Phenology and foraging bias contribute to sex-specific foraging patterns in the rare declining butterfly Argynnis idalia idalia.

Variation in pollinator foraging behavior can influence pollination effectiveness, community diversity, and plant-pollinator network structure. Although effects of interspecific variation have been widely documented, studies of intraspecific variation in pollinator foraging are relatively rare. Sex-specific differences in resource use are a strong potential source of intraspecific variation, especially in species where the phenology of males and females differ. Differences may arise from encountering different flowering communities, sex-specific traits, nutritional requirements, or a combination of these factors. We evaluated sex-specific foraging patterns in the eastern regal fritillary butterfly (Argynnis idalia idalia), leveraging a 21-year floral visitation dataset. Because A. i. idalia is protandrous, we determined whether foraging differences were due to divergent phenology by comparing visitation patterns between the entire season with restricted periods of male-female overlap. We quantified nectar carbohydrate and amino acid contents of the most visited plant species and compared those visited more frequently by males versus females. We demonstrate significant differences in visitation patterns between male and female A. i. idalia over two decades. Females visit a greater diversity of species, while dissimilarity in foraging patterns between sexes is persistent and comparable to differences between species. While differences are diminished or absent in some years during periods of male-female overlap, remaining signatures of foraging dissimilarity during implicate mechanisms other than phenology. Nectar of plants visited more by females had greater concentrations of total carbohydrates, glucose, and fructose and individual amino acids than male-associated plants. Further work can test whether nutritional differences are a cause of visitation patterns or consequence, reflecting seasonal shifts in the nutritional landscape encountered by male and female A. i. idalia. We highlight the importance of considering sex-specific foraging patterns when studying interaction networks, and in making conservation management decisions for this at-risk butterfly and other species exhibiting strong intraspecific variation.

Not Too Warm, Not Too Cold: Thermal Treatments to Slightly Warmer or Colder Conditions from Mother's Origin Can Enhance Performance of Montane Butterfly Larvae.

Climate change alters organismal performance via shifts in temperature. However, we know little about the relative fitness impacts of climate variability and how cold-adapted ectotherms mediate these effects. Here, we advance the field of climate change biology by directly testing for species performance, considering the effects of different thermal environments at the first developmental stage of larvae. We conducted our experiments in climatic chambers (2019-2020) using five cold-adapted butterflies of the genus Erebia (Erebia aethiops, Erebia cassioides, Erebia manto, Erebia tyndarus, Erebia nivalis). Larvae were reared indoors and were treated with higher and lower temperatures than those of their mothers' origins. Overall, we found evidence of better performance at warmer temperatures and a decreased performance at lower temperatures, and larvae were able to tolerate small temperature changes from mother's origin. Warmer conditions, however, were unfavorable for E. nivalis, indicative of its limited elevational range and its poor ability to mediate a variety of thermal conditions. Further, larvae generally performed poorly where there was a large difference in thermal regimen from that of their maternal origin. Future efforts should include additional life history stages and focus on a more mechanistic understanding of species thermal tolerance. Such studies could increase the realism of predicted responses to climate change and could account for asynchronous changes in species development, which will alter community composition and ecosystem functioning.

Species traits affect phenological responses to climate change in a butterfly community.

Diverse taxa have undergone phenological shifts in response to anthropogenic climate change. While such shifts generally follow predicted patterns, they are not uniform, and interspecific variation may have important ecological consequences. We evaluated relationships among species' phenological shifts (mean flight date, duration of flight period), ecological traits (larval trophic specialization, larval diet composition, voltinism), and population trends in a butterfly community in Pennsylvania, USA, where the summer growing season has become warmer, wetter, and longer. Data were collected over 7-19 years from 18 species or species groups, including the extremely rare eastern regal fritillary Speyeria idalia idalia. Both the direction and magnitude of phenological change over time was linked to species traits. Polyphagous species advanced and prolonged the duration of their flight period while oligophagous species delayed and shortened theirs. Herb feeders advanced their flight periods while woody feeders delayed theirs. Multivoltine species consistently prolonged flight periods in response to warmer temperatures, while univoltine species were less consistent. Butterflies that shifted to longer flight durations, and those that had polyphagous diets and multivoltine reproductive strategies tended to decline in population. Our results suggest species' traits shape butterfly phenological responses to climate change, and are linked to important community impacts.

Butterfly phenology in Mediterranean mountains using space-for-time substitution.

Inferring species' responses to climate change in the absence of long-term time series data is a challenge, but can be achieved by substituting space for time. For example, thermal elevational gradients represent suitable proxies to study phenological responses to warming. We used butterfly data from two Mediterranean mountain areas to test whether mean dates of appearance of communities and individual species show a delay with increasing altitude, and an accompanying shortening in the duration of flight periods. We found a 14-day delay in the mean date of appearance per kilometer increase in altitude for butterfly communities overall, and an average 23-day shift for 26 selected species, alongside average summer temperature lapse rates of 3°C per km. At higher elevations, there was a shortening of the flight period for the community of 3 days/km, with an 8.8-day average decline per km for individual species. Rates of phenological delay differed significantly between the two mountain ranges, although this did not seem to result from the respective temperature lapse rates. These results suggest that climate warming could lead to advanced and lengthened flight periods for Mediterranean mountain butterfly communities. However, although multivoltine species showed the expected response of delayed and shortened flight periods at higher elevations, univoltine species showed more pronounced delays in terms of species appearance. Hence, while projections of overall community responses to climate change may benefit from space-for-time substitutions, understanding species-specific responses to local features of habitat and climate may be needed to accurately predict the effects of climate change on phenology.

