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.

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.

Fluctuating insect diversity, abundance and biomass across agricultural landscapes.

Habitat destruction and deterioration of habitat quality caused a severe decline of biodiversity, such as insect diversity. In this study, we analyze insect diversity and biomass across agro-environments. We collected flying insects with 20 malaise traps across a landscape mosaic consisting of organic (eight traps) and conventional (four traps) farmland, as well as across agricultural land that has been recently converted from conventional to organic farming (eight traps). Sampling was conducted over 2 years, in 2019 and 2020, with in total 340 sampling events. We measured the dry weight of the captured organisms and identified species diversity by analyzing Operational Taxonomic Units (OTUs) and Barcode Index Numbers (BINs) via metabarcoding. The results obtained show temporal dynamics. The number of OTUs were always higher than the number of BINs. OTUs and BINs were moderately to highly correlated, while the number of OTUs and BINs were only moderately positively correlated with dry biomass. OTUs and BINs as well as biomass were highest in the recently transformed farmland if compared with pure organic and conventional farmland sites, which showed no significant differences in respect of insect diversity. OTU and BIN numbers but not the OTU/BIN ratio significantly decreased with increasing distance from the nearest forest fringe. The numbers of OTUs, BINs and the OTU/BIN proportion, as well as OTU and BIN/biomass proportions varied strongly over seasons, irrespective of agricultural practice. Based on our findings, we suggest to combine data on insect species richness and biomass measured over a period of time, to derive a largely complete and meaningful assessment of biodiversity for a specific region.

Breakpoints in butterfly decline in Central Europe over the last century.

Recent studies indicated severe decline of insect diversity and abundance across major parts of Central Europe. Theoretical studies showed that the drivers behind biodiversity loss vary considerably over time. However, these scenarios so far have been insufficiently approved by long-term and large-scale data. In this study we analysed the temporal trends of butterflies and Zygaenid moths across the federal state of Salzburg, northern Austria, from 1920 to 2019. Our study area covers a large variety of habitats and altitudes. Various changes of land use and intensification occurred during and shortly before our studied period, with a first wave of habitat destruction starting in the late 19th century, followed by the deterioration of habitat quality since the mid-20th century. We used 59,870 presence-only data of 168 butterfly and burnet moth species. Each of these species was classified according to ecological characteristics. Break point analyses for non-linear temporal trends in the community composition returned two major time windows. These time windows coincide with periods characterized by severe habitat destruction and the deterioration of habitat quality due to agricultural intensification. We found significant reductions of the proportion of species requiring specific habitats since 1920 and until today. We identified additional break points for species requiring high habitat qualities, endangered butterfly species, and sedentary species, particularly after a main break point in the 1960s. Our findings underline that, apart from habitat destruction, the deterioration of habitat quality is a main driver of biodiversity loss in general. Therefore, nature conservation should focus on maintaining the highest possible habitat quality.

Dietary preference of cheetahs (Acinonyx jubatus) in south-eastern Kenya.

The conversion of natural ecosystems due to anthropogenic activities has led to the destruction of natural habitats and to the deterioration of habitat quality. Top predators particularly respond sensitively to changes in habitat structures, including the availability of prey. The cheetah Acinonyx jubatus prefers small-medium-sized, wild ungulate prey due to the cheetah's morphological adaptations. However, the majority of the species' population is found beyond protected areas, where habitat structures, species abundances, and community composition are highly influenced by human activities. Only few studies have analyzed the diet preference of cheetahs in relation to prey availability and abundance for rangelands beyond protected areas in Eastern Africa. The study aimed to determine cheetah prey preference in the rangelands of south-eastern Kenya based on scat analyses. We compared dietary preference of cheetah with prey availability. For this purpose, we conducted standardized game counts. We analyzed 27 cheetah scat samples collected across the same study area where we also conducted game counts. We found that Grant's gazelle Gazella granti contributed the highest portion of cheetah's diet, although Thomson's gazelle Gazella thomsonii was the most abundant medium-sized ungulate prey in the study areas. We also recorded two primate species, yellow baboon Papio cynocephalus and vervet monkey Chlorocebus pygerythrus, as well as the rock hyrax Procavia capensis in the cheetah diet. These species have never been documented as cheetah prey before. Furthermore, our results document livestock as potential prey for cheetahs. These observations underline that cheetah use diverse prey in rangelands outside protected areas, and that the abundance of specific prey does not influence cheetah prey preference.

