Butterfly habitats, broad-scale biotope affiliations, and structural exploitation of vegetation at finer scales: the matrix revisited

Abstract. 1. Biotope and resources data are rarely attached to arthropod (butterfly) synoptic monitoring systems, and invariably not linked to behavioural exploitation of vegetation substrates. Yet, these data allow us to examine resource use within different biotopes and to distinguish more clearly between habitat and the matrix. 2. Comparative data on vegetation exploitation for different behaviours (search flight, direct flight, tactile inspections, perching, feeding, interactions, oviposition) were collected using transect sections over a range of biotopes from bare ground to mature woodland for two closely related satyrine butterflies, Maniola jurtina and Pyronia tithonus with overlapping flight times. Occupancy data were obtained on Pararge aegeria as a marker for the woodland end of the biotope spectrum. 3. There were clear distinctions in biotope occupancy between M. jurtina (grassland bias) and P. tithonus (shrub bias); significant differences in exploitation of vegetation substrates (except for nectar feeding) coincided with this bias in transect sections which comprise both grassland and shrubs. The exception (nectar feeding) is explained by the decline in shrub (Rubus fruticosus) nectar and increase in herb nectar during the later emergence of P. tithonus. 4. Direct flight increased in unsuitable biotopes for both species. However, resource‐exploiting behaviour (>70%) predominated even in biotopes that would be regarded as completely unsuitable for supporting the species and where less than 2% of individuals for each species were observed. 5. Simultaneous collection of biotope, resources, and behavioural data is needed for monitoring affinities of butterflies to vegetation structures and using butterflies as indicators of environmental changes. 6. Much of the landscape is shown to comprise valuable resources for butterflies, even when classified for metapopulation studies as empty matrix.     

Butterfly monitoring in Europe: methods,applications and perspectives

Since the first Butterfly Monitoring Scheme in the UK started in the mid-1970s, butterfly monitoring in Europe has developed in more than ten European countries. These schemes are aimed to assess regional and national trends in butterfly abundance per species. We discuss strengths and weaknesses of methods used in these schemes and give examples of applications of the data. A new development is to establish supra-national trends per species and multispecies indicators. Such indicators enable to report against the target to halt biodiversity loss by 2010. Our preliminary European Grassland Butterfly Indicator shows a decline of 50% between 1990 and 2005. We expect to develop a Grassland Butterfly Indicator with an improved coverage across European countries. We see also good perspectives to develop a supra-national indicator for climate change as well as an indicator for woodland butterflies.     

Biodiversity trends in Europe: development and testing of a species trend indicator for evaluating progress towards the 2010 target

This paper presents a trial of a species population trend indicator for evaluating progress towards the 2010 biodiversity target in Europe, using existing data. The indicator integrates trends on different species (groups), and can be aggregated across habitats and countries. Thus, the indicator can deliver both headline messages for high-level decision-making and detailed information for in-depth analysis, using data from different sources, collected with different methods. International non-governmental organizations mobilized data on over 2800 historical trends in national populations of birds, butterflies and mammals, for a total of 273 species. These were combined by habitat and biogeographical region to generate a pilot pan-European scale indicator. The trial indicator suggests a decline of species populations in nearly all habitats, the largest being in farmland, where species populations declined by an average of 23% between 1970 and 2000. The indicator is potentially useful for monitoring progress towards 2010 biodiversity targets, but constraints include: the limited sensitivity of the historical data, which leads to conservative estimates of species decline; a potential danger of ambiguity because increases in opportunistic species can mask the loss of other species; and failure to account for pre-1970 population declines. We recommend mobilizing additional existing data, particularly for plants and fishes, and elaborating further the criteria for compiling representative sets of species. For a frequent, reliable update of the indicator, sound, sensitive and harmonized biodiversity monitoring programmes are needed in all pan-European countries.     

Contrasting changes in the abundance and diversity of North American bird assemblages from 1971 to 2010

Although it is generally recognized that global biodiversity is declining, few studies have examined long‐term changes in multiple biodiversity dimensions simultaneously. In this study, we quantified and compared temporal changes in the abundance, taxonomic diversity, functional diversity, and phylogenetic diversity of bird assemblages, using roadside monitoring data of the North American Breeding Bird Survey from 1971 to 2010. We calculated 12 abundance and diversity metrics based on 5‐year average abundances of 519 species for each of 768 monitoring routes. We did this for all bird species together as well as for four subgroups based on breeding habitat affinity (grassland, woodland, wetland, and shrubland breeders). The majority of the biodiversity metrics increased or remained constant over the study period, whereas the overall abundance of birds showed a pronounced decrease, primarily driven by declines of the most abundant species. These results highlight how stable or even increasing metrics of taxonomic, functional, or phylogenetic diversity may occur in parallel with substantial losses of individuals. We further found that patterns of change differed among the species subgroups, with both abundance and diversity increasing for woodland birds and decreasing for grassland breeders. The contrasting changes between abundance and diversity and among the breeding habitat groups underscore the relevance of a multifaceted approach to measuring biodiversity change. Our findings further stress the importance of monitoring the overall abundance of individuals in addition to metrics of taxonomic, functional, or phylogenetic diversity, thus confirming the importance of population abundance as an essential biodiversity variable.     

