this study was to address this gap by exploring the morphological changes of a species of carabid beetle (Notonomus resplendens) with a known population response to the Wog Wog Habitat Fragmentation Experiment in Australia. We measured morphological traits associated with body size, head width, and dispersal ability. We quantified patterns of morphological variation over time and between native Eucalyptus forest fragments and the surrounding pine plantation matrix and the continuous intact native Eucalyptus forest controls. 3. We found sexually dimorphic morphological changes in response to the experimental treatments. Males increased in size, had larger legs and had smaller interocular widths in the matrix in both the short and long terms. Conversely, females became comparatively smaller and had increased interocular widths in the same treatments. Effects in the fragments were similar to those in the matrix, but exhibited more uncertainty. 4. Our results demonstrate that species can show morphological change in response to environmental change over very short time periods. We demonstrate that using both population and morphological data allows stronger inferences about the mechanisms behind species responses to environmental change.     

Realized correlated responses to artificial selection on pre-adult life-history traits in a butterfly

We use artificial selection experiments targeted on egg size, development time or pupal mass within a single butterfly population followed by a common-garden experiment to explore the interactions among these life-history traits. Relationships were predicted to be negative between egg size and development time, but to be positive between development time and body size and between egg size and body size. Correlated responses to selection were in part inconsistent with these predictions. Although there was evidence for a positive genetic correlation between egg and body size, there was no support for genetic correlations between larval development time and either egg size or pupal mass. Phenotypic correlations among the three target traits of selection gave comparable results for the relationships between egg mass and development time (no association) as well as between egg mass and pupal mass (positive association), but not for the relation between development time and pupal mass (negative phenotypic correlation). In summary, correlated responses to selection as well as phenotypic correlations were rather unpredictable. The impact of variation in acquisition and allocation of energy as well as of the benign conditions used deserve further investigation.     

Within- and between-generation effects of temperature on life-history traits in a butterfly

Non-genetic parental effects may largely affect offspring phenotype, and such plasticity is potentially adaptive. Despite its potential importance, little is known about cross-generational effects of temperature, at least partly because parental effects were frequently considered a troublesome nuisance, rather than a target of experimental studies. We here investigate effects of parental, developmental and acclimation temperature on life-history traits in the butterfly Bicyclus anynana. Higher developmental temperatures reduced development times and egg size, increased egg number, but did not affect pupal mass. Between-generation temperature effects on larval time, pupal time, larval growth rate and egg size were qualitatively very similar to effects of developmental temperature, and additionally affected pupal mass but not egg number. Parental effects are important mediators of phenotypic plasticity in B. anynana, and partly yielded antagonistic effects on different components of fitness, which may constrain the evolution of cross-generational adaptive plasticity.     

Genetic equilibrium despite habitat fragmentation in an Afrotropical bird

We examined the effects of habitat fragmentation of the white-starred robin Pogonocichla stellata metapopulation in the Taita Hills archipelago, a hotspot for biodiversity which was fragmented approximately 40 years ago. Using seven microsatellite markers, we analysed the robin's genetic structure and tested for equilibrium between migration and drift (testing the probability of decreased dispersal) as well as between mutation and drift (test for recent reduction in effective population size, i.e. bottlenecks). This metapopulation was found to retain relatively high levels of genetic variability (H(E) between 0.63 and 0.71) and to be in migration-drift equilibrium, suggesting that increased isolation between fragments did not have much effect on the dispersal between them. Furthermore, this equilibrium test greatly enhanced the interpretation of parameters (e.g. F(ST)) assumed to have reached an equilibrium value. In contrast to previous findings on the related and sympatric Taita thrush Turdus helleri (which is critically endangered), there were no indications for recent bottlenecks in any of the robin subpopulations. This difference can be attributed to the higher dispersal capacity of the robin compared with the thrush (deduced from both the genetic and capture-recapture data). Our results stress the importance of sustained dispersal for species conservation.     