Ecosystem functions in degraded riparian forests of southeastern Kenya.

Species community structures shape ecosystem functions, which are mostly stronger pronounced in intact than in degraded environments. Riparian forests in semiarid Africa provide important habitats for endangered plant and animal species and provide various ecosystem functions, that is, services to people settling along these streams. Most of these riparian forests are severely disturbed by human activities and dominated by invasive exotic plant species in the meanwhile. Thus, ecosystem functions are negatively influenced. While most studies have analyzed a specific metric to measure the degree of ecosystem function, little is known about how strongly different ecosystem functions respond to anthropogenic disturbances in parallel. In this study, we analyzed a set of four proxies of ecosystem functions, ground-dwelling arthropod abundances, pollination, seed dispersal, and predation, along a highly disturbed riparian forest in southeastern Kenya. We assessed the land cover and land use manually and with an Unmanned Aerial Vehicle. Our data show that ecosystem functions respond differently to vegetation cover, human disturbances, and the availability of the invasive exotic shrub Lantana camara. The occurrence of representatives from the groups Saltatoria and Formicidae profits from heterogeneous habitat structures and natural riparian forest, while representatives of the Araneae profit from high proportion of agricultural fields. In general, predation is higher in mixed land use and natural riparian forest, while pollination and seed dispersal showed no significant trend in regard on land coverage. Along with this, predation also increased with rising proportion of natural riparian forest, while the proportion of agricultural land negatively affects predation, but in parallel showed a slightly significant positive trend with seed dispersal. Human disturbances and the occurrence of the invasive exotic L. camara shrub did not significantly affect our metrics of ecosystem functioning, except of the negative impact of human disturbances on pollinators. In conclusion, our results underpin that ecosystem functions respond highly variable and individually to environmental changes.

Climate change drives mountain butterflies towards the summits.

Climate change impacts biodiversity and is driving range shifts of species and populations across the globe. To understand the effects of climate warming on biota, long-term observations of the occurrence of species and detailed knowledge on their ecology and life-history is crucial. Mountain species particularly suffer under climate warming and often respond to environmental changes by altitudinal range shifts. We assessed long-term distribution trends of mountain butterflies across the eastern Alps and calculated species' specific annual range shifts based on field observations and species distribution models, counterbalancing the potential drawbacks of both approaches. We also compiled details on the ecology, behaviour and life-history, and the climate niche of each species assessed. We found that the highest altitudinal maxima were observed recently in the majority of cases, while the lowest altitudes of observations were recorded before 1980. Mobile and generalist species with a broad ecological amplitude tended to move uphill more than specialist and sedentary species. As main drivers we identified climatic conditions and topographic variables, such as insolation and solar irradiation. This study provides important evidence for responses of high mountain taxa to rapid climate change. Our study underlines the advantage of combining historical surveys and museum collection data with cutting-edge analyses.

Seasonality and landscape characteristics impact species community structure and temporal dynamics of East African butterflies.

Species community structures respond strongly to habitat changes. These are either driven by nature or human activities. The biota of East African drylands responds highly sensitively to natural and anthropogenic impacts. Thus, seasonality strongly influences resource availability in a cyclic manner during the year, with cyclic appearance of the different developmental stages of invertebrates, while man-made landscape transformations profoundly and permanently modify habitat structures and, as a consequence, species communities. Butterflies are an excellent model group for the study of the effects of seasonality, and to test for biodiversity responses to anthropogenic activities such as habitat modification, degradation and destruction. We performed transect counts of adult butterflies in riparian forests and their adjoining areas, either dry savannahs with occasional pasturing (i.e. near-natural status) or farmland areas with fields, gardens and settlements (i.e. highly degraded status with lack of original vegetation). Transects were set along the river beds as well as at 250 m and 500 m distances parallel to these rivers, with eight transects per distance class and site (i.e. 48 transects in total). We recorded habitat structures for each transect. Counts were conducted during the dry and the rainy season, with 16 repetitions for each single transect, i.e. eight per season and transect. We compiled trait data on morphology, geographic distribution, ecology, behaviour, and life-history for all butterfly species encountered. Our results show higher species richness and numbers of individuals in farmland transects compared with the savannah region. Seasonal fluctuations of the detectable species abundances between the rainy and dry season were severe. These fluctuations were much more pronounced for the savannah than the farmland area, i.e. was buffered by human activities. Farmland and savannah support two distinct butterfly communities, with generalist species being more common in the farmland communities. Strict habitat associations were comparatively weak and typical dry savannah and riparian forest species were not clearly restricted to the near natural landscape.

