Butterflies in the highly fragmented prairies of central Iowa: how the landscape affects population isolation

Ever since MacArthur and Wilson presented the Theory of Island Biogeography in 1963, there has been considerable concern about the ability of a system of isolated reserves to retain species diversity or viable populations over extended periods of time. Despite this interest, very little is known about how readily individuals move around in their landscape. Edges and corridors have both been targeted as landscape features that may have important consequences for directing movement and thereby affecting isolation. Edges may form a barrier to movement reducing the probability that an individual will leave a patch. For individuals that have left, they must find a new patch, and corridors may help direct them there. We studied the responses of butterflies to these two important landscape features.;We tracked the responses of two butterflies: the regal fritillary (Speyeria idalia) and the monarch (Danaus plexippus) at four edge types (crop, field, road, treeline) in the highly fragmented prairies of central Iowa. S. idalia responded strongly to all except road edges, but their responses were strongly influenced by conspecific density. Individuals in high density areas were less likely to leave the prairie. Surprisingly, S. idalia showed a very strong response to edges with very subtle differences in vegetation structure, such as a prairie and a pasture. D. plexippus responded only to treeline edges. Their responses were more influenced by factors such as wind, nectar availability, and time of year. We also explored the conservation value of roadside vegetation, both as additional habitat and corridors or stepping stones between prairies.;We examined the abundances of common, open area butterflies and the species richness of grassland specialists in three types of roadsides: grassy, weedy, and restored to prairie. We also considered how distance from a source prairie affected patterns. Two common butterflies, D. plexippus and Everes comyntas showed a significant relationship with roadside type, but not distance. There were no significant associations between grassland species richness and any factor we measured. More intensive sampling may be necessary, though, to detect an effect

Heatwaves and Novel Host Consumption Increase Overwinter Mortality of an Imperiled Wetland Butterfly

Disruptive effects of climate change include range shifts, phenological mismatches among consumers and producers, and population declines. While these biological alterations have been widely documented, studies identifying specific mechanisms linking climate change to population declines are scarce. Extreme events, such as heatwaves can have devastating effects on living organisms and are increasing in frequency as Earth warms. Hence, understanding the effects of heatwaves on insects is necessary to inform conservation efforts and to develop predictions of population dynamics under future climate scenarios. Here, we experimentally evaluated the effects of heatwaves on the survival and phenology of the Baltimore Checkerspot (Euphydryas phaeton phaeton), a wetland butterfly with imperiled populations that has incorporated a novel host. We performed laboratory manipulations (implementing realistic temperature regimes) to assess the effect of heatwaves during summer and winter on the survival and phenology of E. p. phaeton. In addition, we analyzed historical temperature records to quantify the incidence of heatwaves within E. p. phaeton’s range to assess their potential role in the decline of southeastern populations. We found that winter heatwaves with maximum temperatures of 20◦C can have more devastating effects on survival than summer heatwaves (up to 41◦C). Eggs endured acute heat stress during summer with no significant effects on phenology and survival; similarly, pre-overwintering larvae were robust to heatwave exposure, as only themost intense heatwave treatment reduced their survival (37% reduction compared to control conditions). By contrast, dormant larvae were the most vulnerable stage, as they lost from 2 to 6% of their body mass after a three-day summer heatwave. Furthermore, their exposure to winter heatwaves resulted in 75 to 100% mortality. Feeding on the native host provided higher resilience under thermal stress than feeding on the invasive, recently acquired host. Finally, both heatwave incidence and severity have increased in the southern range of E. p. phaeton in the period from 1894 to 2011. We show that warm winter days induced severe mortality, providing a mechanistic explanation of how climate change can trigger population declines in E. p. phaeton and other insects

Inclusion of Host Quality Data Improves Predictions of Herbivore Phenology

Understanding the correspondence between ambient temperature and insect development is necessary to forecast insect phenology under novel environments. In the face of climate change, both conservation and pest control efforts require accurate phenological predictions. Here, we compare a suite of degree-day models to assess their ability to predict the phenology of a common, oligophagous butterfly, the silver-spotted skipper, Epargyreus clarus (Cramer) (Lepidoptera: Hesperiidae). To estimate model parameters, we used development time of eggs and larvae reared in the laboratory at six constant temperatures ranging from 8 to 38 °C and on two host plants of contrasting quality (kudzu and wisteria). We employed three approaches to determine the base temperature to calculate degree days: linear regression, modified reduced major axis regression, and application of a generic base temperature value of 10 °C, which is commonly used in the absence of laboratory data. To calculate the number of degree days required to complete a developmental stage, we used data from caterpillars feeding on high- and low-quality hosts, both in the field and in the laboratory. To test model accuracy, we predicted development time of seven generations of larvae reared in the field on the same host plants across 3 years (2014–2016). To compare performance among models, we regressed predicted vs. observed development time, and found that r2 values were significantly larger when accounting for host plant quality. The accuracy of development time predictions varied across the season, with estimates of the first two generations being more accurate than estimates of the third generation, when ambient temperatures dropped outside the range in which development rate and temperature have a linear relationship. Overall, we show that accounting for variation in host plant quality when calculating development time in the field is more important than the choice of the base temperature for calculating degree days

How Development and Survival Combine to Determine the Thermal Sensitivity of Insects

