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Changes in occupancy of Argynnis adippe sites in eight Morecambe Bay Limestones (dashed ellipses) and South Cumbria Low Fells (dotted ellipses) networks 1994–1997 to 2017. All 12 (re)colonisations occurred between 2008 and 2017 following intensive targeted management intervention on both occupied and potential sites. The only two new populations established in the UK since 1994 are highlighted (circles) as are the two (re)colonisations which have subsequently gone extinct (stars)
Source publication
We describe how a landscape-scale approach has been adopted to conserve the UK’s most threatened butterfly Argynnis adippe. Only 37 populations now remain, with 38 extinctions occurring since 1994 (51% loss). The butterfly has disappeared from most of England and Wales and is now confined to just four landscapes. Since 2005 management in these land...
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Background
The Glanville fritillary (Melitaea cinxia) butterfly is a model system for metapopulation dynamics research in fragmented landscapes. Here, we provide a chromosome-level assembly of the butterfly's genome produced from Pacific Biosciences sequencing of a pool of males, combined with a linkage map from population crosses.
Results
The fin...
While habitat loss is a major threat to species, the effects of habitat fragmentation independent of habitat loss (fragmentation per se) are debated. Metapopulation studies often assert negative fragmentation effects, but they do not measure fragmentation per se. We evaluate the effects of fragmentation per se (patch density) across 20 years of pat...
We apply a molecular approach to quantify the level of hymenopteran parasitoids infestation in the larvae of the marsh fritillary (Euphydryas aurinia), a declining butterfly species, in western Bohemia, Czech Republic, in two subsequent years. We used the novel primer HymR157 in combination with known universal 28SD1F to establish a PCR detection s...
Estimating distribution and abundance of species depends on the probability at which individuals are detected. Butterflies are of conservation interest worldwide, but data collected with Pollard walks - the standard for national monitoring schemes - are often analyzed assuming that changes in detectability are negligible within recommended sampling...
Citations
... Semi-natural grasslands, however, represent a habitat in dramatic decline (Donal et al., 2001;Luoto et al., 2003;Pärtel et al., 1999). In addition, many grassland butterflies require the proximity of forest habitats, with the reasons ranging from habitat associations of their larval host plants (Ellis et al., 2019) to microclimatic factors (e.g. Bergman, 1999). ...
... However, both positions are not necessarily in contradiction, since research into the causes of insect decline does not preclude action where this is already possible and should not be seen as a justification to delay action [26,27]. Being aware of this, many of the recent publications aim to identify the potential causes behind these trends [8,[28][29][30]. However, while many causes, e.g. ...
... A change to better habitat quality, for example, could be accompanied by a decreasing overall abundance whilst an increase of specific characteristic taxa is observed. Nevertheless, negative trends can be observed in many places, i.e. the abundance of insects is decreasing in specific areas, e.g. in nature conservation areas or the agricultural landscape [2,29]. Consequently, the discussion on appropriate policy responses to this reflects the underlying complexity and inter-relationship between these pressures, which is typical of multi-trophic ecosystems with multiple anthropogenic influences [22,26]. ...
Insect declines have been discussed intensively among experts, policymakers, and the public. Albeit, decreasing trends have been reported for a long time for various regions in Europe and North America, but the controversial discussion over the role of specific drivers and pressures still remains. A reason for these uncertainties lies within the complex networks of inter-dependent biotic and abiotic factors as well as anthropogenic activities that influence habitats, communities, populations, and individual organisms. Many recent publications aim to identify both the extent of the observed declines and potential drivers. With this literature analysis, we provide an overview of the drivers and pressures and their inter-relationships, which were concluded in the scientific literature, using some of the best-studied insect groups as examples. We conducted a detailed literature evaluation of publications on Carabidae (Coleoptera) and Lepidoptera trends with data for at least 6 years in countries of Central and Western Europe, with a focus on agricultural landscapes. From the 82 publications identified as relevant, we extracted all reported trends and classified the respective factors described according to the DPSIR model. Further, we analysed the level of scientific verification (presumed vs correlated vs examined) within these papers for these cited stressors. The extracted trends for both species groups underline the reported overall declining trend. Whether negative or positive trends were reported in the papers, our semi-quantitative analysis shows that changes in insect populations are primarily anthropogenically driven by agriculture, climate change, nature conservation activities, urbanisation, and other anthropogenic activities. Most of the identified pressures were found to act on habitat level, only a fraction attributed to direct effects to the insects. While our analysis gives an overview of existing research concerning abundance and biodiversity trends of carabids and lepidopterans, it also shows gaps in scientific data in this area, in particular in monitoring the pressures along with the monitoring of abundance trends. The scientific basis for assessing biodiversity changes in the landscape is essential to help all stakeholders involved to shape, e.g. agriculture and other human activities, in a more sustainable way, balancing human needs such as food production with conservation of nature.
