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At large scales, the mechanisms underpinning stability in natural communities may vary in importance due to changes in species composition, mean abundance, and species richness. Here we link species characteristics (niche positions) and community characteristics (richness and abundance) to evaluate the importance of stability mechanisms in 156 butterfly communities monitored across three European countries and spanning five bioclimatic regions. We construct niche-based hierarchical structural Bayesian models to explain first differences in abundance, population stability, and species richness between the countries, and then explore how these factors impact community stability both directly and indirectly (via synchrony and population stability). Species richness was partially explained by the position of a site relative to the niches of the species pool, and species near the centre of their niche had higher average population stability. The differences in mean abundance, population stability, and species richness then influenced how much variation in community stability they explained across the countries. We found, using variance partitioning, that community stability in Finnish communities was most influenced by community abundance, whereas this aspect was unimportant in Spain with species synchrony explaining most variation; the UK was somewhat intermediate with both factors explaining variation. Across all countries, the diversity–stability relationship was indirect with species richness reducing synchrony which increased community stability, with no direct effects of species richness. Our results suggest that in natural communities, biogeographical variation observed in key drivers of stability, such as population abundance and species richness, leads to community stability being limited by different factors and that this can partially be explained due to the niche characteristics of the European butterfly assemblage.

Species are altering their ranges as a response to climate change, but the magnitude and direction of observed range shifts vary considerably among species. The ability to persist in current areas and colonize new areas plays a crucial role in determining which species will thrive and which decline as climate change progresses. Several studies have sought to identify characteristics, such as morphological and life-history traits, that could explain differences in the capability of species to shift their ranges together with a changing climate. These characteristics have explained variation in range shifts only sporadically, thus offering an uncertain tool for discerning responses among species. As long-term selection to past climates have shaped species’ tolerances, metrics describing species’ contemporary climatic niches may provide an alternative means for understanding responses to on-going climate change. Species that occur in a broader range of climatic conditions may hold greater tolerance to climatic variability and could therefore more readily maintain their historical ranges, while species with more narrow tolerances may only persist if they are able to shift in space to track their climatic niche. Here, we provide a first-filter test of the effect of climatic niche dimensions on shifts in the leading range edges in three relatively well-dispersing species groups. Based on the realized changes in the northern range edges of 383 moth, butterfly, and bird species across a boreal 1,100 km latitudinal gradient over c. 20 years, we show that while most morphological or life-history traits were not strongly connected with range shifts, moths and birds occupying a narrower thermal niche and butterflies occupying a broader moisture niche across their European distribution show stronger shifts towards the north. Our results indicate that the climatic niche may be important for predicting responses under climate change and as such warrants further investigation of potential mechanistic underpinnings.

Aim: It is important to understand the factors affecting community stability because ecosystem function is increasingly at risk from biodiversity loss. Here, we evaluate how a key factor, the position of local environmental conditions within the thermal range of the species, influences the stability of butterfly communities at a continental scale.Location: Spain, UK and Finland.Time period: 1999– 2017.Major taxa studied: Butterflies.Methods: We tested the following hypotheses about how species responses to tem-perature anomalies aggregate to influence stability: Hypothesis 1, species have con-trasting responses to local temperature anomalies at opposing edges of their thermal range; hypothesis 2, communities with central thermal range positions have higher community stability; and the impacts of thermal range position on community stabil-ity are driven by hypothesis 3, population asynchrony, or hypothesis 4, additive popu-lation stability. Data were analysed at 876 sites for 157 species.Results: We found some support for hypothesis 1, because there were interactions be-tween thermal range and response to temperature anomalies such that species at dif-ferent range edges could provide weak compensatory dynamics. However, responses were nonlinear, suggesting strong declines with extreme anomalies, particularly at the hot range edge. Hypothesis 2 was supported in part, because community stabil-ity increased with central thermal range positions and declined at the edges, after accounting for species richness and community abundance. Thermal range position was weakly correlated with asynchrony (hypothesis 3) and population stability (hy-pothesis 4), although species richness and population abundance had larger impacts. Main conclusions: Future extreme heat events will be likely to impact species nega-tively across their thermal range, but might be particularly impactful on populations at the hottest end of the thermal range. Thermal range position influenced community stability because range edge communities were stable. However, the prediction of community stability from thermal range position is challenging because of nonlinear responses to temperature, with small temperature anomalies producing weak com-pensatory dynamics, but large extreme events synchronizing dynamics. The UK butterfly monitoring scheme is organized and funded by Butterfly Conservation, the Centre for Ecology and Hydrology, British Trust for Ornithology and the Joint Nature Conservation Committee. The Catalan butterfly monitoring scheme is funded by the Catalan Government, the Barcelona Provincial Council and other local partners. The Catalan butterfly monitoring scheme also incorpo-rates the Andora butterfly monitoring scheme, which is run by Centre d'Estudis de la Neu i la Mutanya d'Andorra (CENMA) and funded by Govern d'Andorra. The Finnish butterfly monitoring scheme is organ-ized and funded by the Finnish Environment Institute (SYKE).

Non-bee insects have been identified as important crop pollinators globally. However, strategies to protect pollinators and enhance crop pollination usually focus on supporting bees. This study examined the effects of landscape structure, location within field, and floral resources on pollinators’ visits on mass-flowering caraway (Carum carvi L.) in boreal farmland, and the effects of the visits on caraway yield. Pollinator visits on caraway flowers were monitored and caraway yield measured in 30 fields at landscapes ranging from field-dominated to forest-dominated landscapes. Hoverflies were the most abundant flower-visitors of caraway, followed by honeybees. Hoverflies and other flies made more flower visits on caraway than all bee species combined. Pollinator groups differed in their responses to landscape and local factors. Flies were most abundant near field edges and in landscapes with high forest cover. Non-syrphid flies and solitary bees responded positively to the cover of flowering herbs in the adjacent field margins. Flower visits by honeybees, instead, were positively related to the flowering crop cover in the study fields. Caraway seed yield increased with increasing number of flower visits by honeybees, hoverflies and all pollinators together. Pollinator exclusion reduced caraway fruit set (i.e. the number of fruits per flower) by 13% and seed yield by 40%. Our study is the first to report the high importance of flies to crop pollination in boreal farmland, where caraway is an important export crop. The results highlight the need of taking flies and their habitat requirements into account when developing strategies to enhance crop pollination. Highlights • Hoverflies were the most frequent flower-visitors on caraway, followed by honeybees. • Flies made more flower visits than all bee species combined. • Flies were abundant near field edges and in landscapes with high forest cover. • Honeybees responded positively to the flowering crop cover. • Flower visits by both honeybees and hoverflies increased caraway yield.

Climate change is a pervasive threat to biodiversity. While range shifts are a known consequence of climate warming contributing to regional community change, less is known about how species' positions shift within their climatic niches. Furthermore, whether the relative importance of different climatic variables prompting such shifts varies with changing climate remains unclear. Here we analysed four decades of data for 1,478 species of birds, mammals, butterflies, moths, plants and phytoplankton along a 1,200 km high latitudinal gradient. The relative importance of climatic drivers varied non-uniformly with progressing climate change. While species turnover among decades was limited, the relative position of species within their climatic niche shifted substantially. A greater proportion of species responded to climatic change at higher latitudes, where changes were stronger. These diverging climate imprints restructure a full biome, making it difficult to generalize biodiversity responses and raising concerns about ecosystem integrity in the face of accelerating climate change.The authors analyse four decades of distribution data for various taxonomic groups to understand the shift of species within their climatic niches and the changing influences of different climate factors. The diverse and diverging climate imprints raise concerns about future ecosystem integrity.