Tiny terminological disagreements with far reaching consequences for global bird trends

Various combinations of data and expert opinion have been used to select species for indices of bird trends. Commonly these indices break species into groups based on their habitat preference such as woodland specialist, farmland specialist and generalist birds. It is unclear what influence differences in how species are allocated to these groups might have on trends in these indices. There is uncertainty surrounding reported trends in these bird groups with studies variously showing declines or increases in prevalence. This is usually attributed to ecological factors but if studies classify bird groups differently this variation may be due to inconsistency in classification. Disagreement about whether these bird groups are stable, increasing or declining has the potential to obscure important changes in bird prevalence and impede appropriate, timely conservation.We examined how consistently European and Australian researchers classified woodland, farmland and generalist birds, and whether this affected the trends in indices of these groups. Researchers from both regions classified species differently, and the population trends seen in these groups were strongly affected by differences in classification. While all classifications we studied suggest that populations are consistently declining for Australian woodland and European farmland birds and increasing for European woodland birds. European generalist and Australian farmland and generalist birds may be seen as increasing or decreasing in prevalence depending on classification.Our results question the current practice of idiosyncratically classifying indicators in scientific research and conservation. Current practice is making it more difficult to infer whether, when and how to preserve bird groups in Europe and Australia, potentially leading to sub-optimal biodiversity outcomes. We offer suggestions for building consensus on how to classify these bird groups in order to provide more reliable evidence to support conservation decisions.     

Grassland management for meadow birds in the Netherlands is unfavourable to pollinators

Agricultural intensification and loss of semi-natural grassland have contributed to biodiversity decline, including pollinator species, in pastures around the world. To reverse the decline, agri-environmental schemes have been implemented, varying widely in effectiveness. In addition, many countries, including the Netherlands, have established nature reserves in which semi-natural grasslands are restored and are often managed for specific groups of species, e.g. meadow birds or plants. The effects of such measures on insect biodiversity are not well known but recent reports on the dramatic decline of insect biomass in nature reserves have put even more attention to the impact of land use and management on biodiversity. This study compares pollinator abundance and species richness in three common semi-natural grassland management types in the Netherlands: (1) hay meadows, (2) herb-rich grasslands and (3) meadow bird grasslands. Pollinator abundance and species richness were assessed in eleven study areas, each with all three management types present. Standardized transects, insect sampling within a standard 20 min time frame and plot-based flower surveys were used in spring and summer to assess the relationships between management regime, floral abundance and diversity and pollinator communities. The results show that meadow bird grasslands have lower pollinator abundance and diversity and a less unique pollinator assemblage than both other types. Moreover, flower abundance has a positive effect on pollinator abundance and flower diversity has a positive effect on pollinator species richness. These results indicate that meadow-bird grasslands are a comparatively unfavourable habitat for bees, hoverflies and butterflies, which may be explained by a lack of flowers as well as unsuitable mowing practices. Measures benefitting both insectivorous birds and flower-visiting insects, such as rotational mowing, could remediate this imbalance.     

Evaluating butterflies as surrogates for birds and plants in semi-natural grassland buffers

Semi-natural grasslands can support diverse faunal and floral communities, including grassland birds, beneficial insects, and native wildflowers. Monitoring biodiversity of this type of ecosystem is important to assess abundance and richness of grassland-associated species, evaluate success of establishing grasslands, and to assess overall ecosystem health. We tested butterflies as surrogates for birds and plants to assess establishment success of semi-natural grassland buffers in north-central Mississippi using Spearman rank correlation (Spearman’s ρ). Disturbance and grassland butterfly guilds were generally not suitable surrogates for grassland bird metrics, non-grassland bird metrics, or nest density metrics. Butterflies did have consistent positive correlations with plant species richness and forb metrics, as well as consistent negative correlations with grass metrics, but these correlations were generally smaller than what is considered suitable to serve as surrogates. In general, butterflies were not suitable surrogates for birds or plants in semi-natural grassland buffers.     