Linking alternative food sources to winter habitat selection of herbivores in overbrowsed landscapes

During winter, ungulates in boreal forests must cope with high energetic costs related to locomotion in deep snow and reduced forage abundance and quality. At high density, ungulates face additional constraints, because heavy browsing reduces availability of woody browse, the main source of forage during winter. Under these severe conditions, large herbivores might forage on alternative food sources likely independent of browsing pressure, such as litterfall or windblown trees. We investigated the influence of alternative food sources on winter habitat selection, by studying female white-tailed deer (Odocoileus virginianus) living in 2 landscapes with contrasted browse abundance, recently logged and regenerated landscapes, in a population at high density and on a large island free of predators. We fitted 21 female white-tailed deer with Global Positioning System (GPS) collars and delineated winter home ranges and core areas. We measured snow conditions in different habitat categories and sampled vegetation in the core areas and in the rest of the home ranges to determine how forage abundance, protective cover, and snow conditions influenced habitat selection within the home range. In both landscapes, deer were less likely to use open habitat categories as snow accumulated on the ground. At a finer scale, deer inhabiting the regenerated landscape intensively used areas where balsam fir cover was intermediate with greater balsam fir browse density than in the rest of the home range. In the recently logged landscape, deer were more likely to be found near edges between clear-cuts and balsam fir stands and in areas where windblown balsam fir trees were present; the latter being the most influential variable. Although balsam fir browse was sparse and mainly out of reach in this landscape, deer increased the use of areas where it was present. Our results offer novel insights into the resource selection processes of northern ungulates, as we showed that access to winter forage, such as woody browse and alternative food sources, depends on climatic conditions and stochastic events, such as abundant compacted snow or windthrows. To compensate for these scarce and unpredictable food supplies, deer selected habitat categories, but mostly areas within those habitat categories, where the likelihood of finding browse, litterfall, and windblown trees was greatest. © 2011 The Wildlife Society.     

Compounding effects of habitat fragmentation and predation on bird nests

Habitat fragmentation and invasive species are two of the greatest threats to species diversity worldwide. This is particularly relevant for oceanic islands with vulnerable endemics. Here, we examine how habitat fragmentation influences nest predation by Rattus spp. on cup‐nesting birds in Samoan forests. We determined models for predicting predation rates by Rattus on artificial nests at two scales: (i) the position of the bird's nest within the landscape (e.g. proximity to mixed crop plantations, distance to forest edge); and (ii) the microhabitat in the immediate vicinity of the nest (e.g. nest height, ground cover, slope). Nest cameras showed only one mammal predator, the black rat (Rattus rattus), predating artificial nests. The optimal model predicting nest predation rates by black rats included a landscape variable, proximity to plantations and a local nest site variable, the percentage of low (<15 cm) ground cover surrounding the nest tree. Predation rates were 22 ± 13% higher for nests in forest edges near mixed crop plantations than in edges without plantations. In contrast, predation rates did not vary significantly between edge habitat where the matrix did not contain plantations, and interior forest sites (>1 km from the edge). As ground cover reduced, nest predation rates increased. Waxtags containing either coconut or peanut butter were used as a second method for assessing nest predation. The rates at which these were chewed followed patterns similar to the predation of the artificial nests. Rural development in Samoa will increase the proportion of forest edge near plantations. Our results suggest that this will increase the proportion of forest birds that experience nest predation from black rats. Further research is required to determine if rat control is needed to maintain even interior forest sites populations of predator‐sensitive bird species on South Pacific islands.     