Thermal performance curves (TPCs) depict variation in vital rates in response to temperature and have been an important tool to understand ecological and evolutionary constraints on the thermal sensitivity of ectotherms. TPCs allow for the calculation of indicators of thermal tolerance, such as minimum, optimum, and maximum temperatures that allow for a given metabolic function. However, these indicators are computed using only responses from surviving individuals, which can lead to underestimation of deleterious effects of thermal stress, particularly at high temperatures. Here, we advocate for an integrative frame- work for assessing thermal sensitivity, which combines both vital rates and survival probabilities, and focuses on the temperature interval that allows for population persistence. Using a collated data set of Lepidopteran development rate and survival measured on the same individuals, we show that development rate is generally limiting at low temperatures, while survival is limiting at high temperatures. We also uncover differences between life stages and across latitudes, with extended survival at lower temperatures in temperate regions. Our combined performance metric demonstrates similar thermal breadth in temperate and tropical individuals, an effect that only emerges from integration of both development and survival trends. We discuss the benefits of using this framework in future predictive and management contexts

Insecticides, more than herbicides, land use, and climate, are associated with declines in butterfly species richness and abundance in the American Midwest

Mounting evidence shows overall insect abundances are in decline globally. Habitat loss, climate change, and pesticides have all been implicated, but their relative effects have never been evaluated in a comprehensive large-scale study. We harmonized 17 years of land use, climate, multiple classes of pesticides, and butterfly survey data across 81 counties in five states in the US Midwest. We find community-wide declines in total butterfly abundance and species richness to be most strongly associated with insecticides in general, and for butterfly species richness the use of neonicotinoid-treated seeds in particular. This included the abundance of the migratory monarch (Danaus plexippus), whose decline is the focus of intensive debate and public concern. Insect declines cannot be understood without comprehensive data on all putative drivers, and the 2015 cessation of neonicotinoid data releases in the US will impede future research.This article is published as Van Deynze B, Swinton SM, Hennessy DA, Haddad NM, Ries L (2024) Insecticides, more than herbicides, land use, and climate, are associated with declines in butterfly species richness and abundance in the American Midwest. PLoS ONE 19(6): e0304319. https://doi.org/10.1371/journal.pone.0304319. © 2024 Van Deynze et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited

How development and survival combine to determine the thermal sensitivity of insects

Thermal performance curves (TPCs) depict variation in vital rates in response to temperature and have been an important tool to understand ecological and evolutionary constraints on the thermal sensitivity of ectotherms. TPCs allow for the calculation of indicators of thermal tolerance, such as minimum, optimum, and maximum temperatures that allow for a given metabolic function. However, these indicators are computed using only responses from surviving individuals, which can lead to underestimation of deleterious effects of thermal stress, particularly at high temperatures. Here, we advocate for an integrative framework for assessing thermal sensitivity, which combines both vital rates and survival probabilities, and focuses on the temperature interval that allows for population persistence. Using a collated data set of Lepidopteran development rate and survival measured on the same individuals, we show that development rate is generally limiting at low temperatures, while survival is limiting at high temperatures. We also uncover differences between life stages and across latitudes, with extended survival at lower temperatures in temperate regions. Our combined performance metric demonstrates similar thermal breadth in temperate and tropical individuals, an effect that only emerges from integration of both development and survival trends. We discuss the benefits of using this framework in future predictive and management contexts

A regionally informed abundance index for supporting integrative analyses across butterfly monitoring schemes

1. 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. 2. 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). 3. 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. 4. 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

Is habitat fragmentation good for biodiversity?

Habitat loss is a primary threat to biodiversity across the planet, yet contentious debate has ensued on the importance of habitat fragmentation ‘per se’ (i.e., altered spatial configuration of habitat for a given amount of habitat loss). Based on a review of landscape-scale investigations, Fahrig (2017; Ecological responses to habitat fragmentation per se. Annual Review of Ecology, Evolution, and Systematics 48:1-23) reports that biodiversity responses to habitat fragmentation ‘per se’ are more often positive rather than negative and concludes that the widespread belief in negative fragmentation effects is a ‘zombie idea’. We show that Fahrig's conclusions are drawn from a narrow and potentially biased subset of available evidence, which ignore much of the observational, experimental and theoretical evidence for negative effects of altered habitat configuration. We therefore argue that Fahrig's conclusions should be interpreted cautiously as they could be misconstrued by policy makers and managers, and we provide six arguments why they should not be applied in conservation decision-making. Reconciling the scientific disagreement, and informing conservation more effectively, will require research that goes beyond statistical and correlative approaches. This includes a more prudent use of data and conceptual models that appropriately partition direct vs indirect influences of habitat loss and altered spatial configuration, and more clearly discriminate the mechanisms underpinning any changes. Incorporating these issues will deliver greater mechanistic understanding and more predictive power to address the conservation issues arising from habitat loss and fragmentation

Multitaxonomic Diversity Patterns along a Desert Riparian–Upland Gradient

Riparian areas are noted for their high biodiversity, but this has rarely been tested across a wide range of taxonomic groups. We set out to describe species richness, species abundance, and community similarity patterns for 11 taxonomic groups (forbs & grasses, shrubs, trees, solpugids, spiders, scarab beetles, butterflies, lizards, birds, rodents, and mammalian carnivores) individually and for all groups combined along a riparian–upland gradient in semiarid southeastern Arizona, USA. Additionally, we assessed whether biological characteristics could explain variation in diversity along the gradient using five traits (trophic level, body size, life span, thermoregulatory mechanism, and taxonomic affiliation). At the level of individual groups diversity patterns varied along the gradient, with some having greater richness and/or abundance in riparian zones whereas others were more diverse and/or abundant in upland zones. Across all taxa combined, riparian zones contained significantly more species than the uplands. Community similarity between riparian and upland zones was low, and beta diversity was significantly greater than expected for most taxonomic groups, though biological traits explained little variance in diversity along the gradient. These results indicate heterogeneity amongst taxa in how they respond to the factors that structure ecological communities in riparian landscapes. Nevertheless, across taxonomic groups the overall pattern is one of greater species richness and abundance in riparian zones, coupled with a distinct suite of species