Active around the year: Butterflies and moths adapt their life cycles to a warming world.

Living in a warming world requires adaptations to altered annual temperature regimes. In Europe, spring is starting earlier, and the vegetation period is ending later in the year. These climatic changes are leading not only to shifts in distribution ranges of flora and fauna, but also to phenological shifts. Using long-term observation data of butterflies and moths collected during the past decades across northern Austria, we test for phenological shifts over time and changes in the number of generations. On average, Lepidoptera adults emerged earlier in the year and tended to extend their flight periods in autumn. Many species increased the annual number of generations. These changes were more pronounced at lower altitudes than at higher altitudes, leading to an altered phenological zonation. Our findings indicate that climate change does not only affect community composition but also the life history of insects. Increased activity and reproductive periods might alter Lepidoptera-host plant associations and food webs.

Global change drives phenological and spatial shifts in Central European longhorn beetles (Coleoptera, Cerambycidae) during the past 150 years.

Temperature increases and land-use changes induce altered annual activity periods of arthropods. However, sufficiently resolved long-term data sets (> 100 years) are mostly missing. We use a data set of longhorn beetle records (71 species) collected in Luxembourg 1864-2014. Increase of annual temperatures was significantly correlated with an earlier annual appearance. Forty-four species present before and after 1980 appeared on average 8.2 days earlier in the year in the more recent period. Since 1950, the estimated shift was 0.26 days per year. Increase of temperature in spring (March-June) preponed the first appearance of beetles by on average 9.6 days per 1 °C. We found significant changes in the composition of beetle communities, with a net gain in species richness during the last 40 years. Eleven species recorded only after 1997 were characterized by comparatively early annual appearance. Smaller beetles tended to appear earlier in the year in comparison to large-bodied species. Shifts in phenology did not correlate with species Red List status. As also demonstrated by our data, climate change in general affects insect phenologies and changes species composition. However, land-use change has taken place in parallel with climate change. Both aspects of global change are influencing the changes in longhorn beetle occurrences in Luxemburg in their combination. This might be most clearly reflected in the strong decrease of species with continental climate niches dwelling in old-growth deciduous forests that apparently are threatened by the loss of these habitats and increasing spring temperatures.

Holocene shifts in the assembly of plant and animal communities implicate human impacts.

Understanding how ecological communities are organized and how they change through time is critical to predicting the effects of climate change. Recent work documenting the co-occurrence structure of modern communities found that most significant species pairs co-occur less frequently than would be expected by chance. However, little is known about how co-occurrence structure changes through time. Here we evaluate changes in plant and animal community organization over geological time by quantifying the co-occurrence structure of 359,896 unique taxon pairs in 80 assemblages spanning the past 300 million years. Co-occurrences of most taxon pairs were statistically random, but a significant fraction were spatially aggregated or segregated. Aggregated pairs dominated from the Carboniferous period (307 million years ago) to the early Holocene epoch (11,700 years before present), when there was a pronounced shift to more segregated pairs, a trend that continues in modern assemblages. The shift began during the Holocene and coincided with increasing human population size and the spread of agriculture in North America. Before the shift, an average of 64% of significant pairs were aggregated; after the shift, the average dropped to 37%. The organization of modern and late Holocene plant and animal assemblages differs fundamentally from that of assemblages over the past 300 million years that predate the large-scale impacts of humans. Our results suggest that the rules governing the assembly of communities have recently been changed by human activity.

Biogeography of Italy revisited: genetic lineages confirm major phylogeographic patterns and a pre-Pleistocene origin of its biota.

Warm-adapted species survived the cold stages of the past glacial-interglacial cycles in southern European peninsulas and recolonized major parts of Central and Northern Europe in the wake of postglacial warming. However, many of the genetic lineages which differentiated within these refugia predate even the Pleistocene. One of these refugia is the Italian Peninsula with its surrounding islands. In this meta-analysis, we compiled phylogeographic patterns of multiple species across this region. We transformed existing geographic information on 78 animal and plant species (with a total of 471 genetic lineages) within 17 predefined areas into presence/absence matrices. We elaborated three matrices: (i) only old lineages exclusively distinguished by deep splits, (ii) only young lineages distinguished by shallow (i.e. recent) splits, and (iii) presence/absence of the respective species. To infer biogeographic relationships between the predefined areas, we performed bootstrapped neighbour joining cluster analyses on these three matrices. In addition, we reviewed the geological history of Italy to identify causes of the observed biogeographic patterns. We found Sardinia and Corsica to be biogeographically closely linked with each other, and that they diverge strongly from all other regions. Sicily also diverges strongly from all other regions, while the intra-island differentiation was comparatively low. On the Italian mainland, Calabria exhibited the most pronounced biogeographic differentiation, often with several lineages present, resulting from old vicariance events within the region. Furthermore, southern Apulia and the Po Plain with adjoining areas of northern peninsular Italy displayed considerable distinctiveness. Admixture prevailed in the areas between these three regions. The ancient isolation of Sicily, as well as Sardinia plus Corsica, resulted in endemic lineages with only moderate recent exchange with adjacent mainland regions. Pronounced diversification occurs within the Italian Peninsula. The complex tectonic activities, such as shifting (micro)plates, submergence of major parts of peninsular Italy with the genesis of numerous Pliocene islands, in combination with the climatic cycles during the Pleistocene have most likely generated the current biogeographic pattern of species. Immigrations from the Balkan Peninsula into northern Italy partly accounted for the distinctiveness of this region.