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.

Toward a standardized quantitative and qualitative insect monitoring scheme.

The number of insect species and insect abundances decreased severely during the past decades over major parts of Central Europe. Previous studies documented declines of species richness, abundances, shifts in species composition, and decreasing biomass of flying insects. In this study, we present a standardized approach to quantitatively and qualitatively assess insect diversity, biomass, and the abundance of taxa, in parallel. We applied two methods: Malaise traps, and automated and active light trapping. Sampling was conducted from April to October 2018 in southern Germany, at four sites representing conventional and organic farming. Bulk samples obtained from Malaise traps were further analyzed using DNA metabarcoding. Larger moths (Macroheterocera) collected with light trapping were further classified according to their degree of endangerment. Our methods provide valuable quantitative and qualitative data. Our results indicate more biomass and higher species richness, as well as twice the number of Red List lepidopterans in organic farmland than in conventional farmland. This combination of sampling methods with subsequent DNA metabarcoding and assignments of individuals according depending on ecological characteristics and the degree of endangerment allows to evaluate the status of landscapes and represents a suitable setup for large-scale long-term insect monitoring across Central Europe, and elsewhere.

Long-term large-scale decline in relative abundances of butterfly and burnet moth species across south-western Germany.

Current studies have shown a severe general decline in insect species diversity, their abundance, and a biomass reduction of flying insects. Most of previous studies have been performed at single sites, or were spatially restricted at the landscape level. In this study, we analyse trends of species richness and shifts in species composition of butterflies and burnet moth species across the federal state of Baden-Württemberg in south-western Germany, covering an area of 35,750 km2. The data set consists of 233,474 records and covers a period from 1750 until today. We grouped species according to their species´ specific functional traits and analyse how species with different habitat requirements and behaviour respond to land-use changes over time. Our data document a significant loss of relative abundance for most species, especially since the 1950s until today. Species demanding specific habitat requirements are more seriously suffering under this trend than generalists. This in particular affects taxa adapted to extensively used xerothermic grasslands, bogs or other habitats maintained by traditional low-productivity agricultural practices of the past. Our data indicate large-scale decline in relative abundance of many butterfly and burnet moth species, which happened in particular during the past few decades.

Succession matters: Community shifts in moths over three decades increases multifunctionality in intermediate successional stages.

Species composition strongly depends on time, place and resources. In this context, semi-natural grasslands belong to the most species-rich habitats of Europe, and succession may eventually cause local extinction of typical grassland species, but conversely increase species richness due to habitat diversification. Here, we analyse potential effects of succession of calcareous grasslands on moths. Our studied community, assessed over three decades in south-eastern Germany, comprised >1000 species. We compiled data on the ecology of each of these species, considering the larval and adult stages. We assigned Ellenberg indicator values to each main larval food plant species used by these lepidopterans. Changes in the community means of these indicators were applied to test for possible consequences of the changes in habitat structure and quality. Our data revealed increasing multifunctionality of community structure, higher variability of habitat association over time, the appearance of range expanding species, but also local extinction of various typical grassland moth species. These shifts in species composition mirror effects of succession, which frequently transform previously homogenous semi-natural grasslands into a heterogeneous habitat mosaic.

Cross-continental phylogeography of two Holarctic Nymphalid butterflies, Boloria eunomia and Boloria selene.

Pleistocene glaciations had significant effects on the distribution and evolution of species inhabiting the Holarctic region. Phylogeographic studies concerning the entire region are still rare. Here, we compared global phylogeographic patterns of one boreo-montane and one boreo-temperate butterflies with largely overlapping distribution ranges across the Northern Hemisphere, but with different levels of range fragmentation and food specialization. We reconstructed the global phylogeographic history of the boreo-montane specialist Boloria eunomia (n = 223) and of the boreo-temperate generalist Boloria selene (n = 106) based on mitochondrial and nuclear DNA markers, and with species distribution modelling (SDM). According to the genetic structures obtained, both species show a Siberian origin and considerable split among populations from Nearctic and Palaearctic regions. According to SDMs and molecular data, both butterflies could inhabit vast areas during the moderate glacials. In the case of B. selene, high haplotype diversity and low geographic structure suggest long-lasting interconnected gene flow among populations. A stronger geographic structuring between populations was identified in the specialist B. eunomia, presumably due to the less widespread, heterogeneously distributed food resources, associated with cooler and more humid climatic conditions. Populations of both species show opposite patterns across major parts of North America and in the case of B. eunomia also across Asia. Our data underline the relevance to cover entire distribution ranges to reconstruct the correct phylogeographic history of species.