Funding Information: The shares of financing of the investigated TSCs are reported in Table S9. The differences in financing illustrate the absence of a systematic nationwide approach for financing TSCs. The farmers’ share of the costs was rather low in the six pilot studies, averaging less than 10%. Most financing has been arranged through individual, externally funded research and development projects. Currently, in Finland, a TSC project may obtain state financial support in the form of either a drainage subsidy from Agrifood Research and Development Fund (MAKERA) managed by the ELY Centers (Regional State Authority) or from CAP-AES’s non-productive investments as wetland investment and maintenance allowance. However, overlapping subsidies cannot be received for the same drainage area. Support can cover no more than 40% of the costs, but environmentally friendly (i.e., nature-based) solutions, and particularly expensive structures, may increase the support percentage. In recent years, MAKERA has granted approximately 2.5 million €/year of state support for about 50 dredging projects. Funding Information: Funding: This research was funded by the Joint Research Centre of European Commission (contract no. 939642), Academy of Finland (grant no. 330217), the Finnish Cultural Foundation (grant no. 00201229) and by the Valumavesi project. Publisher Copyright: © 2021 by the authors. Licensee MDPI, Basel, Switzerland. Conventional dredging of ditches and streams to ensure agricultural drainage and flood mitigation can have severe environmental impacts. The aim of this paper is to investigate the potential benefits of an alternative, nature-based two-stage channel (TSC) design with floodplains excavated along the main channel. Through a literature survey, investigations at Finnish field sites and expert interviews, we assessed the performance, costs, and monetary environmental benefits of TSCs in comparison to conventional dredging, as well as the bottlenecks in their financing and governance. We found evidence supporting the expected longer-term functioning of drainage as well as larger plant and fish biodiversity in TSCs compared to conventional dredging. The TSC design likely improves water quality since the floodplains retain suspended sediment and phosphorus and remove nitrogen. In the investigated case, the additional value of phosphorus retention and conservation of protected species through the TSC design was 2.4 times higher than the total costs. We demonstrate how TSCs can be made eligible for the obligatory vegetated riparian buffer of the European Union agri-environmental subsidy scheme (CAP-AES) by optimising their spatial application with respect to other buffer measures, and recommend to publicly finance their additional costs compared to conventional dredging at priority sites. Further studies on biodiversity impacts and long-term performance of two-stage channels are required.

Significance Cities represent novel environments with altered seasonality; they are warmer, which may accelerate growth, but light pollution can also lengthen days, misleading organisms that use daylength to predict seasonal change. Using long-term observational data, we show that urban populations of a butterfly and a moth have longer flight seasons than neighboring rural populations for six Nordic city regions. Next, using laboratory experiments, we show that the induction of diapause by daylength has evolved in urban populations in the direction predicted by urban warming. We thus show that the altered seasonality of urban environments can lead to corresponding evolutionary changes in the seasonal responses of urban populations, a pattern that may be repeated in other species., Urbanization is gaining force globally, which challenges biodiversity, and it has recently also emerged as an agent of evolutionary change. Seasonal phenology and life cycle regulation are essential processes that urbanization is likely to alter through both the urban heat island effect (UHI) and artificial light at night (ALAN). However, how UHI and ALAN affect the evolution of seasonal adaptations has received little attention. Here, we test for the urban evolution of seasonal life-history plasticity, specifically changes in the photoperiodic induction of diapause in two lepidopterans, Pieris napi (Pieridae) and Chiasmia clathrata (Geometridae). We used long-term data from standardized monitoring and citizen science observation schemes to compare yearly phenological flight curves in six cities in Finland and Sweden to those of adjacent rural populations. This analysis showed for both species that flight seasons are longer and end later in most cities, suggesting a difference in the timing of diapause induction. Then, we used common garden experiments to test whether the evolution of the photoperiodic reaction norm for diapause could explain these phenological changes for a subset of these cities. These experiments demonstrated a genetic shift for both species in urban areas toward a lower daylength threshold for direct development, consistent with predictions based on the UHI but not ALAN. The correspondence of this genetic change to the results of our larger-scale observational analysis of in situ flight phenology indicates that it may be widespread. These findings suggest that seasonal life cycle regulation evolves in urban ectotherms and may contribute to ecoevolutionary dynamics in cities.

Species can adapt to climate change by adjusting in situ or by dispersing to new areas, and these strategies may complement or enhance each other. Here, we investigate temporal shifts in phenology and spatial shifts in northern range boundaries for 289 Lepidoptera species by using long-term data sampled over two decades. While 40% of the species neither advanced phenology nor moved northward, nearly half (47%) -used one of the two strategies. The strongest positive population trends was observed for the minority of species (13%) that both advanced flight phenology and shifted their northern range boundaries northward. We show that, for Boreal Lepidoptera, a combination of phenology and range shifts is the most viable strategy under a changing climate. Effectively, this may divide species into winners and losers based on their propensity to capitalize on this combination, with potentially large consequences on future community composition.

Recent actions to mitigate biodiversity loss in agricultural environments appear insufficient despite the considerable efforts channeled via the European Union’s Common Agricultural Policy. One likely reason for this failure is the limited attention paid to the regional and landscape level ecological characteristics in farmland conservation planning. We demonstrate how to obtain conservation prioritization solutions that would address simultaneously three goals, including two landscape level targets: minimizing local habitat quality loss, maximizing habitat connectivity, and incorporating landscape heterogeneity. As these goals may be contradictory, we investigate the potential trade-offs between them. We used the Zonation prioritization tool to examine how our three goals could be implemented in the agricultural landscapes of southwest Finland. We used measures of (i) biodiversity value of grasslands, (ii) connectivity between grasslands, and (iii) landscape heterogeneity which comprised of (land cover type based) compositional heterogeneity and (field margin based) configurational heterogeneity. Integration of landscape heterogeneity measures and habitat connectivity resulted in some tradeoffs with local habitat quality, the most prominent observation being that landscape heterogeneity co-varied with grassland connectivity. Among the two landscape heterogeneity parameters, inclusion of compositional heterogeneity resulted in more clustered prioritization solutions than configurational heterogeneity, which had a spatially more balanced impact. Concordance among landscape scale factors implies high potential for reconstruction of a functioning network of semi-natural grasslands in areas under intensive agricultural use. Broader scale multi-objective planning approaches can thus importantly support targeting biodiversity conservation planning and mediating the implementation of Common Agricultural Policy objectives. peerReviewed