Primary productivity and species richness: relationships among functional guilds, residency groups and vagility classes at multiple spatial scales

One of the major determinants of species richness is the amount of energy available, often measured as primary productivity. Heterogeneity of environmental variables has also been found to influence species richness. Predicting species distributions across landscapes and identifying areas that have high species richness, or vulnerable groups of species, is useful for land management. Remotely sensed data may help identify such areas, with the Normalized Difference Vegetation Index (NDVI) providing an estimate of primary productivity. We examined the relationship between maximum productivity (NDVI), heterogeneity of productivity, and species richness of birds and butterflies at multiple spatial scales. We also explored relationships between productivity, functional guilds and residency groups of birds, and vagility classes of butterflies. Positive linear relationships between maximum NDVI and number of functional guilds of birds were found at two spatial scales. We also found positive linear relationships between maximum NDVI and species richness of neotropical migrant birds at two scales. Heterogeneity of NDVI, by contrast, was negatively associated with number of functional guilds of birds and species richness of resident birds. Maximum NDVI was associated with species richness of all butterflies and of the most vagile butterflies. No association was found between heterogeneity of NDVI and species richness of butterflies. In the Great Basin, where high greenness and availability of water correspond to areas of high species richness and maximum NDVI, our results suggest that NDVI can provide a reliable basis for stratifying surveys of biodiversity, by highlighting areas of potentially high biodiversity across large areas. Measures of heterogeneity of NDVI appear to be less useful in explaining species richness.     

Assessing the Sensitivity of Seabed Species and Biotopes – The Marine Life Information Network (MarLIN)

Science-based approaches to support the conservation of marine biodiversity have been developed in recent years. They include measures of ‘rarity’, ‘diversity’, ‘importance’, biological indicators of water ‘quality’ and measures of ‘sensitivity’. Identifying the sensitivity of species and biotopes, the main topic of this contribution, relies on accessing and interpreting available scientific data in a structured way and then making use of information technology to disseminate suitably presented information to decision makers. The Marine Life Information Network (MarLIN) has achieved that research for a range of environmentally critical species and biotopes over the past four years and has published the reviews on the MarLIN Web site (www.marlin.ac.uk). Now, by linking the sensitivity database and databases of survey information, sensitivity mapping approaches using GIS are being developed. The methods used to assess sensitivity are described and the approach is advocated for wider application in Europe.     

Importance of boreal grasslands in Sweden for butterfly diversity and effects of local and landscape habitat factors

A widespread decline in biodiversity in agro-ecosystems has been reported for several groups of organisms in Western Europe. The butterfly fauna was studied in 60 selected semi-natural grasslands in a coniferous-dominated boreal landscape in south-eastern Sweden. The aim was to investigate how butterfly assemblages were affected by the amount of semi-natural grasslands in the surrounding landscape. Furthermore, we wanted to determine if semi-natural grasslands in boreal landscapes harboured species otherwise declining in other parts of Europe. For each study site, the amounts of semi-natural grasslands in the landscape within radii of 500, 2,000 and 5,000 m were studied. Nine local habitat factors were also recorded. Only the amount of semi-natural grasslands within a 5,000 m radius could explain a significant part of the variation in butterfly composition, but there was no clear relationship between the amount of semi-natural grassland and butterfly diversity. Instead, this study showed that local habitat quality was very important for butterfly diversity at individual sites. Flower abundance, sward height and herb composition were the most important local factors. Patches surrounded by a small amount of semi-natural grasslands had high butterfly diversity, contrary to expectations. This may be explained by the fact that forest habitat provides a matrix with several features suitable for butterflies. The butterfly fauna was rich in species representative of low-productivity grasslands, species that are declining in other countries in Western Europe.     

Biotope prioritisation in the Central Apennines (Italy): species rarity and cross-taxon congruence

The conservation status of invertebrates is usually lesser known than that of vertebrates, and strategies to identify biotopes to preserve invertebrate diversity are typically based on a single surrogate taxon, or even on the use of vertebrates as surrogates. Aim of this research is to illustrate a method for biotope prioritisation that can be easily adapted to different animal groups and geographical contexts. A two-step protocol for biotope prioritisation is proposed on the basis of a multidimensional characterisation of species vulnerability. Firstly, species vulnerability is estimated from rarity measures which include geographical range, abundance and biotope specialisation. Then, these values of vulnerability are used to rank biotopes. The method was applied here to the tenebrionid beetles, the butterflies, the birds and the mammals of the Central Apennines, a montane area of high conservation concern for South Europe. This study provides evidence for the importance of including insects in conservation decisions, because vertebrates are poor surrogates for insects. Conservation efforts in the reserves included in the study area are mostly focused on vertebrates, for which woodlands are considered particularly important. However high altitude open biotopes are crucial for both tenebrionids and butterflies, and preservation of such kind of biotopes would be beneficial also for vertebrates. The approach applied here demonstrates that (1) vertebrates are poor surrogates for insects, and (2) measures of species rarity, typically used in vertebrate conservation, can be obtained also for insects, for which a veritable amount of data are hidden in specialised literature and museum collections.     