Evolution of contact and alarm calls in the Kenyan endemic Hinde's babbler (Aves: Passeriformes).

BACKGROUND: Spatial isolation, diverging environmental conditions and social structures may lead to the differentiation of various traits, e.g. molecules, morphology and behaviour. Bird calls may provide important information on effects of geographic isolation and may reflect diverging ecological conditions related to altitude. Furthermore, bird calls are strongly shaped by the social behaviour of species. The Kenyan endemic bird Hinde's Babbler, Turdoides hindei, is a cooperative breeder existing in distinct family groups. The species occurs in five isolated population groups at different altitudes across its distribution range in south-eastern Kenya. With this model species we test for potential effects of geographic isolation, diverging altitudes, and social structures. We recorded and analysed contact and alarm calls of T. hindei, including its entire distribution range and all existing population groups. RESULTS: Our data show significant differentiation of call characteristics among population groups across the species' distribution range. This differentiation is correlated with geographical distance, but also with altitude. We also found strong call differentiation among neighbouring family groups. Call differentiation of contact calls was comparatively high in comparison to alarm calls, which showed a lower degree of divergence. CONCLUSION: Our data show that call differentiation is governed by geographic isolation as well as altitude. Diverging degrees of call differentiation in contact and alarm calls suggests that both call types are under different selective pressures. Alarm calls are required to be understood by all members of the species across the entire distribution range and thus call differentiation is lower. In contrast, contact calls are more specific and differ even among neighbouring families supporting the maintenance of distinct bird families and groups.

Bi-dimensional null model analysis of presence-absence binary matrices.

Comparing the structure of presence/absence (i.e., binary) matrices with those of randomized counterparts is a common practice in ecology. However, differences in the randomization procedures (null models) can affect the results of the comparisons, leading matrix structural patterns to appear either "random" or not. Subjectivity in the choice of one particular null model over another makes it often advisable to compare the results obtained using several different approaches. Yet, available algorithms to randomize binary matrices differ substantially in respect to the constraints they impose on the discrepancy between observed and randomized row and column marginal totals, which complicates the interpretation of contrasting patterns. This calls for new strategies both to explore intermediate scenarios of restrictiveness in-between extreme constraint assumptions, and to properly synthesize the resulting information. Here we introduce a new modeling framework based on a flexible matrix randomization algorithm (named the "Tuning Peg" algorithm) that addresses both issues. The algorithm consists of a modified swap procedure in which the discrepancy between the row and column marginal totals of the target matrix and those of its randomized counterpart can be "tuned" in a continuous way by two parameters (controlling, respectively, row and column discrepancy). We show how combining the Tuning Peg with a wise random walk procedure makes it possible to explore the complete null space embraced by existing algorithms. This exploration allows researchers to visualize matrix structural patterns in an innovative bi-dimensional landscape of significance/effect size. We demonstrate the rational and potential of our approach with a set of simulated and real matrices, showing how the simultaneous investigation of a comprehensive and continuous portion of the null space can be extremely informative, and possibly key to resolving longstanding debates in the analysis of ecological matrices.

Increasing aridity reduces soil microbial diversity and abundance in global drylands.