Ecological intensification has been proposed as a strategy to mitigate the impacts of conventional intensive land use on pollinators and insect pollination. However, research has concentrated on the effects of land management on pollinator abundance and diversity, while studies directly measuring pollination services are still lacking for many crops and regions. This study examines the effects of landscape heterogeneity, pesticide use intensity and flowering time on insect pollination of turnip rape Brassica rapa ssp. oleifera, a major oil crop at high latitudes. The field experiment included monitoring pollinator visits on turnip rape flowers and measuring yield in 34 spring-sown turnip rape fields in Southern Finland. The fields were situated in two landscape types that differed in the cover of arable land and represented independent gradients of pesticide use intensity and flowering time. Based on flower visits and number of seeds per silique, turnip rape was best pollinated in fields with late flowering time. The result suggests a temporal mismatch between crop flowering and the availability of pollination service in early sown fields. The increase in flower visits during summer was steeper among fields with low than high pesticide use, possibly due to a faster colony growth of important bee pollinators. Pollinator community in turnip rape fields was more diverse in heterogeneous landscapes with less arable land than in homogeneous field-dominated landscapes, suggesting higher stability of pollination services in diverse landscapes. Total yield per plant was positively related to pesticide use intensity. The relationship between insect pollination and total yield was weakened by high compensation capacity of turnip rape, the plants producing more flowers under poor pollination.Synthesis and applications. Crop pollination can be enhanced by shifting sowing time to better match crop flowering to the peak availability of the most important pollinators. Reduced pesticide use may also enhance pollination services but can lead to the total yield reduction due to pests. This highlights the need for pollinator-friendly pest control methods to maintain turnip rape yields while conserving pollination services. Foreign Language AbstractTiivistelma Maatalouden ekologista tehostamista on esitetty strategiaksi, jonka avulla intensiivisen maankayton kielteisia vaikutuksia polyttajiin ja hyonteispolytykseen voidaan hillita. Tutkimukset maankayton vaikutuksista viljelykasvien hyonteispolytykseen kuitenkin puuttuvat yha monilta viljelykasveilta ja alueilta. Tama tutkimus tarkasteli maiseman monimuotoisuuden, torjunta-aineiden kayton ja kukinta-ajan vaikutusta rypsin hyonteispolytykseen. Peltokokeessa seurattiin polyttajien vierailuja rypsin kukilla ja mitattiin satoa 34 kevatrypsipellolla Uudellamaalla. Pellot sijaitsivat kahdessa maisematyypissa, joista toinen oli peltovaltainen ja toinen monimuotoisempi ja vahemman peltoa sisaltava. Lisaksi pellot erosivat toisistaan torjunta-aineiden kayton ja kukinta-ajan suhteen. Polyttajien kukkavierailut ja siementen maara per litu olivat korkeimmillaan myohaan kukkineilla rypsipelloilla, mika kertoo hyvasta polytyksesta. Tuloksen perusteella aikaisin kylvetyt rypsipellot saattavat kukkia liian aikaisin polyttajien saatavuuden nakokulmasta. Voimakkaimmin polyttajien maara lisaantyi kesan aikana niiden peltojen joukossa, joilla ei kaytetty torjunta-aineita. Mahdollinen selitys on, etta tarhamehilaisten ja kimalaisten yhdyskunnat kasvoivat nopeimmin naiden peltojen laheisyydessa. Polyttajien monimuotoisuus oli suurempi monimuotoisessa kuin peltovaltaisessa maisemassa, mika parantaa polytyspalvelun vakautta muuttuvissa olosuhteissa. Kokonaissato per kasvi nousi torjunta-aineiden kayton lisaantyessa. Hyonteispolytyksen ja sadon valista yhteytta heikensi rypsin hyva kompensaatiokyky, jonka ansiosta heikosti polyttyneet kasvit tuottivat enemman kukkia.Yhteenveto ja sovellukset. Viljelykasvin polytysta voi parantaa saatamalla kylvoaikaa siten, etta kasvin kukinta ajoittuu tarkeimpien polyttajien runsaushuippuun. Torjunta-aineiden kayton vahentaminen voi parantaa polytysta mutta laskea samaan aikaan satoa tuholaisten ja rikkakasvien lisaantymisen takia. Polyttajaystavallisten kasvinsuojelukeinojen kehittaminen on tarkeaa korkeiden rypsisatojen ja polytyspalveluiden sailymisen kannalta. Crop pollination can be enhanced by shifting sowing time to better match crop flowering to the peak availability of the most important pollinators. Reduced pesticide use may also enhance pollination services but can lead to the total yield reduction due to pests. This highlights the need for pollinator-friendly pest control methods to maintain turnip rape yields while conserving pollination services. peerReviewed

Good knowledge on how increasing urbanization affects biodiversity is essential in order to preserve biodiversity in urban green spaces. We examined how urban development affects species richness and total abundance of butterflies as well as the occurrence and abundance of individual species within the Helsinki metropolitan area in Northern Europe. Repeated butterfly counts in 167 separate 1-km-long transects within Helsinki covered the entire urbanization gradient, quantified by human population density and the proportion of built-up area (within a 50-m buffer surrounding each butterfly transect). We found consistently negative effects of both human population density and built-up area on all studied butterfly variables, though butterflies responded markedly more negatively to increasing human population density than to built-up area. Responses in butterfly species richness and total abundance showed higher variability in relation to proportion of built-up area than to human density, especially in areas of high human density. Increasing human density negatively affected both the abundance and the occurrence of 47% of the 19 most abundant species, whereas, for the proportion of built-up area, the corresponding percentages were 32% and 32%, respectively. Species with high habitat specificity and low mobility showed higher sensitivity to urbanization (especially high human population density) than habitat generalists and mobile species that dominated the urban butterfly communities. Our results suggest that human population density provides a better indicator of urbanization effects on butterflies compared to the proportion of built-up area. The generality of this finding should be verified in other contexts and taxonomic groups.

Climatic anomalies are increasing in intensity and frequency due to rapid rates of global change, leading to increased extinction risk for many species. The impacts of anomalies are likely to vary between species due to different degrees of sensitivity and extents of local adaptation. Here, we used long-term butterfly monitoring data of 143 species across six European bioclimatic regions to show how species' population dynamics have responded to local or globally-calculated climatic anomalies, and how species attributes mediate these responses. Contrary to expectations, degree of apparent local adaptation, estimated from the relative population sensitivity to local versus global anomalies, showed no associations with species mobility or reproductive rate but did contain a strong phylogenetic signal. The existence of phylogenetically-patterned local adaptation to climate has important implications for forecasting species responses to current and future climatic conditions and for developing appropriate conservation practices.

Uncultivated field margins are important refugia for pollinating insects in agricultural landscapes. However, the spill-over of pollination services from field margins to adjacent crops is poorly understood. This study (i) examined the effects of landscape heterogeneity on pollinator occurrence in permanent field margins and pollinator visitation to adjacent mass-flowering turnip rape (Brassica rapa ssp. oleifera) in boreal agricultural landscapes, and (ii) tested whether pollinator abundance and species richness in field margins predict abundance and species richness of crop visitors. Pollinators visiting the crop were more affected by landscape heterogeneity than pollinators in adjacent margins. Species richness, total abundance, and the abundance of syrphid flies visiting the crop increased with increasing landscape heterogeneity, whereas, in field margins, landscape heterogeneity had little effect on pollinators. In field-dominated homogeneous landscapes, wild pollinators rarely visited the crop even if they occurred in adjacent margins, whereas in heterogeneous landscapes, differences between the two habitats were smaller. Total pollinator abundance and species richness in field margins were poor predictors of pollinator visitation to adjacent crop. However, high abundances of honeybees and bumblebees in margins were related to high numbers of crop visitors from these taxa. Our results suggest that, while uncultivated field margins help pollinators persist in boreal agricultural landscapes, they do not always result in enhanced pollinator visitation to the adjacent crop. More studies quantifying pollination service delivery from semi-natural habitats to crops in different landscape settings will help develop management approaches to support crop pollination.

