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.

Causes and consequences of variation in heterospecific pollen receipt in Oenothera fruticosa.

PREMISE: Heterospecific pollen transfer, the transfer of pollen between species, is common among co-flowering plants, yet the amount of pollen received is extremely variable among species. Intraspecific variation in heterospecific pollen receipt can be even greater, but we lack an understanding of its causes and fitness consequences in wild populations. METHODS: We examined potential drivers of variation in heterospecific pollen receipt in Oenothera fruticosa. We evaluated the relationship between heterospecific and conspecific pollen receipt and considered how visitation by different pollinator groups, local floral neighborhood composition, and flowering phenology affect the total amount and proportion of heterospecific pollen received. Finally, we tested whether variation in heterospecific pollen receipt translated into lower seed production. RESULTS: Heterospecific pollen was ubiquitous on O. fruticosa stigmas, but the amount received was highly variable and unrelated to conspecific pollen receipt. Heterospecific pollen receipt depended on pollinator type, the proportion of nearby conspecific flowers, and flowering date. Significant interactions revealed that the effects of pollinator type and neighborhood were not independent, further contributing to variation in heterospecific pollen. Naturally occurring levels of heterospecific pollen were sufficient to negatively impact seed set, but large amounts of conspecific pollen counteracted this detrimental effect. CONCLUSIONS: Although selection could act on floral traits that attract quality pollinators and promote synchronous flowering in O. fruticosa, the risk of heterospecific pollen is equally dependent on local floral context. This work highlights how extrinsic and intrinsic factors contribute to intraspecific variation in heterospecific pollen receipt in wild plants, with significant fitness consequences.

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.

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.