... Due to the conservation status and specialised habitat requirements of the High Brown Fritillary, management should be directed towards this species, however the Dark Green Fritillary and other Viola-feeding fritillaries with wider niches (e.g. Pearl-bordered and Small Pearl-bordered Fritillaries) are also likely to benefit (Ellis et al. 2019). ...
The survival of butterfly populations depends on successful oviposition strategies. The limited mobility of early life stages requires females to select sites that reflect larval requirements. However, as land use and climate changes are altering habitat conditions and micro-climate, some species may adapt ovipositing strategies and flourish while others, with narrow niche requirements, may be unable to respond. Oviposition site selection and micro-habitat niche is examined for two closely related butterfly species—the specialist High Brown Fritillary (Fabriciana adippe) and relative generalist Dark Green Fritillary (Speyeria aglaja) through field observations of egg-laying females and analysis of micro-habitat characteristics. A total of 104 oviposition behaviour observations across both species were recorded in 69 1 m² quadrats, with the habitat characteristics compared to randomly selected quadrats in the same area. Results show that higher host plant density was a positively significant factor for oviposition site selection only for the High Brown Fritillary. Moreover, the cover of live Bracken (Pteridium aquilinum) and grass were important for site selection in both species, with High Brown Fritillaries tolerating less live Bracken and grass cover than Dark Green Fritillaries. This confirms the more specific requirements and narrower micro-habitat niche of the High Brown Fritillary, which appears to be more sensitive to micro-habitat cooling.
Implications for insect conservation
The management of Bracken mosaic habitats for these two species should aim to supress grass growth and maintain Bracken density within limits, by opening the Bracken canopy on a rotation through grazing or manual cutting, ensuring a continuous supply of suitable micro-habitat.
... Fabriciana adippe reaches as far north as central Sweden (Warren, 1995;Franzén and Johannesson, 2007;Fox et al., 2011;Tolman and Lewington, 2012), while F. niobe's northern range limit is southern Sweden (van Swaay and Warren, 1999;Eliasson et al., 2005;Reinhardt and Bolz, 2011). Fabriciana adippe is relatively constrained to open areas within woodland landscapes, and is facing steep declines in some parts of northern Europe, such as the UK (Warren, 1995;Zimmermann et al., 2009;van Swaay et al., 2010;Fox et al., 2011;Ellis et al., 2019), while F. niobe prefers nutrient-poor warm micro-habitats within grasslands with a high abundance of Viola spp. from sea-level up to sub-alpine altitudes (Salz and Fartmann, 2009;Spitzer et al., 2009;Salz and Fartmann, 2017). ...
... Speyeria aglaja can breed at lower temperatures than the other two studied species, and its distribution range reaches further north (Warren, 1995;Ellis et al., 2010;Fox et al., 2011;Tolman and Lewington, 2012). The observed trait correlation might contribute to S. aglaja's ecological generalism (it occupies a broad range of habitats), while the studied Fabriciana species are restricted to warm microclimates (Salz and Fartmann, 2009;Ellis et al., 2019). ...