Linking habitat specialization with species' traits in European birds

Ecological specialization provides information about adaptations of species to their environment. However, identification of traits representing the relevant dimensions of ecological space remains challenging. Here we endeavoured to explain how complex habitat specializations relate to various ecological traits of European birds. We employed phylogenetic generalized least squares and information theoretic approach statistically controlling for differences in geographic range size among species. Habitat specialists had narrower diet niche, wider climatic niche, higher wing length/tail length ratio and migrated on shorter distances than habitat generalists. Our results support an expected positive link between habitat and diet niche breadth estimates, however a negative relationship between habitat and climate niche breadths is surprising. It implies that habitat specialists occur mostly in spatially restricted environments with high climatic variability such as mountain areas. This, however, complicates our understanding of predicted impacts of climatic changes on avian geographical distributions. Our results further corroborate that habitat specialization reflects occupation of morphological space, when specialists depend more on manoeuvrability of the flight and are thus more closely associated to open habitats than habitat generalists. Finally, our results indicate that long distance movements might hamper narrow habitat preferences. In conclusion, we have shown that species’ distributions across habitats are informative about their positions along other axes of ecological space and can explain states of particular functional traits, however, our results also reveal that the links between different niche estimates cannot be always straightforwardly predicted.     

Effect of habitat fragmentation on dispersal in the butterfly Proclossiana eunomia

Comparison of dispersal rates of the bog fritillary butterfly between continuous and fragmented landscapes indicates that between patch dispersal is significantly lower in the fragmented landscape, while population densities are of the same order of magnitude. Analyses of the dynamics of the suitable habitat for the butterfly in the fragmented landscape reveal a severe, non linear increase in spatial isolation of patches over a time period of 30 years (i.e. 30 butterfly generations), but simulations of the butterfly metapopulation dynamics using a structured population model show that the lower dispersal rates in the fragmented landscape are far above the critical threshold leading to metapopulation extinction. These results indicate that changes in individual behaviour leading to the decrease of dispersal rates in the fragmented landscape were rapidly selected for when patch spatial isolation increased. The evidence of such an adaptive answer to habitat fragmentation suggests that dispersal mortality is a key factor for metapopulation persistence in fragmented landscapes. We emphasise that landscape spatial configuration and patch isolation have to be taken into account in the debate about large-scale conservation strategies.     

Linking habitat use to range expansion rates in fragmented landscapes: a metapopulation approach

Temperature increases because of climate change are expected to cause expansions at the high latitude margins of species distributions, but, in practice, fragmented landscapes act as barriers to colonization for most species. Understanding how species distributions will shift in response to climate change therefore requires techniques that incorporate the combined effects of climate and landscape‐scale habitat availability on colonization rates. We use a metapopulation model (Incidence Function Model, IFM) to test effects of fine‐scale habitat use on patterns and rates of range expansion by the butterfly Hesperia comma. At its northern range margin in Britain, this species has increased its breadth of microhabitat use because of climate warming, leading to increased colonization rates. We validated the IFM by reconstructing expansions in five habitat networks between 1982 and 2000, before using it to predict metapopulation dynamics over 100 yr, for three scenarios based on observed changes to habitat use. We define the scenarios as “cold‐world” (only hot, south‐facing 150–250° hillsides are deemed warm enough), “warm‐world” in which 100–300° hillsides can be populated, and “hot‐world”, where the background climate is warm enough to enable use of all aspects (as increasingly observed). In the simulations, increased habitat availability in the hot‐world scenario led to faster range expansion rates, and to long‐term differences in distribution size and pattern. Thus, fine‐scale changes in the distribution of suitable microclimates led to landscape‐scale changes in population size and colonization rate, resulting in coarse‐scale changes to the species distribution. Despite use of a wider range of habitats associated with climate change, H. comma is still expected to occupy a small fraction of available habitat in 100 yr. The research shows that metapopulation models represent a potential framework to identify barriers to range expansion, and to predict the effects of environmental change or conservation interventions on species distributions and persistence.     

Forest edges have high conservation value for bird communities in mosaic landscapes