The European Union's Natura 2000 (N2000) is among the largest international networks of protected areas. One of its aims is to secure the status of a predetermined set of (targeted) bird and butterfly species. However, nontarget species may also benefit from N2000. We evaluated how the terrestrial component of this network affects the abundance of nontargeted, more common bird and butterfly species based on data from long-term volunteer-based monitoring programs in 9602 sites for birds and 2001 sites for butterflies. In almost half of the 155 bird species assessed, and particularly among woodland specialists, abundance increased (slope estimates ranged from 0.101 [SD 0.042] to 3.51 [SD 1.30]) as the proportion of landscape covered by N2000 sites increased. This positive relationship existed for 27 of the 104 butterfly species (estimates ranged from 0.382 [SD 0.163] to 4.28 [SD 0.768]), although most butterflies were generalists. For most species, when land-cover covariates were accounted for these positive relationships were not evident, meaning land cover may be a determinant of positive effects of the N2000 network. The increase in abundance as N2000 coverage increased correlated with the specialization index for birds, but not for butterflies. Although the N2000 network supports high abundance of a large spectrum of species, the low number of specialist butterflies with a positive association with the N2000 network shows the need to improve the habitat quality of N2000 sites that could harbor open-land butterfly specialists. For a better understanding of the processes involved, we advocate for standardized collection of data at N2000 sites.Efectos de Natura 2000 sobre las Especies No Focales de Aves y Mariposas con Base en Datos de Ciencia Ciudadana Resumen La red Natura 2000 (N2000) de la Unión Europea está entre las redes internacionales más grandes de áreas protegidas. Uno de sus objetivos es asegurar el estado de un conjunto predeterminado de especies de aves y mariposas (focales). Sin embargo, las especies no focales también pueden beneficiarse con la N2000. Evaluamos cómo el componente terrestre de esta red afecta la abundancia de las especies de aves y mariposas no focales más comunes con base en los datos de programas de monitoreo voluntario a largo plazo en 9,602 sitios para aves y en 2,001 sitios para mariposas. En casi la mitad de las 155 especies de aves evaluadas, particularmente entre aquellas especies especialistas en zonas boscosas, la abundancia incrementó (los estimaciones de la pendiente variaron desde 0.101 [DS 0.042] hasta 3.51 [DS 1.30]) conforme incrementó la proporción del paisaje cubierto por sitios de la N2000. Esta relación positiva existió en 27 de las 104 especies de mariposas (con una variación de estimaciones desde 0.382 [DS 0.163] hasta 4.28 [DS 0.768]), aunque la mayoría de las especies de mariposas fueron generalistas. Cuando se consideraron las covarianzas de cobertura de suelo estas relaciones positivas no fueron evidentes para la mayoría de las especies, lo que significa que la cobertura de suelo puede ser una determinante de los efectos positivos de la red N2000. El incremento en la abundancia conforme aumentó la cobertura de la N2000 estuvo correlacionado con el índice de especialización de las aves, pero no el de las mariposas. Aunque la red N2000 sostiene la abundancia alta de un espectro amplio de especies, el bajo número de mariposas especialistas con una asociación positiva a la red N2000 demuestra la necesidad de mejorar la calidad del hábitat de los sitios N2000 que podrían albergar a mariposas especialistas de campo abierto. Para un mejor entendimiento de los procesos involucrados, promovemos una recolección estandarizada de datos en los sitios de la red N2000.欧盟的 Natura 2000(N2000) 是全球最大的保护地网络之一, 它的目标之一是保护一批目标鸟类和蝴蝶物种的现状。不过, 非目标物种也可能从 N2000 中获益。本研究评估了该网络陆地保护区如何影响更为常见的非目标鸟类和蝴蝶的丰度, 所用数据来自长期的志愿者监测项目, 包括了 9602 个鸟类监测点和 2001 个蝴蝶监测点。在评估的 155 种鸟类中, 几乎一半的鸟类, 特别是林地专性种, 在 N2000 位点覆盖的景观比例上升时数量增加 (斜率估计值在 0.101 [SD 0.042] 到3.51 [SD 1.30] 之间) 。虽然大多数蝴蝶都是广幅种, 但 104 种蝴蝶中也有 27 种存在这种正相关关系 (斜率估计值在 0.382 [SD 0.163] 到 4.28 [SD 0.768]) 。对于大多数物种来说, 当考虑土地覆盖协变量时, 这样的正相关关系并不明显, 这意味着土地覆盖可能是 N2000 网络能否对物种产生积极影响的关键因素。物种的丰度随着 N2000 覆盖度增加而增加的现象与鸟类的生境特化指数相关, 而在蝴蝶中则无关。虽然 N2000 网络使许多物种得以维持较高丰度, 但获得其积极影响的蝴蝶专性种数量仍较少, 这说明需要在 N2000 网络中提高专性生活在开阔地的蝴蝶的生境质量。为了更好地理解其中的过程, 我们建议收集更多标准化的 N2000 位点数据。 【翻译: 胡怡思; 审校: 聂永刚】.

The amount of High Nature Value farmland (HNVf) is a commonly used environmental indicator for assessing the performance of the Common Agricultural Policy, to support sustainable agriculture and monitor changes in agricultural land use in Europe. HNVf comprises agricultural areas of semi-natural state, low-intensity farming and fine-scale landscape mosaics of different habitat types. For a successful implementation, the identification of HNVf should correctly reflect the variation in biodiversity values between different agricultural landscapes. We examined how well the Finnish HNVf indicator and the sub-indicators constituting it – recalculated for the purposes of this study for five study regions – reflect the variation in bird and butterfly species richness and diversity patterns at different spatial scales. We found that butterfly diversity index was positively associated with the HNVf indicator at the finest scale of 0.5 km × 0.5 km squares. Among the HNVf sub-indicators, extensive cultivation of grasslands was most strongly related to the farmland bird diversity and the density of edge to the butterfly diversity. Thus, the HNVf concept reflects well the distribution of butterflies in the Finnish agricultural landscapes but insufficiently the diversity patterns of farmland birds. Importantly, semi-natural vegetation and long-term pastures – the backbone of the concept – presently occur in small and highly fragmented patches in agricultural landscapes in Finland. The Pan-European concept of HNVf has restricted application to farmland birds of this boreal country and the national HNVf concept may need to be revised.

Set-asides sown to wildflowers are introduced to promote agrobiodiversity. Often, sown seed mixtures have been used to enhance aboveground biodiversity while belowground biodiversity has received less attention. We studied in a 15-year-old set-aside experiment, the impact of seed mixtures (grass vs. meadow) and mowing (mown vs. unmown) on the abundance and diversity of plants, two aboveground invertebrate groups (pollinators and natural enemies of pest species) and one key belowground group, earthworms. We also studied the treatment effects on selected soil chemical properties. In general, a sown meadow seed mixture mostly benefitted aboveground biodiversity, whereas the impacts of late-season mowing were most discernible for earthworms. Plant species richness or coverage did not differ between seed mixtures, but plant coverage showed a positive response to mowing, while the impact on species richness was minor. The meadow seed mixture was dominated by Centaurea jacea. A non-mown meadow treatment supported the highest earthworm density and species richness, while the values were lowest in the mown meadow, where earthworm total mass was also at its lowest. In a grass seed mixture of intermediate density and richness, the mowing had no measurable effect on the earthworm community. Topsoil carbon, nitrogen and soluble phosphorus concentrations showed a typical gradient for non-inverted soils, with the highest concentrations in the 0–2.5 cm layer. The two uppermost soil layers (to 5 cm depth) had higher soluble phosphorus content under grass than under meadow seed mixture. Abundance of natural enemies of pests was not significantly different between the treatments, but for carabids there was a marginally significant positive response to mowing. Bumblebee species richness and the abundance of bumblebees and honeybees were highest in the mown meadow seed mixture plots. Butterfly species richness was marginally significantly higher in mown than in non-mown treatments in both seed mixtures. The results showed the potential of set-asides to promote both aboveground and belowground biodiversity through increased plant diversity.

Conservation translocation is a controversial, yet sometimes the only applicable, conservation tool in the face of the human-induced environmental change. One important aspect often neglected in translocation experiments is predicting the spatial expansion potential of the translocated population in the release area. In this study, we used a spatially explicit movement model to project the expansion rate of a translocated clouded Apollo (Parnassius mnemosyne) population. The model was parameterised with data collected from the source population inhabiting an open landscape with well-connected habitat patches. We then used these parameter estimates to simulate the spread of the translocated population in the new area, which was a forested landscape with more fragmented habitat. We tested the model predictions by comparing them to the observed expansion rate between 2000 and 2016. Additionally, we tested whether including annually varying weather conditions would increase realism in the projections. Model predictions were qualitatively correct and quantitatively most accurate for flight seasons with prevailing weather conditions similar to those of the season when the data for parameter estimation was collected. We conclude that spatially explicit movement models are potential tools to improve the planning of future conservation translocations in terms of choosing the optimal release area, as they enable the comparison of functional connectivities even between structurally different landscapes. However, our results suggest that a model fitted based on movement data from a single year may not be representative for the overall dispersal rates due to temporal variation in dispersal.

Agricultural intensification has caused drastic declines in the area and species richness of semi-natural grasslands across Europe. Novel habitats, such as power line clearings, provide alternative habitats and niches for grassland species, and might therefore mitigate these declines. However, it is not fully understood which environmental factors determine the occurrence of grassland species in the clearings. Identifying the most important drivers for grassland species occurrence would help understand the value of the clearings for grassland conservation and target enhanced management into clearings with most potential as grassland habitat. We studied the effects of local environmental conditions, and past and present connectivity to semi-natural grasslands, on the species richness of grassland plants and butterflies in 43 power line clearings in Finland. The results of generalized linear models and hierarchical partitioning showed that increasing time since clear-cut and amount of clearing residue decreased the species richness of both species groups, while the cover of mesic habitats increased it. However, the two species groups showed also divergent responses. Present-day local environmental conditions appeared to be the sole driver of grassland butterfly species richness, whereas the richness of grassland plants was related both to current conditions and historical connectivity to grasslands in 1870–1880s. This suggests the presence of an extinction debt in the studied grassland plant communities, emphasizing the need for enhanced management to increase suitable grassland habitat in the clearings. This would diminish the potential future losses of grassland plant species in the clearings and create valuable habitat for grassland butterflies as well.