Understanding how large-scale environmental variability may shape the distribution of phenotypic variation remains central to evolutionary biology. Across-species comparisons of trait variation alongside environmental gradients may offer valuable insights into how different species may respond to similar selective pressures. We conducted a comparative morphological study (>32° latitude and >47° longitude) on three closely related Argynnini butterfly species, Speyeria aglaja, Fabriciana adippe, and F. niobe. We measured wing size and coloration to assess (1) whether they respond similarly or differently to environmental factors (longitude, latitude, altitude, temperature, precipitation, solar radiation, wind speed); (2) if these factors correspond with those associated with the species’ genetic structure based on a previous study; and (3) whether correlations between phenotypic traits within individuals are species-specific. We found common and species-specific associations of climatic (precipitation, wind speed) and geographic (longitude, altitude) factors with the composite phenotypic variation. Wing size was associated with different variables in the studied species, while melanisation mainly increased in cooler regions in all species, suggesting that the need for temperature regulation is a strong selective pressure on melanisation. Wing size was associated with the genetic structure in all species, highlighting the functional importance of this trait. The environmental drivers associated with the phenotypic variation in S. aglaja and F. adippe were largely the same as those associated with their genetic structure, hinting at a genetic underpinning of the observed morphological variation due to local adaption. We report some distinct intraspecific trait correlations in S. aglaja and F. adippe, indicative of independent phenotypic integration. These phenotypes seem to be associated with protection against predators and thermal regulation in the respective habitats of both species, suggesting that similar selective pressures have resulted in the evolution of different trait combinations. Some of the inter-specific differences could be related to diverging niche breadths and dispersal capacities, exemplifying that the evolution of trait integration and spatial phenotypic differentiation may differ between closely related species with overlapping distribution ranges. Our findings highlight the importance of comparative assessments of variation, and demonstrate that the relative effects of drivers of variability may vary between sister species.
... The temporary removal of livestock or a reduction in grazing intensity may be required if vegetation is too short, whereas grazing or cutting can be reinstated where vegetation is becoming too tall for T. acteon. The challenge of managing habitat for mid-successional species in fragmented landscapes (Dolman et al. 2007, Ellis et al. 2019, Johansson et al. 2020, Mestre et al. 2020) could therefore be met using intermediate management intensity or frequency, for example by coordinated rotational management among networks of habitat patches. ...
Landscape‐scale approaches are increasingly advocated for species conservation but ensuring landscape level persistence by enlarging the size of patches or increasing their physical connectivity is often impractical. Here, we test how such barriers can be overcome by management of habitat at the local (site‐based) level, using a rare butterfly as an exemplar. We used four surveys of the entire UK distribution of the Lulworth skipper Thymelicus acteon over 40 years to test how local habitat influences population density and colonization/extinction dynamics, and parameterized, validated and applied a metapopulation model to simulate effects of varying local habitat quality on regional persistence. We found the total number of populations in four distribution snapshots between 1978 and 2017 varied between 59 and 84, and from 1997 to 2017 34% of local populations showed turnover (colonization or extinction). Population density was closely linked to vegetation characteristics indicative of management, namely height and food plant frequency, both of which changed through time. Simulating effects of habitat quality on metapopulation dynamics 40 years into the future suggests coordinated changes to two key components of quality (vegetation height and food plant frequency) would increase patch occupancy above the range observed in the past 40 years (50–80%). In contrast, deterioration of either component below threshold levels leads to metapopulation retraction to core sub‐networks of patches, or eventual extirpation. Our results indicate that changes to habitat quality can overcome constraints imposed by habitat patch area and spatial location on relative rates of colonization and local extinction, demonstrating the sensitivity of regional dynamics to targeted in situ management. Local habitat management therefore plays a key role in landscape‐scale conservation. Monitoring of population density, and the monitoring and management of local (site‐level) habitat quality, therefore represent effective and important components of conservation strategies in fragmented landscapes.
... While F. adippe is considered a species of Least Concern according to the IUCN Red List for Europe (van Swaay et al., 2010), it is listed as endangered on the UK Red List , with only 37 populations remaining in the UK (Ellis et al., 2019). Decreased coppicing and local conservation efforts to implement changes in forest management have enjoyed some success in protecting this species (Ellis et al., 2019). ...
... While F. adippe is considered a species of Least Concern according to the IUCN Red List for Europe (van Swaay et al., 2010), it is listed as endangered on the UK Red List , with only 37 populations remaining in the UK (Ellis et al., 2019). Decreased coppicing and local conservation efforts to implement changes in forest management have enjoyed some success in protecting this species (Ellis et al., 2019). Today, F. adippe is restricted to a few localities in western England and Wales. ...
