Extinction risk modeling predicts range-wide differences of climate change impact on Karner blue butterfly (Lycaeides melissa samuelis).

The Karner blue butterfly (Lycaeides melissa samuelis, or Kbb), a federally endangered species under the U.S. Endangered Species Act in decline due to habitat loss, can be further threatened by climate change. Evaluating how climate shapes the population trend of the Kbb can help in the development of adaptive management plans. Current demographic models for the Kbb incorporate in either a density-dependent or density-independent manner. We instead created mixed density-dependent and -independent (hereafter "endo-exogenous") models for Kbbs based on long-term count data of five isolated populations in the upper Midwest, United States during two flight periods (May to June and July to August) to understand how the growth rates were related to previous population densities and abiotic environmental conditions, including various macro- and micro-climatic variables. Our endo-exogenous extinction risk models showed that both density-dependent and -independent components were vital drivers of the historical population trends. However, climate change impacts were not always detrimental to Kbbs. Despite the decrease of population growth rate with higher overwinter temperatures and spring precipitations in the first generation, the growth rate increased with higher summer temperatures and precipitations in the second generation. We concluded that finer spatiotemporally scaled models could be more rewarding in guiding the decision-making process of Kbb restoration under climate change.

Potential ecological impacts of climate intervention by reflecting sunlight to cool Earth.

As the effects of anthropogenic climate change become more severe, several approaches for deliberate climate intervention to reduce or stabilize Earth's surface temperature have been proposed. Solar radiation modification (SRM) is one potential approach to partially counteract anthropogenic warming by reflecting a small proportion of the incoming solar radiation to increase Earth's albedo. While climate science research has focused on the predicted climate effects of SRM, almost no studies have investigated the impacts that SRM would have on ecological systems. The impacts and risks posed by SRM would vary by implementation scenario, anthropogenic climate effects, geographic region, and by ecosystem, community, population, and organism. Complex interactions among Earth's climate system and living systems would further affect SRM impacts and risks. We focus here on stratospheric aerosol intervention (SAI), a well-studied and relatively feasible SRM scheme that is likely to have a large impact on Earth's surface temperature. We outline current gaps in knowledge about both helpful and harmful predicted effects of SAI on ecological systems. Desired ecological outcomes might also inform development of future SAI implementation scenarios. In addition to filling these knowledge gaps, increased collaboration between ecologists and climate scientists would identify a common set of SAI research goals and improve the communication about potential SAI impacts and risks with the public. Without this collaboration, forecasts of SAI impacts will overlook potential effects on biodiversity and ecosystem services for humanity.

Preparing interdisciplinary leadership for a sustainable future.

Urgent sustainability challenges require effective leadership for inter- and trans-disciplinary (ITD) institutions. Based on the diverse experiences of 20 ITD institutional leaders and specific case studies, this article distills key lessons learned from multiple pathways to building successful programs. The lessons reflect both the successes and failures our group has experienced, to suggest how to cultivate appropriate and effective leadership, and generate the resources necessary for leading ITD programs. We present two contrasting pathways toward ITD organizations: one is to establish a new organization and the other is to merge existing organizations. We illustrate how both benefit from a real-world focus, with multiple examples of trajectories of ITD organizations. Our diverse international experiences demonstrate ways to cultivate appropriate leadership qualities and skills, especially the ability to create and foster vision beyond the status quo; collaborative leadership and partnerships; shared culture; communications to multiple audiences; appropriate monitoring and evaluation; and perseverance. We identified five kinds of resources for success: (1) intellectual resources; (2) institutional policies; (3) financial resources; (4) physical infrastructure; and (5) governing boards. We provide illustrations based on our extensive experience in supporting success and learning from failure, and provide a framework that articulates the major facets of leadership in inter- and trans-disciplinary organizations: learning, supporting, sharing, and training.

Five financial incentives to revive the Gulf of Mexico dead zone and Mississippi basin soils.

A central challenge in the Mississippi River Basin is how to continue to support profitable agricultural production, provide water supply, flood control, transportation, and other benefits, while reducing the current burden of environmental degradation. Several practices have been shown to reduce nutrient runoff and water pollution, and improve soil fertility, while often yielding profits for farmers. Yet many of these beneficial practices remain underutilized. Participants at an expert workshop identified five candidate financial mechanisms that could increase adoption of these beneficial farming practices in four focal Midwest states in the next five years: crop insurance premium subsidies, transformation of the private service provider business model, expansion and targeting of 2019 U.S. Farm Bill funding, development of new state funds, and direction of post-disaster federal funds towards habitat restoration, particularly in floodplains. This study provides rough approximations of the change in nutrient runoff and greenhouse gas (GHG) emissions, the annualized costs, and the nutrient and GHG reductions per dollar likely to result from deployment of each financial mechanism. Based upon these approximations, the adoption of these programs could reduce annual nitrate flows at the outlet of the Ohio and Upper Mississippi River Basins by 25%, surpassing the intermediate 2025 target (20% reduction) and achieving more than half of the long-term target (45% reduction) set by the Mississippi River/Gulf of Mexico Hypoxia Task Force. These approximations also illustrate that these five mechanisms could provide the same GHG reductions (∼43 Tg CO2e yr-1) as taking 12 coal-fired energy plants offline. The total cost of these five financial mechanisms is estimated at ∼$2.6 billion, or 64 g of nitrates and ∼17 kg of CO2e per dollar spent. These proposed solutions all face political, financial, cultural or institutional challenges, but with industry support, creative political action, and continued communication of both private and public benefits, they can create meaningful nutrient reductions and rebuild soils by 2022.