Gregarious, social caterpillars have stimulated research because group size may affect survival, growth rate, thermoregulation, and interactions with other species, yet group size is often variable both within and among populations. We used a combination of observations and experiments to study the importance of group size for egg and larval survival in the Glanville fritillary butterfly, Melitaea cinxia, which lives in groups from egg hatching until the last larval instar. Both experimental manipulation of egg clutches placed in the field and observations of naturally occurring groups showed that survival increased with increasing group size. This pattern was present independently during all four developmental stages studied: eggs, prediapause larvae, diapausing larvae and post-diapause larvae. However, it was significant only during two stages: pre-diapause (in one year only) and diapausing larvae (in all years). Large group size increased survival of entire larval groups as well as that of individual larvae within surviving groups. These results may explain why cluster size is large and why adults oviposit infrequently. Large cluster size, coupled with correlated survival of group members, in turn helps to explain the unstable local dynamics and short average persistence time of local M. cinxia populations.

Clumped distributions of herbivorous insect eggs often result from independent assess- ments of individual plants by different ovipositing females. Here we ask whether, in addition, plants might be rendered more or less attractive to ovipositing Melitaea cinxia butter ies by presence of conspeci c eggs and/or by prior larval attack. Both eggs and larval damage rendered Veronica spicata plants signi cantly more accept- able; the effect of eggs was particularly strong. Larval damage caused a marginally signi cant increase in acceptability of Plantago lanceolata, but there was no trend for an effect of eggs on this host. Variable oviposition preferences of Melitaeine butter ies are known to drive their metapopulation dynamics by affecting rates of emigration and patch colonization. Therefore variable host acceptability, as documented here, should do likewise, reducing emigration rates at high population densities where V. spicata is present in the landscape and complementing Allee effects that are already known in this system.

High cover of forest in the landscape matrix has been shown to weaken the negative effects of habitat fragmentation on grassland butterflies. No studies have however focused on examining species-specific responses of grassland butterflies to forest. The data from 3 years of butterfly monitoring in Southern Finland were used to test whether the amount of forest cover in the surrounding landscape affected the abundance of grassland butterfly species in semi-natural grasslands, field margins, and forest edges. More than half of the studied species benefitted from high cover of forest. Species with the strongest preference for forested landscapes were Lycaena virgaureae, Argynnis adippe, Argynnis aglaja, and Boloria selene, which probably find suitable resources in herbaceous habitats at forest edges and clearings. Several small-sized species were positively affected by surrounding forest cover in field margins but not in the other habitat types. Although field margins are suboptimal habitats for grassland butterflies, they provide important corridors for dispersal. High cover of forest in the landscape matrix may enhance butterfly dispersal along field margins by reducing windiness, which is likely to be most important for small-sized species with poor dispersal capacity. The most abundant grassland species showed little or no preference for forested landscapes. Our results suggest that high cover of forest enhances the persistence of most grassland butterflies, including declining species, in boreal agricultural landscapes. The responses to forest are however strongly dependent on species-specific properties and habitat types.

Measures of functional diversity are expected to predict community responses to land use and environmental change because, in contrast to taxonomic diversity, it is based on species traits rather than their identity. Here, we investigated the impact of landscape homogenisation on plants, butterflies and birds in terms of the proportion of arable field cover in southern Finland at local (0.25 km2) and regional (> 10 000 km2) scales using four functional diversity indices: functional richness, functional evenness, functional divergence and functional dispersion. No uniform response in functional diversity across taxa or scales was found. However, in all cases where we found a relationship between increasing arable field cover and any index of functional diversity, this relationship was negative. Butterfly functional richness decreased with increasing arable field cover, as did butterfly and bird functional evenness. For butterfly functional evenness, this was only evident in the most homogeneous regions. Butterfly and bird functional dispersion decreased in homogeneous regions regardless of the proportion of arable field cover locally. No effect of landscape heterogeneity on plant functional diversity was found at any spatial scale, but plant species richness decreased locally with increasing arable field cover. Overall, species richness responded more consistently to landscape homogenisation than did the functional diversity indices, with both positive and negative effects across species groups. Functional diversity indices are in theory valuable instruments for assessing effects of land use scenarios on ecosystem functioning. However, the applicability of empirical data requires deeper understanding of which traits reliably capture species' vulnerability to environmental factors and of the ecological interpretation of the functional diversity indices. Our study provides novel insights into how the functional diversity of communities changes in response to agriculturally derived landscape homogenisation; however, the low explanatory power of the functional diversity indices hampers the ability to reliably anticipate impacts on ecosystem functioning. (Less)

Essential Biodiversity Variables (EBVs) have been suggested to harmonize biodiversity monitoring worldwide. Their aim is to provide a small but comprehensive set of monitoring variables that would give a balanced picture of the development of biodiversity and the reaching of international and national biodiversity targets. Globally, GEO BON (Group on Earth Observations Biodiversity Observation Network) has suggested 22 candidate EBVs to be monitored. In this article we regard EBVs as a conceptual tool that may help in making national scale biodiversity monitoring more robust by pointing out where to focus further development resources. We look at one country –Finland –with a relatively advanced biodiversity monitoring scheme and study how well Finland’s current biodiversity state indicators correspond with EBVs. In particular, we look at how national biodiversity monitoring could be improved by using available remote sensing (RS) applications. Rapidly emerging new technologies from drones to airborne laser scanning and new satellite sensors providing imagery with very high resolution (VHR) open a whole new world of opportunities for monitoring the state of biodiversity and ecosystems at low cost. In Finland, several RS applications already exist that could be expanded into national indicators. These include the monitoring of shore habitats and water quality parameters, among others. We hope that our analysis and examples help other countries with similar challenges. Along with RS opportunities, our analysis revealed also some needs to develop the EBV framework itself.

Land-use change is one of the primary drivers of species loss, yet little is known about its effect on other components of biodiversity that may be at risk. Here, we ask whether, and to what extent, landscape simplification, measured as the percentage of arable land in the landscape, disrupts the functional and phylogenetic association between primary producers and consumers. Across seven European regions, we inferred the potential associations (functional and phylogenetic) between host plants and butterflies in 561 semi-natural grasslands. Local plant diversity showed a strong bottom-up effect on butterfly diversity in the most complex landscapes, but this effect disappeared in simple landscapes. The functional associations between plant and butterflies are, therefore, the results of processes that act not only locally but are also dependent on the surrounding landscape context. Similarly, landscape simplification reduced the phylogenetic congruence among host plants and butterflies indicating that closely related butterflies become more generalist in the resources used. These processes occurred without any detectable change in species richness of plants or butterflies along the gradient of arable land. The structural properties of ecosystems are experiencing substantial erosion, with potentially pervasive effects on ecosystem functions and future evolutionary trajectories. Loss of interacting species might trigger cascading extinction events and reduce the stability of trophic interactions, as well as influence the longer-term resilience of ecosystem functions. This underscores a growing realization that species richness is a crude and insensitive metric and that both functional and phylogenetic associations, measured across multiple trophic levels, are likely to provide additional and deeper insights into the resilience of ecosystems, and the functions they provide.