Prescribed fire maintains host plants of a rare grassland butterfly.

As grassland ecosystems transform globally due to anthropogenic pressures, improvements in our understanding of the effect of management on rare and threatened species in such landscapes has become urgent. Although prescribed fire is a very efficient tool for habitat restoration and endangered species management on fire-adapted ecosystems, the specific mechanisms underlying potential effects of burning on population dynamics of butterfly host plants are poorly understood. We analyzed a 12-year dataset (2004-2015), combining violet abundance, habitat physiognomy and fire history data from a fire-managed system, to determine factors influencing the spatiotemporal distribution and abundance of violets (Viola spp.), the host plants of the threatened eastern regal fritillary (Speyeria idalia idalia) butterfly. Our results demonstrate a critical role for fire in driving both presence and abundance of violets, suggesting management with prescribed fires can effectively promote butterfly host plants. In addition, we determined the character of habitats associated with violet presence and abundance, in particular a strong positive association with biocrusts. These results provide a roadmap for efficient site selection to increase the effectiveness of restoration efforts, including assessment of potential reintroduction sites for regal fritillary and other grassland butterflies and actions to promote the re-establishment of host plants in these sites.

Discovering the still unexplored arachnofauna of the National Park of Dadia-Lefkimi-Soufli, NE Greece: a taxonomic review with description of new species.

The National Park of Dadia in NE Greece (Thrace) was established as a nature reserve in 1980, mainly due to its great diversity in birds of prey. Since then many studies have taken place, focusing on other birds, reptiles, amphibians and some invertebrates (grasshoppers, beetles and butterflies), but up to now none was conducted on spiders. The aim of the present paper was to create the first extensive checklist on the spiders of this important natural reserve. For this purpose, pitfall traps were set in 15 sites located in and around the National Park, resulting in a large spider collection. The results of the taxonomical revision of this collection are here presented, giving rise to 132 species in total, which belong to 24 families. Of them, 11 species (Centromerus valkanovi Deltshev, 1983, Crosbyarachne silvestris (Georgescu, 1973), Ipa terrenus (L. Koch, 1879), Sintula spiniger (Balogh, 1935), Tenuiphantes floriana (van Helsdingen, 1977), Alopecosa taeniopus (Kulczyñski, 1895), Liocranum rupicola (Walckenaer, 1830), Zodarion turcicum Wunderlich, 1980, Gnaphosa modestior Kulczyñski, 1897, Philodromus krausi Muster & Thaler, 2004, Cozyptila thaleri Marusik & Kovblyuk, 2005) are new records for the Greek territory. Seven species (Dysdera kati sp. n., Dysdera krisis sp. n., Harpactea ice sp. n., Harpactea wolfgangi sp. n.-Dysderidae, Phrurolithus thracia sp. n.-Phrurolithidae, Zodarion beroni sp. n.-Zodariidae, Drassyllus dadia sp. n.-Gnaphosidae) are here proposed as new species for science.

Signals of climate change in butterfly communities in a Mediterranean protected area.

The European protected-area network will cease to be efficient for biodiversity conservation, particularly in the Mediterranean region, if species are driven out of protected areas by climate warming. Yet, no empirical evidence of how climate change influences ecological communities in Mediterranean nature reserves really exists. Here, we examine long-term (1998-2011/2012) and short-term (2011-2012) changes in the butterfly fauna of Dadia National Park (Greece) by revisiting 21 and 18 transects in 2011 and 2012 respectively, that were initially surveyed in 1998. We evaluate the temperature trend for the study area for a 22-year-period (1990-2012) in which all three butterfly surveys are included. We also assess changes in community composition and species richness in butterfly communities using information on (a) species' elevational distributions in Greece and (b) Community Temperature Index (calculated from the average temperature of species' geographical ranges in Europe, weighted by species' abundance per transect and year). Despite the protected status of Dadia NP and the subsequent stability of land use regimes, we found a marked change in butterfly community composition over a 13 year period, concomitant with an increase of annual average temperature of 0.95°C. Our analysis gave no evidence of significant year-to-year (2011-2012) variability in butterfly community composition, suggesting that the community composition change we recorded is likely the consequence of long-term environmental change, such as climate warming. We observe an increased abundance of low-elevation species whereas species mainly occurring at higher elevations in the region declined. The Community Temperature Index was found to increase in all habitats except agricultural areas. If equivalent changes occur in other protected areas and taxonomic groups across Mediterranean Europe, new conservation options and approaches for increasing species' resilience may have to be devised.