A major conservation challenge in mosaic landscapes is to understand how trait‐specific responses to habitat edges affect bird communities, including potential cascading effects on bird functions providing ecosystem services to forests, such as pest control. Here, we examined how bird species richness, abundance and community composition varied from interior forest habitats and their edges into adjacent open habitats, within a multi‐regional sampling scheme. We further analyzed variations in Conservation Value Index (CVI), Community Specialization Index (CSI) and functional traits across the forest‐edge‐open habitat gradient. Bird species richness, total abundance and CVI were significantly higher at forest edges while CSI peaked at interior open habitats, i.e., furthest from forest edge. In addition, there were important variations in trait‐ and species‐specific responses to forest edges among bird communities. Positive responses to forest edges were found for several forest bird species with unfavorable conservation status. These species were in general insectivores, understorey gleaners, cavity nesters and long‐distance migrants, all traits that displayed higher abundance at forest edges than in forest interiors or adjacent open habitats. Furthermore, consistently with predictions, negative edge effects were recorded in some forest specialist birds and in most open‐habitat birds, showing increasing densities from edges to interior habitats. We thus suggest that increasing landscape‐scale habitat complexity would be beneficial to declining species living in mosaic landscapes combining small woodlands and open habitats. Edge effects between forests and adjacent open habitats may also favor bird functional guilds providing valuable ecosystem services to forests in longstanding fragmented landscapes.     

The impact of hedge-forest connectivity and microhabitat conditions on spider and carabid beetle assemblages in agricultural landscapes

Agricultural intensification in terms of decreasing landscape complexity and connectivity has negatively affected biodiversity. Linear landscape elements composed of woody vegetation like hedges may counteract this negative trend by providing habitats and enhancing habitat connectivity for different organisms. Here, we tested the impacts of habitat type (forest edges vs. hedges) and hedges’ isolation (connected vs. isolated hedges) from forests as well as microhabitat conditions (percentage of bare ground and width) on trait-specific occurrence of ground-dwelling arthropods, namely spiders and carabids. Arthropods were grouped by habitat specialisation (forest vs. open-habitat species vs. generalists), hunting strategy (web-building or hunting spiders) and dispersal ability (wing morphology of carabids). Spider and carabid assemblage composition was strongly influenced by habitat type and isolation, but not by microhabitat conditions. Activity density of forest species and brachypterous carabids was higher in forest edges compared to hedges, whereas open-habitat species and macropterous carabids showed reverse patterns, with no effects of isolation. Occurrence of generalist carabids, but not spiders, was higher in hedges compared to forest edges. Habitat type and isolation did not affect spiders with different hunting strategy. Microhabitat conditions were less important for spider and carabid occurrence. Our study concludes that on a landscape scale, type of linear woody habitat is more important for arthropod occurrence than isolation effects and microhabitat conditions, depending on traits. Hedges provide refuges for species specialised to open habitats and species with high dispersal ability, such as macropterous carabids. Forest edges enhance persistence of species specialised to forests and species with low dispersal ability, such as brachypterous carabids.     

Spatial and habitat variation in aphid,butterfly, moth and bird phenologies over the last half century

Global warming has advanced the timing of biological events, potentially leading to disruption across trophic levels. The potential importance of phenological change as a driver of population trends has been suggested. To fully understand the possible impacts, there is a need to quantify the scale of these changes spatially and according to habitat type. We studied the relationship between phenological trends, space and habitat type between 1965 and 2012 using an extensive UK dataset comprising 269 aphid, bird, butterfly and moth species. We modelled phenologies using generalized additive mixed models that included covariates for geographical (latitude, longitude, altitude), temporal (year, season) and habitat terms (woodland, scrub, grassland). Model selection showed that a baseline model with geographical and temporal components explained the variation in phenologies better than either a model in which space and time interacted or a habitat model without spatial terms. This baseline model showed strongly that phenologies shifted progressively earlier over time, that increasing altitude produced later phenologies and that a strong spatial component determined phenological timings, particularly latitude. The seasonal timing of a phenological event, in terms of whether it fell in the first or second half of the year, did not result in substantially different trends for butterflies. For moths, early season phenologies advanced more rapidly than those recorded later. Whilst temporal trends across all habitats resulted in earlier phenologies over time, agricultural habitats produced significantly later phenologies than most other habitats studied, probably because of nonclimatic drivers. A model with a significant habitat‐time interaction was the best‐fitting model for birds, moths and butterflies, emphasizing that the rates of phenological advance also differ among habitats for these groups. Our results suggest the presence of strong spatial gradients in mean seasonal timing and nonlinear trends towards earlier seasonal timing that varies in form and rate among habitat types.     