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.

Neutral colonisations drive high beta-diversity in cavernicole springtails (Collembola).

The theory of island biogeography predicts the effects of habitat isolation and size on species richness, community assembly, and the persistence of species. Various studies showed that habitat conditions and the ecology of species are also of relevance in explaining community assembly. Geographically isolated habitats like caves with rather constant environmental conditions provide models to test for the relevance of the above described variables. In this study we analysed springtails living in karst caves of the Romanian Carpathians and Dobrogea region. We considered phylogenetic relatedness, habitat and species characteristics to identify the relevant drivers of community assembly. Our data show that species richness of single caves is low. Neither phylogenetic relatedness nor habitat filtering and competitive interactions seem to shape species composition or to affect species richness. We found that glacial-interglacial cycles with subsequent range contractions and expansions might have led to independent and multiple colonisations of caves. Furthermore, single caves might have acted as refugia and thus might have provided the prerequisite for distinct evolution processes, leading to a high level of endemicity of these animal species.

A new amplicon based approach of whole mitogenome sequencing for phylogenetic and phylogeographic analysis: An example of East African white-eyes (Aves, Zosteropidae).

Classical Sanger sequencing is still frequently used to generate sequence data for phylogenetic and phylogeographic inference. In this contribution we present a novel approach to genotype whole mitogenomic haplotypes using Illumina MiSeq reads from indexed amplicons. Our new approach reduces preparation time by multiplexing loci within a single or few PCR reactions and by plate format library construction. The use of paired-end reads allows covering amplicons of about 0.5kb and thus no nebulisation and assembly are necessary. We tested the power and effectiveness of this technique by analysing the mitogenomic diversity of East African white-eye bird species (Zosteropidae), a taxonomically highly diverse and complex species flock found in various ecosystems spread across major parts of Africa. We compare the newly generated mitogenomic data set with published data of three mitochondrial genes for a similar set of populations and taxa. The comparison demonstrates that our new procedure represents a cost effective use of NGS for medium throughput phylogenetic analyses. Using this method, we were able to increase the amount of phylogenetic information significantly, while reducing the costs and effort in the laboratory. The mitogenomic data show a higher resolution than previous studies providing higher support and new insights in the relationships of Zosterops species. Our data suggest to split Z. poliogaster into four distinct species, three of which had previously been proposed: Z. silvanus, Z. mbulensis, Z. kikyuensis and Z. kulalensis. Our approach allows the genotyping of whole mitogenomes for a large number of individuals and thus allows more reliable reconstruction of phylogenetic and phylogeographic relationships - also for non-model organisms.

Kenyan endemic bird species at home in novel ecosystem.

Riparian thickets of East Africa harbor a large number of endemic animal and plant species, but also provide important ecosystem services for the human being settling along streams. This creates a conflicting situation between nature conservation and land-use activities. Today, most of this former pristine vegetation is highly degraded and became replaced by the invasive exotic Lantana camara shrub species. In this study, we analyze the movement behavior and habitat use of a diverse range of riparian bird species and model the habitat availability of each of these species. We selected the following four riparian bird species: Bare-eyed Thrush Turdus tephronotus, Rufous Chatterer Turdoides rubiginosus, Zanzibar Sombre Greenbul Andropadus importunus insularis, and the Kenyan endemic Hinde's Babbler Turdoides hindei. We collected telemetric data of 14 individuals during a 2 months radio-tracking campaign along the Nzeeu River in southeast Kenya. We found that (1) all four species had similar home-range sizes, all geographically restricted and nearby the river; (2) all species mainly use dense thicket, in particular the invasive L. camara; (3) human settlements were avoided by the bird individuals observed; (4) the birds' movement, indicating foraging behavior, was comparatively slow within thickets, but significantly faster over open, agricultural areas; and (5) habitat suitability models underline the relevance of L. camara as suitable surrogate habitat for all understoreyed bird species, but also show that the clearance of thickets has led to a vanishing of large and interconnected thickets and thus might have negative effects on the population viability in the long run.

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.