Soil bacteria and fungi play key roles in the functioning of terrestrial ecosystems, yet our understanding of their responses to climate change lags significantly behind that of other organisms. This gap in our understanding is particularly true for drylands, which occupy ∼41% of Earth´s surface, because no global, systematic assessments of the joint diversity of soil bacteria and fungi have been conducted in these environments to date. Here we present results from a study conducted across 80 dryland sites from all continents, except Antarctica, to assess how changes in aridity affect the composition, abundance, and diversity of soil bacteria and fungi. The diversity and abundance of soil bacteria and fungi was reduced as aridity increased. These results were largely driven by the negative impacts of aridity on soil organic carbon content, which positively affected the abundance and diversity of both bacteria and fungi. Aridity promoted shifts in the composition of soil bacteria, with increases in the relative abundance of Chloroflexi and α-Proteobacteria and decreases in Acidobacteria and Verrucomicrobia. Contrary to what has been reported by previous continental and global-scale studies, soil pH was not a major driver of bacterial diversity, and fungal communities were dominated by Ascomycota. Our results fill a critical gap in our understanding of soil microbial communities in terrestrial ecosystems. They suggest that changes in aridity, such as those predicted by climate-change models, may reduce microbial abundance and diversity, a response that will likely impact the provision of key ecosystem services by global drylands.

Butterfly community shifts over two centuries.

Environmental changes strongly impact the distribution of species and subsequently the composition of species assemblages. Although most community ecology studies represent temporal snap shots, long-term observations are rather rare. However, only such time series allow the identification of species composition shifts over several decades or even centuries. We analyzed changes in the species composition of a southeastern German butterfly and burnet moth community over nearly 2 centuries (1840-2013). We classified all species observed over this period according to their ecological tolerance, thereby assessing their degree of habitat specialisation. This classification was based on traits of the butterfly and burnet moth species and on their larval host plants. We collected data on temperature and precipitation for our study area over the same period. The number of species declined substantially from 1840 (117 species) to 2013 (71 species). The proportion of habitat specialists decreased, and most of these are currently endangered. In contrast, the proportion of habitat generalists increased. Species with restricted dispersal behavior and species in need of areas poor in soil nutrients had severe losses. Furthermore, our data indicated a decrease in species composition similarity between different decades over time. These data on species composition changes and the general trends of modifications may reflect effects from climate change and atmospheric nitrogen loads, as indicated by the ecological characteristics of host plant species and local changes in habitat configuration with increasing fragmentation. Our observation of major declines over time of currently threatened and protected species shows the importance of efficient conservation strategies.

Environmental correlates of species rank - abundance distributions in global drylands.

Theoretical models predict lognormal species abundance distributions (SADs) in stable and productive environments, with log-series SADs in less stable, dispersal driven communities. We studied patterns of relative species abundances of perennial vascular plants in global dryland communities to: i) assess the influence of climatic and soil characteristics on the observed SADs, ii) infer how environmental variability influences relative abundances, and iii) evaluate how colonisation dynamics and environmental filters shape abundance distributions. We fitted lognormal and log-series SADs to 91 sites containing at least 15 species of perennial vascular plants. The dependence of species relative abundances on soil and climate variables was assessed using general linear models. Irrespective of habitat type and latitude, the majority of the SADs (70.3%) were best described by a lognormal distribution. Lognormal SADs were associated with low annual precipitation, higher aridity, high soil carbon content, and higher variability of climate variables and soil nitrate. Our results do not corroborate models predicting the prevalence of log-series SADs in dryland communities. As lognormal SADs were particularly associated with sites with drier conditions and a higher environmental variability, we reject models linking lognormality to environmental stability and high productivity conditions. Instead our results point to the prevalence of lognormal SADs in heterogeneous environments, allowing for more evenly distributed plant communities, or in stressful ecosystems, which are generally shaped by strong habitat filters and limited colonisation. This suggests that drylands may be resilient to environmental changes because the many species with intermediate relative abundances could take over ecosystem functioning if the environment becomes suboptimal for dominant species.

Intransitive competition is widespread in plant communities and maintains their species richness.

Intransitive competition networks, those in which there is no single best competitor, may ensure species coexistence. However, their frequency and importance in maintaining diversity in real-world ecosystems remain unclear. We used two large data sets from drylands and agricultural grasslands to assess: (1) the generality of intransitive competition, (2) intransitivity-richness relationships and (3) effects of two major drivers of biodiversity loss (aridity and land-use intensification) on intransitivity and species richness. Intransitive competition occurred in > 65% of sites and was associated with higher species richness. Intransitivity increased with aridity, partly buffering its negative effects on diversity, but was decreased by intensive land use, enhancing its negative effects on diversity. These contrasting responses likely arise because intransitivity is promoted by temporal heterogeneity, which is enhanced by aridity but may decline with land-use intensity. We show that intransitivity is widespread in nature and increases diversity, but it can be lost with environmental homogenisation.

Temporal patterns of energy equivalence in temperate soil invertebrates.