Aim Nitrogen deposition is a major global driver of change in plant communities, but its impacts on higher trophic levels are insufficiently understood. Here, we introduce and test a novel conceptual trait-based model describing how the effects of soil eutrophication cascade to higher trophic levels across differential plant–herbivore interactions. Location Northern Europe. Methods We synthesize previous literature on the effects of nitrogen on plants and herbivorous insects as well as relevant multispecies patterns of insect communities concerning species dietary breadth, body size, dispersal propensity and voltinism in order to derive the model. We empirically evaluate the proposed, hitherto untested, four main model pathways using statistical modelling and data on 1064 northern European butterfly and moth species, their life-history traits, phylogeny and population trends. Results We show that across all species: (1) larval dietary breadth and host plant foliar nitrogen content are positively and equally strongly related to insect body size, and that (2) multivoltinism, host plant preferences for soil nitrogen, body size and larval dietary breadth are positively related to population trends of butterflies and moths as predicted by the model. Positive relationships between plant foliar nitrogen content and body size as well as multivoltinism and population trends are the first multispecies demonstrations for these patterns. Main conclusions Soil nitrogen enrichment amplifies the diverging trends of herbivorous insects feeding on nitrophilous versus nitrophobous plants through differential plant–herbivore interactions, causing predictable changes in community composition at higher trophic levels. A positive foliar nitrogen–insect body size relationship, now empirically supported, is the integrating link within this cascade. As nitrogen deposition is a global driver, our model suggests that a major future trend may be an increased dominance of insects that are large, dispersive, multivoltine, dietary generalists or specialized on nitrophilous plant species at the expense of species preferring oligotrophic environments.

Red Lists are very valuable tools in nature conservation at global, continental and (sub-) national scales. In an attempt to prioritise conservation actions for European butterflies, we compiled a database with species lists and Red Lists of all European countries, including the Macaronesian archipelagos (Azores, Madeira and Canary Islands). In total, we compiled national species lists for 42 countries and national Red Lists for 34 of these. The most species-rich countries in Europe are Italy, Russia and France with more than 250 species each. Endemic species are mainly found on the Macaronesian archipelagos and on the Mediterranean islands. By attributing numerical values proportionate to the threat statuses in the different national Red List categories, we calculated a mean Red List value for every country (cRLV) and a weighted Red List value for every species (wsRLV) using the square root of the country’s area as a weighting factor. Countries with the highest cRLV were industrialised (NW) European countries such as the Netherlands, Belgium, the Czech Republic and Denmark, whereas large Mediterranean countries such as Spain and Italy had the lowest cRLV. Species for which a Red List assessment was available in at least two European countries and with a relatively high wsRLV (≥ 50) are Colias myrmidone, Pseudochazara orestes, Tomares nogelii, Colias chrysotheme and Coenonympha oedippus. We compared these wsRLVs with the species statuses on the European Red List to identify possible mismatches. We discuss how this complementary method can help to prioritise butterfly conservation on the continental and/or the (sub-)national scale.

Fallows (i.e. fields temporarily taken out of production) provide important habitat for flower-visiting insects in intensively cultivated agricultural landscapes. Cost-efficiency of fallowing schemes could be enhanced through improved understanding of key characteristics of fallows and surrounding landscape that determine community composition and provide support for species of conservation concern. Impacts of fallow characteristics and landscape structure on the species composition of butterflies and bumblebees were studied in two types of perennial fallows in boreal farmland. To understand species’ responses to environmental factors from a conservation perspective, community composition was examined with respect to two species traits—niche breadth and dispersal capacity. Whereas overall species composition of butterflies and bumblebees was strongly affected by forest cover in the surrounding landscape, the studied species traits were most related to fallow type and the cover of perennial grasslands. Habitat breadth of butterflies was narrowest in long-term grassland fallows in landscapes with high grassland cover. Dispersal capacity of butterflies was also lowest in grassland-rich landscapes. Diet breadth of bumblebees was narrower in long-term grassland fallows than in short-term fallows. The results confirm that the diversity of butterflies and bumblebees can be enhanced by establishing and managing fallows both in open and forested landscapes. For conservation of habitat specialists and less mobile species, retention of long-term fallows in grassland-rich landscapes is apparently the best option. The results provide no justification for exempting forested regions or farms with high grassland cover from the ecological focus area requirement under the European Union’s current agricultural policy.

Success rates for conservation translocations of species are low and there is a need for increased understanding of how this activity is best applied. Here, using static species distribution models and a spatially-explicit dynamic simulation model, RangeShifter, we examine the impacts of habitat cover in recipient landscapes, allocation of individuals into multiple sites and species trait characteristics on the long-term fate of hypothetical translocations of a grassland specialist butterfly, Maniola jurtina, in Finland. While persistence of populations introduced to climatically suitable locations northwards of the current range can be increased by selecting sites with increasing habitat cover and by allocation of individuals to multiple release sites, local population growth rate is shown to be the key parameter in determining likely translocation success. We conclude that the long-term persistence of translocated habitat specialist butterflies, particularly with low growth rates, appears to be uncertain in modern-day fragmented grassland networks and that translocation activities should prioritize management that improves local growth rate.

Translocations have been advocated as a conservation tool helping species adapt to climate and land-use change, but well-documented examples of invertebrates’ translocations are rare. The paper describes a successful translocation of the threatened Clouded Apollo butterfly ( Parnassius mnemosyne ) in Finland, compares this to a specific failed translocation, and presents conclusions for conservation planning as to factors contributing to the success. Two apparent key characteristics of the successful translocation were greater abundance of larval resources and less open landscape. The successful site was surrounded by forest, which strongly restricted emigration, crucially supporting the survival of the small initial population. Based on 20 mated females’ translocation in 2000, the local population increased slowly, reaching 600 butterflies in 2011. A large translocation patch together with host-plant abundance enabled successful establishment of the local population. Availability of other suitable grassland patches sufficiently nearby was an additional key characteristic, facilitating the Clouded Apollo’s expansion. However, the expansion rate was low; it took seven years for the butterflies to colonise the five nearest patches, only 10–200 m from the translocation patch. By 2013, they had colonised all suitable semi-natural grassland patches within 2 km from the translocation site and established a seemingly viable metapopulation with 11 subpopulations. The results point to the significance of local habitat area and landscape quality, along with conditions restricting emigration, in determination of suitable translocation sites.

1. Kleptoparasitic and parasitoid insects are expected to be particularly sensitive to changes in habitat availability due to their high trophic position and small population sizes compared with their hosts, but there are only few quantitative studies on their population changes. 2. Here, we studied the distribution and abundance of 48 kleptoparasitic and parasitoid species of cuckoo wasps (Chrysididae) and eight selected host species recorded in Finland from 1840 to 2015 based on an extensive survey of entomological collections. Population trends were assessed by studying changes in occupancy in 10 9 10 km grid squares between two study periods, 1840-1967 and 1968-2015. 3. Statistically significant decreases in occurrence were found for 11 cuckoo wasp species and one host species, while significant increases were not observed for any species. Trends of cuckoo wasps and their hosts were positively correlated, and changes were generally stronger in cuckoo wasps than in their hosts. 4. In a comparative analysis of species traits, abundance, body size and nesting type of host were related to occurrence changes of cuckoo wasps. Scarce and small species that use above ground-nesting hosts declined more than abundant and large species that use ground-nesting hosts. 5. Cuckoo wasp species dependent on dead wood are more vulnerable to changes in the environment than species associated with open sandy habitats. While both groups of species have probably suffered from habitat loss, the emergence of secondary habitats may have benefitted species living in sandy areas and compensated for the negative impact of habitat destruction.

Mitigation tools aimed at counteracting the loss of farmland insect diversity and associated ecosystem services are usually conducted on cultivated fields. Possibilities of managing field-forest ecotones for biodiversity have been rarely discussed, despite their importance for flower-visiting insects. The effectiveness of logging in enhancing three aspects of insect diversity (bumblebee abundance, total species richness, and habitat specialist butterfly abundance) and changes in insect community composition were studied in a three-year field experiment. The study included 15 field-forest ecotones, each with a 50 m long and 25 m wide treated area and a corresponding control. At forest edge, a 5 m wide strip was clear-cut and behind this strip, a 20 m wide belt was thinned to a basal area of 8 m2 ha−1. Logging simultaneously promoted all three aspects of insect diversity. Changes were more apparent at clear-cut edge than in the thinned interior. Community composition changed via increased abundance of open habitat species, especially butterflies. Logging promoted warm microclimates and flower availability at clear-cut edges. All three aspects of insect diversity showed a positive response to flower coverage, which in turn was negatively correlated with the amount of logging residue. Creating open field-forest ecotones is beneficial for flower-visiting insects and open habitat species in particular. This approach is expected to be most beneficial at sunny forest edges and sites which become overgrown slowly. In addition, we suggest logging residue to be removed when aiming to promote flower-visitors.