Climate-mediated hybrid zone movement revealed with genomics, museum collection, and simulation modeling.

Climate-mediated changes in hybridization will dramatically alter the genetic diversity, adaptive capacity, and evolutionary trajectory of interbreeding species. Our ability to predict the consequences of such changes will be key to future conservation and management decisions. Here we tested through simulations how recent warming (over the course of a 32-y period) is affecting the geographic extent of a climate-mediated developmental threshold implicated in maintaining a butterfly hybrid zone (Papilio glaucus and Papilio canadensis; Lepidoptera: Papilionidae). These simulations predict a 68-km shift of this hybrid zone. To empirically test this prediction, we assessed genetic and phenotypic changes using contemporary and museum collections and document a 40-km northward shift of this hybrid zone. Interactions between the two species appear relatively unchanged during hybrid zone movement. We found no change in the frequency of hybridization, and regions of the genome that experience little to no introgression moved largely in concert with the shifting hybrid zone. Model predictions based on climate scenarios predict this hybrid zone will continue to move northward, but with substantial spatial heterogeneity in the velocity (55-144 km/1 °C), shape, and contiguity of movement. Our findings suggest that the presence of nonclimatic barriers (e.g., genetic incompatibilities) and/or nonlinear responses to climatic gradients may preserve species boundaries as the species shift. Further, we show that variation in the geography of hybrid zone movement could result in evolutionary responses that differ for geographically distinct populations spanning hybrid zones, and thus have implications for the conservation and management of genetic diversity.

Patterns of divergence across the geographic and genomic landscape of a butterfly hybrid zone associated with a climatic gradient.

Hybrid zones are a valuable tool for studying the process of speciation and for identifying the genomic regions undergoing divergence and the ecological (extrinsic) and nonecological (intrinsic) factors involved. Here, we explored the genomic and geographic landscape of divergence in a hybrid zone between Papilio glaucus and Papilio canadensis. Using a genome scan of 28,417 ddRAD SNPs, we identified genomic regions under possible selection and examined their distribution in the context of previously identified candidate genes for ecological adaptations. We showed that differentiation was genomewide, including multiple candidate genes for ecological adaptations, particularly those involved in seasonal adaptation and host plant detoxification. The Z chromosome and four autosomes showed a disproportionate amount of differentiation, suggesting genes on these chromosomes play a potential role in reproductive isolation. Cline analyses of significantly differentiated genomic SNPs, and of species-diagnostic genetic markers, showed a high degree of geographic coincidence (81%) and concordance (80%) and were associated with the geographic distribution of a climate-mediated developmental threshold (length of the growing season). A relatively large proportion (1.3%) of the outliers for divergent selection were not associated with candidate genes for ecological adaptations and may reflect the presence of previously unrecognized intrinsic barriers between these species. These results suggest that exogenous (climate-mediated) and endogenous (unknown) clines may have become coupled and act together to reinforce reproductive isolation. This approach of assessing divergence across both the genomic and geographic landscape can provide insight about the interplay between the genetic architecture of reproductive isolation and endogenous and exogenous selection.

Merging paleobiology with conservation biology to guide the future of terrestrial ecosystems.

Conservation of species and ecosystems is increasingly difficult because anthropogenic impacts are pervasive and accelerating. Under this rapid global change, maximizing conservation success requires a paradigm shift from maintaining ecosystems in idealized past states toward facilitating their adaptive and functional capacities, even as species ebb and flow individually. Developing effective strategies under this new paradigm will require deeper understanding of the long-term dynamics that govern ecosystem persistence and reconciliation of conflicts among approaches to conserving historical versus novel ecosystems. Integrating emerging information from conservation biology, paleobiology, and the Earth sciences is an important step forward on the path to success. Maintaining nature in all its aspects will also entail immediately addressing the overarching threats of growing human population, overconsumption, pollution, and climate change.

Climate Vulnerability and Human Migration in Global Perspective.