The question whether total population energy use is invariant to species body size (the energy equivalence hypothesis) is central to metabolic ecology and continues to be controversial. While recent comparative field work and meta-analyses pointed to systematic deviations of the underlying allometric scaling laws from predictions of metabolic theory none of these studies included the variability of metabolic scaling in ecological time. Here we used extensive data on the invertebrate soil fauna of Kampinos National Park (Poland) obtained from six consecutive quantitative sampling seasons to show that phylogenetically corrected species density-body weight and population energy use-body weight relationships across all soil fauna species and within trophic groups and body weight classes were highly variable in time. On average, population energy use tended to increase with species body weight in decomposers and phytophages, but not in predators. Despite these trends, our data do not exclude the possibility that energy equivalence marks the central tendency of energy use in the edaphon. Our results highlight the need for long-term studies on energy use to unequivocally assess predictions of metabolic theory.

Prevalence, virulence, and antimicrobial resistance of Campylobacter jejuni and Campylobacter coli in white stork Ciconia ciconia in Poland.

The aim of this study was to investigate the role of white stork Ciconia ciconia as a potential reservoir of Campylobacter spp. Antimicrobial resistance and the presence of putative virulence genes of the isolates were also examined. A total of 398 white stork chicks sampled in Western Poland in habitats with high density of breeding were examined. Rectal swabs were collected during breeding season 2009-2012 from storks developing in a relatively pure environment (Odra meadows), in polluted areas (a copper mining-smelting complex), and in suburbs. Of the anal swabs collected, 7.6% were positive for Campylobacter among chicks (5.3% samples positive for C. jejuni and 2.3% samples positive for C. coli). Samples from polluted areas had the highest prevalence of Campylobacter (12.2%). The prevalence of resistance among C. jejuni and C. coli isolates from young storks was as follows: to ciprofloxacin (52.4%, 44.4%), and to tetracycline (19%, 77.8%). All of the analyzed isolates were susceptible to macrolides. The resistance to both classes of antibiotics was found in the 23.3% of Campylobacter spp. All Campylobacter spp. isolates had cadF gene and flaA gene responsible for adherence and motility. CdtB gene associated with toxin production was present in 88.9% of C. coli isolates and 57.1% of C. jejuni isolates. The iam marker was found more often in C. coli strains (55.6%) compared to C. jejuni isolates (42.9%). Our results confirm the prevalence of Campylobacter spp. in the white stork in natural conditions and, because it lives in open farmlands with access to marshy wetlands, the environmental sources such as water reservoirs and soil-water can be contaminated from white stork feces and the pathogens can be widely disseminated. We can thus conclude that Campylobacter spp. may easily be transmitted to waterfowl, other birds, and humans via its environmental sources and/or by immediate contact.

The evolution of contact calls in isolated and overlapping populations of two white-eye congeners in East Africa (Aves, Zosterops).

BACKGROUND: Closely related species often occur in geographic isolation, yet sometimes form contact zones with the potential to hybridize. Pre-zygotic barriers may prevent cross breeding in such contact zones. In East Africa, White-eye birds have evolved into various species, inhabiting different habitat types. Zosterops poliogaster is found in cool and moist cloud forests at higher elevations, whereas Z. abyssinicus is distributed across the dry and hot lowland savannahs. In most areas, these two species occur allopatrically, but in the contact zone where the mountain meets the savannah, the distributions of these species sometimes overlap (parapatry), and in a few areas the two taxa occur sympatrically. Acoustic communication is thought to be an important species recognition mechanism in birds and an effective prezygotic barrier for hybridisation. We recorded contact calls of both the lowland and highland species in (i) distinct populations (allopatry), (ii) along contact zones (parapatry), and (iii) in overlapping populations (sympatry) to test for species and population differentiation. RESULTS: We found significant differences in call characteristics between the highland and lowland species, in addition to call differentiation within species. The highland Z. poliogaster shows a strong call differentiation among local populations, accompanied by comparatively low variability in their contact calls within populations (i.e. a small acoustic space). In contrast, calls of the lowland Z. abyssinicus are not differentiated among local sites but show relatively high variability in calls within single populations. Call patterns in both species show geographic clines in relation to latitude and longitude. Calls from parapatric populations from both species showed greater similarity to the other taxon in comparison to heterospecific populations found in allopatry. However, where the two species occur sympatrically, contact calls of both species are more distinct from each other than in either allopatric or parapatric populations. CONCLUSION: The contrasting patterns reflect divergent spatial distributions: the highland Z. poliogaster populations are highly disjunct, while Z. abyssinicus lowland populations are interconnected. Higher similarity in contact calls of heterospecific populations might be due to intermixing. In contrast, sympatric populations show reproductive character displacement which leads to strongly divergent call patterns.