We present a genome assembly from an individual female Fabriciana adippe (the high brown fritillary; Arthropoda; Insecta; Lepidoptera; Nymphalidae). The genome sequence is 485 megabases in span. Most of the assembly (99.98%) is scaffolded into 29 chromosomal pseudomolecules with the Z sex chromosome assembled. The complete mitochondrial genome was also assembled and is 15.1 kilobases in length. Gene annotation of this assembly in Ensembl identified 13,536 protein coding genes.
... The traits pointing towards specialism include a single adult generation per year, narrow host plant ranges and narrow requirements for host plant conditions (Ellis et al., 2019;Salz & Fartmann, 2009Spitzer, Beneš, & Konvička, 2009;Warren, 1995). Their conservation status varies regionally; some are among the most rapidly declining butterflies in respective regions (Powell et al., 2007;Sanford, 2011;Swartz et al., 2015), while others are still safe (Mikitova et al., 2021;Zimmermann et al., 2009). ...
... Argynnis adippe uses larger-sized violets (e.g., Viola canina, V. hirta, V. riviniana) growing at woodland clearings and edges (Britain: Warren, 1995;Polic et al., 2021). It is of least concern continentally (Maes et al., 2019;Van Swaay et al., 2010), but severely threatened in Britain, for example (Bonsall et al., 2014;Ellis et al., 2019;Fox et al., 2011). In the Czech Republic, the species had been declining until two decades ago, when it was classified as vulnerable (Benes et al., 2002), but a remarkable re-expansion followed and is continuing at present (Figure 2) across Central Europe (Reinhardt et al., 2020) Argynnis niobe prefers violets growing at desiccating conditions, amidst patches of near-bare ground (Salz & Fartmann, 2009Spitzer, Beneš, & Konvička, 2009). ...
... A difference between the two Argynnis species also manifested from comparison of conditions at capture spots. A. adippe occurred more closely to woodland edges, agreeing with its affinity for open forests and edges (Ellis et al., 2019;Fartmann et al., 2013;Reinhardt et al., 2020;Warren, 1995). A. niobe occurred in more open conditions, in agreement with earlier studies (Salz & Fartmann, 2009Stevens et al., 2010). ...
In human‐altered landscapes, specialist butterflies typically form spatially restricted populations, genetically differentiated due to dispersal restrictions. Generalists, in contrast, display minimum differentiation but high genetic diversity. While local‐level actions suffice to conserve specialists and landscape‐level actions are necessary for generalists, minimum information exists regarding conservation of species with intermediate features. We targeted two congeneric butterflies, the recently re‐expanding Argynnis adippe and the strongly declining A. niobe, co‐occurring in the pastoral landscape of the Carpathian Mountains, Czech Republic. We integrated species distribution models, mark‐recapture, and microsatellite analysis to compare their habitat requirements, adult demography, dispersal, and genetic patterns, and expanded the genetic analysis across the Carpathian Arc and beyond to delimit spatial conservation units. In two mountain valleys, both species formed interconnected populations numbering thousands of individuals. Mobility patterns suggested the populations’ interconnection across the Czech Carpathians. Genetic diversity was extremely poor in the non‐threatened A. adippe and moderate in the declining A. niobe. No population differentiation was detected within the Czech Carpathians (ca 1500 km2). Low genetic diversity and no differentiation was preserved in A. adippe across East Central Europe, whereas in A. niobe, populations from Serbia were differentiated from the Carpathian Arc + Alps. The high adult mobility linked to low differentiation probably reflect the distribution of larval resources, historically widespread but sparse and currently declining for A. niobe (grazing‐disturbed grounds), while currently increasing for A. adippe (abandonment scrub, disturbed woodlands). Units as large as entire mountain systems define population boundaries, and hence conservation management units, for both species.
... butte rfly-conse rvati on. org), including the High Brown Fritillary (Fabriciana [Argynnis] adippe), which has been observed to frequently lay its eggs in dead bracken and to complete pupation close to the ground under bracken (Ellis et al. 2019). But apart from these observations, dense bracken stands are generally assumed to be poor in animal species (Pakeman and Marrs 1992). ...