The conservation of declining butterfly populations in Britain and Europe: priorities, problems and successes

The status, ecology and conservation of butterflies in Europe and Britain are reviewed, as a background to the National Trust's past and future contribution to British conservation. Britain has a poor butterfly fauna by European standards, the main areas of endemism and species richness being in the Alps and southern Europe. To date, the main declines among European butterfly populations have occurred across central-northern Europe, with slightly higher extinction rates in mainland countries than in Britain. The main causes of decline are biotope destruction, the loss of certain species' habitats within surviving semi-natural biotopes due to changed land management, and a failure by several species to track the patches of their habitat that are still being generated in modern fragmented landscapes. Until recently, most conservation programmes failed to take account of the latter two factors, resulting in many local extinctions of rare butterfly species even in conservation areas. Recent measures have been much more successful; many were first tested on National Trust properties.     

Patches of bare ground as a staple commodity for declining ground-foraging insectivorous farmland birds

Conceived to combat widescale biodiversity erosion in farmland, agri-environment schemes have largely failed to deliver their promises despite massive financial support. While several common species have shown to react positively to existing measures, rare species have continued to decline in most European countries. Of particular concern is the status of insectivorous farmland birds that forage on the ground. We modelled the foraging habitat preferences of four declining insectivorous bird species (hoopoe, wryneck, woodlark, common redstart) inhabiting fruit tree plantations, orchards and vineyards. All species preferred foraging in habitat mosaics consisting of patches of grass and bare ground, with an optimal, species-specific bare ground coverage of 30-70% at the foraging patch scale. In the study areas, birds thrived in intensively cultivated farmland where such ground vegetation mosaics existed. Not promoted by conventional agri-environment schemes until now, patches of bare ground should be implemented throughout grassland in order to prevent further decline of insectivorous farmland birds.     

The role of linear mown firebreaks in conserving butterfly diversity: Effects of adjacent vegetation and management

Semi‐natural grasslands are vital for maintaining grassland butterflies in Japan, as well as in Europe. However, severe decline in these grassland environments has recently attracted attention to linear grasslands, such as firebreaks and power‐line corridors, as alternative habitats for grassland insects. We surveyed butterflies in an abandoned grassland and nearby linear mown firebreaks adjacent to different vegetation at the northern foot of Mt. Fuji, central Japan, over 5 successive years, particularly focusing on species on the 2012 Japanese Red List of Threatened Species (“red‐list” species). We found that the firebreaks were consistently higher in species richness and abundance of butterflies than the long‐term abandoned grassland, and that species composition differed among the firebreaks depending on conditions of the adjacent vegetation. The firebreaks surrounded by forests were mainly utilized by forest and edge species, whereas the firebreaks adjacent to the grassland were essential for conserving red‐list grassland species. Thus, only the mown firebreaks adjacent to the grassland were regarded as a high‐quality alternative habitat for many grassland butterflies, but the area was limited. Therefore, creating heterogeneity in the abandoned grassland by infrequent mowing could help conserve grassland butterflies, including red‐list species.     

Climate,landscape, habitat,and woodland management associations with hazel dormouse Muscardinus avellanarius population status

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<Emphasis Type="Italic">Polylepis</Emphasis> woodland remnants as biodiversity islands in the Bolivian high Andes

Mountain forests deserve special attention from ecologists and conservation biologists given the ecosystem services they provide to society, and their threat under global change. In the subalpine region of the Andes, Polylepis woodlands occur as arboreal islands in a matrix of grassland and scrub. Due to overgrazing and burning, however, these woodland patches are believed to cover only 11% of their potential area in Bolivia, core area for Polylepis. We reviewed the knowledge on the species diversity for the Bolivian Polylepis woodland remnants, assessed the conservation status of the occurring species, determined their trophic niche, and related species richness with climatic variables and elevation. Based in 31 publications, we found 780 identified species occurring in Polylepis woodlands: 425 plants, 266 birds, 46 mammals, 35 butterflies and 8 reptiles. Ten of the 13 Bolivian Polylepis species, as well as 7 other plant species, 14 bird species and 4 mammal species were categorized as threatened or near threatened according to IUCN criteria. In general, plant species richness increased with increased precipitation and length of the growth season, while it decreased with increasing elevation. There was a positive relationship between bird species richness, precipitation and length of the growth season. The highest bird endemism in Polylepis woodland remnants occurred at intermediate elevations, temperatures and precipitation. Mammal species richness decreased with increasing maximum temperature. Finally, we discuss the most important knowledge gaps regarding biodiversity in Bolivian Polylepis woodland remnants.     