Modelling the effect of habitat fragmentation on range expansion in a butterfly

There is an increasing need for conservation programmes to make quantitative predictions of biodiversity responses to changed environments. Such predictions will be particularly important to promote species recovery in fragmented landscapes, and to understand and facilitate distribution responses to climate change. Here, we model expansion rates of a test species (a rare butterfly, Hesperia comma) in five landscapes over 18 years (generations), using a metapopulation model (the incidence function model). Expansion rates increased with the area, quality and proximity of habitat patches available for colonization, with predicted expansion rates closely matching observed rates in test landscapes. Habitat fragmentation constrained expansion, but in a predictable way, suggesting that it will prove feasible both to understand variation in expansion rates and to develop conservation programmes to increase rates of range expansion in such species.     

Hybridisation between two cyprinid fishes in a novel habitat: genetics,morphology and life-history traits

Background  

The potential role hybridisation in adaptive radiation and the evolution of new lineages has received much recent attention. Hybridisation between roach (Rutilus rutilus L.) and bream (Abramis brama L.) is well documented throughout Europe, however hybrids in Ireland occur at an unprecedented frequency, often exceeding that of both parental species. Utilising an integrated approach, which incorporates geometric morphometrics, life history and molecular genetic analyses we identify the levels and processes of hybridisation present, while also determining the direction of hybridisation, through the analysis of mitochondrial DNA.     

Sex-specific genetic variances in life-history and morphological traits of the seed beetle Callosobruchus maculatus

Knowledge of heritability and genetic correlations are of central importance in the study of adaptive trait evolution and genetic constraints. We use a paternal half-sib-full-sib breeding design to investigate the genetic architecture of three life-history and morphological traits in the seed beetle, Callosobruchus maculatus. Heritability was significant for all traits under observation and genetic correlations between traits (r(A)) were low. Interestingly, we found substantial sex-specific genetic effects and low genetic correlations between sexes (r(MF)) in traits that are only moderately (weight at emergence) to slightly (longevity) sexually dimorphic. Furthermore, we found an increased sire ([Formula: see text]) compared to dam ([Formula: see text]) variance component within trait and sex. Our results highlight that the genetic architecture even of the same trait should not be assumed to be the same for males and females. Furthermore, it raises the issue of the presence of unnoticed environmental effects that may inflate estimates of heritability. Overall, our study stresses the fact that estimates of quantitative genetic parameters are not only population, time, environment, but also sex specific. Thus, extrapolation between sexes and studies should be treated with caution.     

Nestedness of butterfly assemblages in the Zhoushan Archipelago,China: area effects,life-history traits and conservation implications

The nested subset pattern (nestedness) of faunal assemblages has been a research focus in the fields of island biogeography and conservation biology in recent decades. However, relatively few studies have described nestedness in butterfly assemblages in oceanic archipelago systems. Moreover, previous studies often quantified nestedness using inappropriate nestedness metrics and random fill algorithms with high Type I errors. The aims of this study are to examine the existence of nestedness and underlying causal mechanisms of butterfly assemblages in the Zhoushan Archipelago, China. We used the line-transect method to determine butterfly occupancy and abundance on 42 study islands from July to August 2014. We obtained butterfly life-history traits (wingspan, body weight and minimum area requirement) by field work and island geographical features (area and isolation) from the literature. We used the recently developed metric WNODF to estimate nestedness. Partial Spearman rank correlation was used to evaluate the associations of nestedness and island geographical features as well as butterfly life-history traits related to species extinction risk and colonization ability. The butterfly assemblages were significantly nested. Island area and minimum area requirement of butterflies were significantly correlated with nestedness after controlling for other independent variables. In contrast, the nestedness of butterflies did not appear to result from passive sampling or selective colonization. However, multi-year studies are needed to confirm that target effects are not muddling these results. Our results indicate that selective extinction may be the main driver of nestedness of butterfly assemblages in our study system. From a conservation viewpoint, we should protect both large islands and species with large area requirement to maximize the number of species preserved.