Long-term grassland fallow and short-term meadow fallow represent alternative fallowing strategies for biodiversity conservation on farmland. Conventional grassland mixtures are used to establish long-term grassland fallows, whereas short-term meadow fallows are sown with specific meadow seed mixtures and require regular re-establishment to maintain the abundance of sown species. This study examined the impact of fallow type and landscape structure on the species richness and abundance of four taxa: plants, bumblebees, butterflies and birds. Long-term grassland fallows and short-term meadow fallows were studied in four contrasting landscape types in Finland, differing in the cover of forest and perennial grasslands. The fallow type and landscape structure affected each species group differently, and the value of a particular fallow type was also modified by landscape setting. Bumblebees were most abundant in short-term meadow fallows, whereas butterflies benefited more from long-term grassland fallows. Species richness of plants and butterflies were higher in forested than in open landscapes. Long-term grassland fallows promoted species richness of butterflies as well as the abundance of foraging edge birds in forested landscapes, whereas short-term meadow fallows enhanced the abundance of foraging edge birds in open landscapes. In landscapes with high grassland cover, the breeding density of open farmland birds was higher in meadow fallows than in grassland fallows, while the reverse was true in landscapes with low perennial grassland cover. The fallowing strategy thus can be adapted to the landscape context depending on the specific objectives. For overall biodiversity enhancement, establishing and managing different kinds of fallows is important.

Mobility is a key factor determining lepidopteran species responses to environmental change. However, direct multispecies comparisons of mobility are rare and empirical comparisons between butterflies and moths have not been previously conducted. Here, we compared mobility between butterflies and diurnal moths and studied species traits affecting butterfly mobility. We experimentally marked and released 2011 butterfly and 2367 moth individuals belonging to 32 and 28 species, respectively, in a 25 m × 25 m release area within an 11-ha, 8-year-old set-aside field. Distance moved and emigration rate from the release habitat were recorded by species. The release experiment produced directly comparable mobility data in 18 butterfly and 9 moth species with almost 500 individuals recaptured. Butterflies were found more mobile than geometroid moths in terms of both distance moved (mean 315 m vs. 63 m, respectively) and emigration rate (mean 54% vs. 17%, respectively). Release habitat suitability had a strong effect on emigration rate and distance moved, because butterflies tended to leave the set-aside, if it was not suitable for breeding. In addition, emigration rate and distance moved increased significantly with increasing body size. When phylogenetic relatedness among species was included in the analyses, the significant effect of body size disappeared, but habitat suitability remained significant for distance moved. The higher mobility of butterflies than geometroid moths can largely be explained by morphological differences, as butterflies are more robust fliers. The important role of release habitat suitability in butterfly mobility was expected, but seems not to have been empirically documented before. The observed positive correlation between butterfly size and mobility is in agreement with our previous findings on butterfly colonization speed in a long-term set-aside experiment and recent meta-analyses on butterfly mobility.

This paper studies joint provision of two environmental non-market commodities related to agriculture: biodiversity conservation and water protection. We provide an optimising tool for analysing spatial dependencies of multifunctional agriculture at catchment scale. We show that efficiency gains can be achieved by spatial allocation and choice of the type of vegetation. In particular, inclusion of meadow nectar plants in the founding grass seed mixture of set-asides was found out to be an economically efficient measure to promote biodiversity and water protection on warm, steep slopes.

Biodiversity indicators are assumed to reflect changes in e.g. species richness of multiple taxa, but correlations in species richness between taxa have often been shown to be weak. However, only few studies are based on data allowing for rigorous tests whether strengths of correlations differ between habitat and landscape factors. We compared strengths of correlations between species richness of butterflies, plants and farmland birds between habitats (semi-natural grasslands, forest verges or field boundaries), spatial scales (0.8 ha, 25 ha and 50 ha) and landscapes differing in heterogeneity and regional land-use intensity. Between habitats, the correlation between butterflies and plants was strongest in semi-natural grasslands. Also concerning butterflies and plants, the correlation was weakest at the 0.8 ha scale, but no consistent scale-dependent patterns were found between plants and farmland birds. In a regional context, butterfly and plant species richness were consistently positively correlated, whereas when involving farmland birds we found correlations between taxa to be weaker and/or not significant in regions with high agricultural land-use intensity and in homogeneous landscapes. In general, species richness was consistently congruent only between butterflies and plants, whereas correlations involving farmland birds were mainly weak and showed contrasting patterns depending on regional context. Increasing landscape heterogeneity thus increased congruence amongst all studied taxa, but in different contexts and due to different underlying mechanisms. Although plants were involved in most of the significant correlations we cannot recommend a particular taxon as a general diversity indicator.

Sown wildflower strips have been used to mitigate against the loss of farmland biodiversity e.g. as a part of agri-environmental schemes. In this four-year field experiment, conducted on six field parcels, the ability of wildflower strips to simultaneously promote the availability of pollination services (using bumblebee abundance as a proxy), species diversity (total species richness of bumblebees, butterflies and diurnal moths) and conservation (abundance of habitat specialist butterflies) was evaluated. The results showed that the first two indicators increased strongly and rapidly in wildflower strips until the third year of the experiment. The conservation indicator increased more slowly and this response was related to the forest cover of the surrounding landscape. Furthermore, the benefits of wildflower strips were largely independent of experimentally varied strip properties. It is concluded that wildflower strips are a flexible mitigation tool to promote different biodiversity targets in agricultural systems. Pollination services and species diversity can be promoted in different kinds of landscapes by ensuring local habitat quality. In the promotion of habitat specialists, landscape matrix quality is more critical.

Agricultural intensification is a major driver of biodiversity decline throughout Europe. Agri-environment schemes governed by EU regulation are a significant tool in combating this decline but despite high spending, experiences of their effectiveness have been mixed. Their effectiveness might be improved by targeting them to locations with high biodiversity value, and particularly by spatial coordination to enhance habitat connectivity and the associated ecological processes, such as dispersal. We show, with an example of semi-natural grassland conservation in South-Western Finland, how spatial conservation planning tools, here the Zonation software, could help in assessing the habitat connectivity and allocating management actions. We assign highest priority to sites that have been classified as nationally important and that have been under management, and let connectivity influence the Zonation prioritizations. According to Zonation outputs, 25–30% of highest-ranking grasslands in our study area are without management contracts, indicating weak connectivity of managed sites, whereas Natura 2000 areas are spatially better located. A 50% expansion of the current network would be adequate to bring its value close to that of a network created from scratch, but as the contracts are temporary, reallocation of the contracts from the least valuable, funded sites to more valuable, unfunded ones would be even more effective. Current policy instruments supporting farmland biodiversity are strongly constrained by EU regulation, and appear to be too inflexible to take the spatial differences in conservation values into account. Better communication and incentives to encourage farmer participation to these voluntary programs are needed.

Losses of both habitat area and connectivity have been identified as important drivers of species richness declines, but little theoretical and empirical work exists that addresses the effect of fragmentation on relative commonness of highly mobile species such as pollinating insects. With a large dataset of wild bee and butterfly abundances collected across Europe, we first tested the effect of habitat area and connectivity on evenness in pollinator communities using a large array of indexes that give different weight to dominance and rarity. Second, we tested if traits related to mobility and diet breadth could explain the observed evenness patterns. We found a clear negative effect of area and a weaker, but positive effect of connectivity on evenness. Communities in small habitat fragments were mainly composed of mobile and generalist species. The higher evenness in small fragments could thereby be generated by highly mobile species that maintain local populations with frequent inter-fragment movements. Trait analysis suggested an increasing importance of dispersal over local recruitment, as we move from large to small fragments and from less to more connected fragments. Species richness and evenness were negatively correlated indicating that the two variables responded differently to habitat area and connectivity, although the mechanisms underlying the observed patterns are difficult to isolate. Even though habitat area and connectivity often decrease simultaneously due to habitat fragmentation, an interesting practical implication of the contrasting effect of the two variables is that the resulting community composition will depend on the relative strength of these two processes.