More losers than winners: investigating Anthropocene defaunation through the diversity of population trends

The global-scale decline of animal biodiversity (‘defaunation’) represents one of the most alarming consequences of human impacts on the planet. The quantification of this extinction crisis has traditionally relied on the use of IUCN Red List conservation categories assigned to each assessed species. This approach reveals that a quarter of the world's animal species are currently threatened with extinction, and ~1% have been declared extinct. However, extinctions are preceded by progressive population declines through time that leave demographic ‘footprints’ that can alert us about the trajectories of species towards extinction. Therefore, an exclusive focus on IUCN conservation categories, without consideration of dynamic population trends, may underestimate the true extent of the processes of ongoing extinctions across nature. In fact, emerging evidence (e.g. the Living Planet Report), reveals a widespread tendency for sustained demographic declines (an average 69% decline in population abundances) of species globally. Yet, animal species are not only declining. Many species worldwide exhibit stable populations, while others are even thriving. Here, using population trend data for >71,000 animal species spanning all five groups of vertebrates (mammals, birds, reptiles, amphibians and fishes) and insects, we provide a comprehensive global-scale assessment of the diversity of population trends across species undergoing not only declines, but also population stability and increases. We show a widespread global erosion of species, with 48% undergoing declines, while 49% and 3% of species currently remain stable or are increasing, respectively. Geographically, we reveal an intriguing pattern similar to that of threatened species, whereby declines tend to concentrate around tropical regions, whereas stability and increases show a tendency to expand towards temperate climates. Importantly, we find that for species currently classed by the IUCN Red List as ‘non-threatened’, 33% are declining. Critically, in contrast with previous mass extinction events, our assessment shows that the Anthropocene extinction crisis is undergoing a rapid biodiversity imbalance, with levels of declines (a symptom of extinction) greatly exceeding levels of increases (a symptom of ecological expansion and potentially of evolution) for all groups. Our study contributes a further signal indicating that global biodiversity is entering a mass extinction, with ecosystem heterogeneity and functioning, biodiversity persistence, and human well-being under increasing threat.     

Developing indicators for European birds

The global pledge to deliver 'a significant reduction in the current rate of biodiversity loss by 2010' is echoed in a number of regional and national level targets. There is broad consensus, however, that in the absence of conservation action, biodiversity will continue to be lost at a rate unprecedented in the recent era. Remarkably, we lack a basic system to measure progress towards these targets and, in particular, we lack standard measures of biodiversity and procedures to construct and assess summary statistics. Here, we develop a simple classification of biodiversity indicators to assist their development and clarify purpose. We use European birds, as example taxa, to show how robust indicators can be constructed and how they can be interpreted. We have developed statistical methods to calculate supranational, multi-species indices using population data from national annual breeding bird surveys in Europe. Skilled volunteers using standardized field methods undertake data collection where methods and survey designs differ slightly across countries. Survey plots tend to be widely distributed at a national level, covering many bird species and habitats with reasonable representation. National species' indices are calculated using log-linear regression, which allows for plot turnover. Supranational species' indices are constructed by combining the national species' indices weighted by national population sizes of each species. Supranational, multi-species indicators are calculated by averaging the resulting indices. We show that common farmland birds in Europe have declined steeply over the last two decades, whereas woodland birds have not. Evidence elsewhere shows that the main driver of farmland bird declines is increased agricultural intensification. We argue that the farmland bird indicator is a useful surrogate for trends in other elements of biodiversity in this habitat.     

蝴蝶在生境和生物小区之间的移位伴随着对外来蜜源的利用

在发育成熟的温带环境中,植食性昆虫群落能迅速适应引入植物并将其作为幼虫的食物和花蜜.我们研究了经过森林砍伐的热带环境中蝴蝶对利用引入植物作为蜜源植物的适应快慢程度,并研究了蝴蝶-显花植物在新的生物小区中出现的范围,发现蝴蝶对引入显花植物的利用和探访多于本地植物,这与引入植物在调查地点、生物小区和植物丰度中的普遍性有关.此外,取食花蜜的蝴蝶和显花植物与正在形成的的生物小区有关,例如路边、农田、集约耕地以及花园.在这些新生物小区中,引入植物很重要,因为它们为蝴蝶提供了蜜源.