Small-scale spatial variability in phylogenetic community structure during early plant succession depends on soil properties.

During early plant succession, the phylogenetic structure of a community changes in response to important environmental filters and emerging species interactions. We traced the development of temperate-zone plant communities during the first 7 years of primary succession on catchment soils to explore patterns of initial species assembly. We found pronounced small-scale differences in the phylogenetic composition of neighbouring plant assemblages and a large-scale trend towards phylogenetic evenness. This small-scale variability appears to be mediated by soil properties, particularly carbonate content. Therefore, abiotic environmental conditions might counteract or even supersede the effects of interspecific competition among closely related species, which are usually predicted to exhibit patterns of phylogenetic evenness. We conclude that theories on phylogenetic community composition need to incorporate effects of small-scale variability of environmental factors.

Out of the alps: the biogeography of a disjunctly distributed mountain butterfly, the almond-eyed ringlet Erebia alberganus (Lepidoptera, Satyrinae).

Many studies on the biogeography of thermophilic and arctic-alpine species were performed during the past. Only little is known about species with intermediate characteristics. We analyzed the molecular biogeography of the butterfly Erebia alberganus (30 populations, representing 1106 individuals), sampled over the Alps, Apennines (Italy), and the Stara Planina (Bulgaria) using allozyme electrophoresis (17 loci). Genetic analyses revealed 3 major splits, with the strongest between the Stara Planina populations and all other populations, and a weaker split between the Alps and the Apennines. Individuals from the Apennines were genetically nested within the Alps group. The Alps cluster was segregated into 3 groups: the Southwestern, Western/Central, and Eastern Alps. The genetic diversities were highest for the Alps populations and significantly lower in the 2 isolates (Apennines, Stara Planina). The remarkable genetic split between Stara Planina and all other populations and the genetic distinctiveness of the former cluster might be interpreted as an ancient colonization event of this Balkan mountain range. The Apennines populations derive from a more recent expansion out of the Southwestern Alps. After surviving the Würm ice age most probably in the central Apennines, accompanied by genetic modification of some of these populations, northward expansion might have started from the western parts of the central Apennines reaching the northern Apennines during the early postglacial. The subtle genetic differentiation found among the Alps populations probably reflects 3 geographically disjunct Würm glacial centers located at the western slopes of the Southwestern Alps, at the southern slopes of the Central Alps, and in the Southeastern Alps.

Species-driven phases and increasing structure in early-successional plant communities.

Successional phases describe changes in ecological communities that proceed in steps rather than continuously. Despite their importance for the understanding of ecosystem development, there still exists no reliable definition of phases and no quantitative measure of phase transitions. In order to obtain these data, we investigated primary succession in an artificial catchment (6 ha) in eastern Germany over a period of 6 years. The data set consists of records of plant species and their cover values, and initial substrate properties, both from plots in a regular grid (20 m × 20 m) suitable for spatial data analysis. Community assembly was studied by analyses of species co-occurrence and nestedness. Additionally, we correlated lognormal and log series distributions of species abundance to each community. We here introduce a new general method for detection of successional phases based on the degree of transient spatial homogeneity in the study system. Spatially coherent vegetation patterns revealed nonoverlapping partitions within this sequence of primary succession and were characterized as two distinct ecological phases. Patterns of species co-occurrence were increasingly less random, and hence the importance of demographic stochasticity and neutral community assembly decreased during the study period. Our findings highlight the spatial dimension of successional phases and quantify the degree of change between these steps. They are an element for advancing a more reliable terminology of ecological successions.

Butterfly species respond differently to climate warming and land use change in the northern Alps.

Climate change has a worldwide impact on biodiversity and ecosystem functions, in particular by causing shifts in species distributions and changes in species communities. Here, we analyse altitudinal range shifts of 30,604 lowland butterfly and burnet moth records from 119 species over the past seven decades across the federal state of Salzburg (northern Austria) spanning an altitudinal gradient of >2500 m. For each species, we compiled species-specific traits on their ecology, behaviour, and life-cycle. During the study period, the butterflies have shifted their average occurrence and also lower and upper occurrence limits >300 m uphill. This shift is particularly obvious for the last ten years. Habitat generalist and mobile species exhibited strongest and habitat specialist and sedentary species weakest shifts. Our results underline that the effects of climate change have a strong and currently increasing impact on the patterns of species distribution and local community composition. Hence, we confirm the observation that ubiquistic, mobile species with a broad ecological amplitude can cope better with environmental changes than specialist and sedentary species. Furthermore, the strong changes in land use in lowland areas might additionally enhanced this up-hill shift.