The rapid expansion of systematic monitoring schemes necessitates robust methods to reliably assess species' status and trends. Insect monitoring poses a challenge where there are strong seasonal patterns, requiring repeated counts to reliably assess abundance. Butterfly monitoring schemes (BMSs) operate in an increasing number of countries with broadly the same methodology, yet they differ in their observation frequency and in the methods used to compute annual abundance indices. Using simulated and observed data, we performed an extensive comparison of two approaches used to derive abundance indices from count data collected via BMS, under a range of sampling frequencies. Linear interpolation is most commonly used to estimate abundance indices from seasonal count series. A second method, hereafter the regional generalized additive model (GAM), fits a GAM to repeated counts within sites across a climatic region. For the two methods, we estimated bias in abundance indices and the statistical power for detecting trends, given different proportions of missing counts. We also compared the accuracy of trend estimates using systematically degraded observed counts of the Gatekeeper Pyronia tithonus (Linnaeus 1767). The regional GAM method generally outperforms the linear interpolation method. When the proportion of missing counts increased beyond 50%, indices derived via the linear interpolation method showed substantially higher estimation error as well as clear biases, in comparison to the regional GAM method. The regional GAM method also showed higher power to detect trends when the proportion of missing counts was substantial.Synthesis and applications. Monitoring offers invaluable data to support conservation policy and management, but requires robust analysis approaches and guidance for new and expanding schemes. Based on our findings, we recommend the regional generalized additive model approach when conducting integrative analyses across schemes, or when analysing scheme data with reduced sampling efforts. This method enables existing schemes to be expanded or new schemes to be developed with reduced within-year sampling frequency, as well as affording options to adapt protocols to more efficiently assess species status and trends across large geographical scales.Monitoring offers invaluable data to support conservation policy and management, but requires robust analysis approaches and guidance for new and expanding schemes. Based on our findings, we recommend the regional generalized additive model approach when conducting integrative analyses across schemes, or when analysing scheme data with reduced sampling efforts. This method enables existing schemes to be expanded or new schemes to be developed with reduced within-year sampling frequency, as well as affording options to adapt protocols to more efficiently assess species status and trends across large geographical scales.

Aim We test the prediction that beta diversity (species turnover) and the decay of community similarity with distance depend on spatial resolution (grain). We also study whether patterns of beta diversity are related to variability in climate, land cover or geographic distance and how the independent effects of these variables depend on the spatial grain of the data. Location Europe, Great Britain, Finland and Catalonia. Methods We used data on European birds, plants, butterflies, amphibians and reptiles, and data on British plants, Catalonian birds and Finnish butterflies. We fitted two or three nested grids of varying resolutions to each of these datasets. For each grid we calculated differences in climate, differences in land-cover composition (CORINE) and beta diversity (bsim, bJaccard) between all pairs of grid cells. In a separate analysis we looked specifically at pairs of adjacent grid cells (the first distance class). We then used variation partitioning to identify the magnitude of independent statistical associations (i.e. independent effects in the statistical sense) of climate, land cover and geographic distance with spatial patterns of beta

Local species richness of butterflies can be expected to benefit from both local habitat properties as well as the availability of suitable habitats and source populations in the surrounding landscape. Whether local species richness is dependent on local or landscape factors can be assessed by examining the relationship between local and landscape species richness. Here we studied how local species richness is related to landscape-level species richness in landscapes differing in agricultural intensity. The relationship was linear for field boundaries in intensively cultivated landscapes and non-linear in less-intensively cultivated landscapes. In landscapes containing semi-natural grasslands (on average 4% of overall land-use), the relationship was non-linear for field boundaries, but linear when considering local species richness of the grasslands themselves. These results show that local factors are more important than landscape factors in determining local species richness in landscapes which contained semi-natural grasslands. Local species richness was limited by landscape factors in intensively cultivated landscapes. This interpretation was supported by the relationship between local species richness and landscape-scale average mobility and generalist percentage of butterfly assemblages. We conclude that the management of field boundary habitat quality for butterflies is expected to be most effective in landscapes with semi-natural grasslands, the species composition of which in turn is dependent on the regional occurrence of grasslands. Based on our results, managing non-crop habitats for the conservation of habitat specialists and species with poor mobility will be most efficient in regions where patches of semi-natural grasslands occur.

Viljelymaiden lajistollinen monimuotoisuus on vähentynyt maataloustuotannon tehostumisen myötä. Monimuotoisuuden ja ekosysteemipalveluiden vähenemistä on yritetty pysäyttää maatalouden ympäristötuen toimenpiteillä, joiden huonona puolena pidetään kalleutta ja tehottomuutta. Tutkitulle tiedolle toimenpiteiden kustannusvaikuttavuudesta on siis tarvetta. Tässä tutkimuksessa vertaillaan pellolla ja peltoon rajoittuvan metsän puolella tehtävien monimuotoisuustoimenpiteiden kustannusvaikuttavuutta. Tarkasteltavat toimenpiteet ovat: 1) metsän puolelle peltoon rajoittuvan metsän reunaan perustettava monimuotoisuusvyöhyke, 2) pellon puolelle metsään rajoittuvan pellon reunaan perustettava monimuotoisuuskaista ja 3) luonnonhoitopelto. Peltoon rajoittuvan metsän reunaan perustettu 25 metriä leveä monimuotoisuusvyöhyke koostuu lähempänä peltoa olevasta 5 metriä leveästä niittymäisestä kaistasta ja syvemmällä metsässä olevasta 20 metriä leveästä vaihettumisvyöhykkeestä. Niittymäinen kapea kaista hakataan paljaaksi ja pidetään puuttomana 6–7 vuoden välein toistuvin raivauksin. Leveämpi vaihettumisvyöhyke puolestaan harvennetaan 8 m2/ha:n pohjapinta-alaan ja sitä käsitellään 20 vuoden välein toistuvin poimintahakkuin. Tässä tutkimuksessa pellolla olevalla monimuotoisuuskaistalla tarkoitetaan pellon reunassa olevaa 5 metriä leveää kaistaa, jolla kasvaa monivuotista nurmea tai niittykasveja. Tarkasteltavat luonnonhoitopellot ovat monivuotisia nurmipeltoja ja monivuotisten niittykasvien siemenseoksilla perustettuja monimuotoisuuspeltoja. Maaperän ravinteiden köyhdyttämiseksi ja metsittymisen ehkäisemiseksi monimuotoisuuskaistojen ja luonnonhoitopeltojen kasvusto niitetään ja korjataan pois kerran vuodessa. Lannoitteita tai torjunta-aineita ei käytetä. Toimenpiteiden vaikuttavuutta mitataan kimalaisten yksilömäärien lisääntymisellä kontrollikäsittelyyn eli vallitsevaan maankäyttötyyppiin verrattuna. Kimalaisten yksilömäärän kasvu suhteessa kontrolliin kuvaa monimuotoisuustoimenpiteen seurauksena syntyvää pölytyspalvelujen lisäystä. Monimuotoisuustoimenpiteistä maanomistajalle aiheutuvat kustannukset lasketaan vähentämällä monimuotoisuustoimenpiteen alalta saatavan nettotulovirran nykyarvo puu- tai peltokasvituotannosta vastaavalta alalta saatavan nettotulovirran nykyarvosta. Tulokset osoittavat, että peltotoimenpiteiden kustannusvaikuttavuus on metsätoimenpiteitä parempi. Peltotoimenpiteissä niittykasviseosten merkittävä positiivinen vaikutus kimalaisten runsauteen kumoaa seosten suuret siemenkustannukset. Näin ollen niittykasvien siemenseoksilla perustettujen monimuotoisuuskaistojen ja –peltojen kustannusvaikuttavuus on nurmiseoksella perustettuja kaistoja ja peltoja parempi.