The native bracken fern ( Pteridium aquilinum (L.) Kuhn) has become increasingly invasive in abandoned or undergrazed Alpine grasslands. Bracken stands are generally assumed to be poor in species, but there is still very little information about the impact on the fauna. We recorded Orthoptera communities of 24 sample plots with varying bracken cover. Compared to plots with only sparse or medium bracken cover, species richness was highly significantly lower in densely overgrown plots, while the former two did not differ markedly. Multivariate ordination analyses revealed gradients with unequal orthopteran communities, showing distinct patterns of bracken cover clustering. Based on indicator values, 13 of 23 Orthoptera species turned out to be significantly or marginally significantly associated with a single bracken cover stratum or a combination of two strata, whereby all nine Red List species were primarily restricted to sparse or medium bracken cover. However, some generalist species were found to favor denser bracken stands, such as Tettigonia viridissima , contributing to the local Orthoptera beta diversity. Our data suggest that ongoing rotational sheep grazing with temporary paddocks is regarded as the most preferable management system. However, selected parts of the study area, which have only sparse vegetation and no pressure of bracken or scrub overgrowth, are recommended to be alternately left ungrazed for a few years to allow for the coexistence of different successional stages. Low-intensity bracken control measures, situationally by mowing or uprooting, are considered appropriate and should be pursued.
Implications for insect conservation
Even though most specialized and rare Orthoptera species in Western Europe tend to prefer sparsely vegetated areas, subareas with medium to even dense vegetation cover might promote certain generalist species, leading to a higher local species richness.
... Unfortunately, the life histories of many insect groups are not well understood. The one exception is the butterflies and moths (Lepidoptera), undoubtedly the best-studied insect lineage, and is already responding to climate change, habitat loss, and habitat fragmentation (Warren et al., 2001;Fox, 2013;Belitz et al., 2018;Maurer et al., 2018;Ellis et al., 2019). ...
Butterflies and moths (Lepidoptera) are one of the most studied, diverse, and widespread animal groups, making them an ideal model for climate change research. They are a particularly informative model for studying the effects of climate change on species ecology because they are ectotherms that thermoregulate with a suite of physiological, behavioural, and phenotypic traits. While some species have been negatively impacted by climatic disturbances, others have prospered, largely in accordance with their diversity in life‐history traits. Here we take advantage of a large repertoire of studies on butterflies and moths to provide a review of the many ways in which climate change is impacting insects, animals, and ecosystems. By studying these climate‐based impacts on ecological processes of Lepidoptera, we propose appropriate strategies for species conservation and habitat management broadly across animals.
... Markrecapture indicates that the male population in 2019 (the last year of population surveying) was less than 200 individuals, with bees not seen outside of the Seaford Head reserve, including in surveys conducted in 2020. The small size of the population is unlikely to be because A. retusa forages on rare plants or specialises on a narrow range of species, as is the case in some rare bees (Zayed and Packer 2007) and insect herbivores (Ellis et al. 2019). The plant species that was predominantly used for foraging was ground ivy, G. hederacea, which is extremely abundant in the local area and nationally. ...
Anthophora retusa is a rare solitary bee which has declined throughout Britain and other European countries since the 1990s. It is thought to be restricted to five sites in Britain. However, information on these remaining populations is limited. Knowledge on population size, habitat and forage requirements and foraging distance, are important for successful conservation of species. The population of A. retusa at the Seaford Head Nature reserve in East Sussex was surveyed. Transects within the reserve were conducted and population estimates using mark recapture were made for 2018 and 2019. Pollen from foraging females was analysed alongside visual sightings to determine forage requirements. The total population was estimated to be 91 in 2018 (males and females) with an estimated male population of 167 in 2019. The most visited flower species by females was Glechoma hederacea (66% of visits) but flower preference changed throughout the flight season, shifting to Fabaceae species and Iris foetidissima with 16 plant groups identified in pollen samples. Bees were geographically restricted to a small area within the reserve (approximately 30 ha). Although the exact location of nesting sites was not determined with certainty it is thought nests are in the loess deposits at the top of the inaccessible sea cliff face. This project suggests the presence of appropriate nesting sites may be limiting A. retusa distribution as they appear to forage on common plant species. More research is needed on the exact nesting requirements of the species.
Implications for Insect Conservation
The findings from this paper help contribute to the limited understanding of the ecology of this rare and declining species. By knowing the forage requirements of A. retusa , other areas where it is found can ensure these are present within a short distance of nest sites, hopefully ensuring the survival of individual populations and therefore the species.