The relationship between climate change and human migration is not homogenous and depends critically on the differential vulnerability of population and places. If places and populations are not vulnerable, or susceptible, to climate change, then the climate-migration relationship may not materialize. The key to understanding and, from a policy perspective, planning for whether and how climate change will impact future migration patterns is therefore knowledge of the link between climate vulnerability and migration. However, beyond specific case studies, little is known about this association in global perspective. We therefore provide a descriptive, country-level portrait of this relationship. We show that the negative association between climate vulnerability and international migration holds only for countries least vulnerable to climate change, which suggests the potential for trapped populations in more vulnerable countries. However, when analyzed separately by life supporting sector (food, water, health, ecosystem services, human habitat, and infrastructure) and vulnerability dimension (exposure, sensitivity, and adaptive capacity), we detect evidence of a relationship among more, but not the most, vulnerable countries. The bilateral (i.e., country-to-country) migration show that, on average, people move from countries of higher vulnerability to lower vulnerability, reducing global risk by 15%. This finding is consistent with the idea that migration is a climate adaptation strategy. Still, ~6% of bilateral migration is maladaptive with respect to climate change, with some movement toward countries with greater climate change vulnerability.

A call to insect scientists: challenges and opportunities of managing insect communities under climate change.

As climate change moves insect systems into uncharted territory, more knowledge about insect dynamics and the factors that drive them could enable us to better manage and conserve insect communities. Climate change may also require us to revisit insect management goals and strategies and lead to a new kind of scientific engagement in management decision-making. Here we make five key points about the role of insect science in aiding and crafting management decisions, and we illustrate those points with the monarch butterfly and the Karner blue butterfly, two species undergoing considerable change and facing new management dilemmas. Insect biology has a strong history of engagement in applied problems, and as the impacts of climate change increase, a reimagined ethic of entomology in service of broader society may emerge. We hope to motivate insect biologists to contribute time and effort toward solving the challenges of climate change.

Addressing potential local adaptation in species distribution models: implications for conservation under climate change.

Species distribution models (SDMs) have been criticized for involving assumptions that ignore or categorize many ecologically relevant factors such as dispersal ability and biotic interactions. Another potential source of model error is the assumption that species are ecologically uniform in their climatic tolerances across their range. Typically, SDMs treat a species as a single entity, although populations of many species differ due to local adaptation or other genetic differentiation. Not taking local adaptation into account may lead to incorrect range prediction and therefore misplaced conservation efforts. A constraint is that we often do not know the degree to which populations are locally adapted. Lacking experimental evidence, we still can evaluate niche differentiation within a species' range to promote better conservation decisions. We explore possible conservation implications of making type I or type II errors in this context. For each of two species, we construct three separate Max-Ent models, one considering the species as a single population and two of disjunct populations. Principal component analyses and response curves indicate different climate characteristics in the current environments of the populations. Model projections into future climates indicate minimal overlap between areas predicted to be climatically suitable by the whole species vs. population-based models. We present a workflow for addressing uncertainty surrounding local adaptation in SDM application and illustrate the value of conducting population-based models to compare with whole-species models. These comparisons might result in more cautious management actions when alternative range outcomes are considered.

Cold hardiness and deacclimation of overwintering Papilio zelicaon pupae.

Seasonally-acquired cold tolerance can be reversed at warm temperatures, leaving temperate ectotherms vulnerable to cold snaps. However, deacclimation, and its underlying mechanisms, has not been well-explored in insects. Swallowtail butterflies are widely distributed but in some cases their range is limited by low temperature and their cold tolerance is seasonally acquired, implying that they experience mortality resulting from deacclimation. We investigated cold tolerance and hemolymph composition of Anise swallowtail (Papilio zelicaon) pupae during overwintering in the laboratory, and after four days exposure to warm temperatures in spring. Overwintering pupae had supercooling points around -20.5°C and survived brief exposures to -30°C, suggesting partial freeze tolerance. Overwintering pupae had hemolymph osmolality of approximately 920 mOsm, imparted by high concentrations of glycerol, K⁺ and Na⁺. After exposure to spring warming, supercooling points increased to approximately -17°C, and survival of a 1h exposure to -20°C decreased from 100% to 0%. This deacclimation was associated with decreased hemolymph osmolality and reduced glycerol, trehalose, Na⁺ and Ca²âº concentrations. We compared cold tolerance of pupae to weather conditions at and beyond the species' northern range boundary. Minimum temperatures at the range boundary approached the lower lethal temperature of pupae, and were colder north of the range, suggesting that cold hardiness may set northern range limits. Minimum temperatures following warm snaps were likely to cause mortality in at least one of the past three years. Cold snaps in the spring are increasing in frequency as a result of global climate change, so are likely to be a significant source of mortality for this species, and other temperate ectotherms.

Gene expression in closely related species mirrors local adaptation: consequences for responses to a warming world.

Local adaptation of populations could preclude or slow range expansions in response to changing climate, particularly when dispersal is limited. To investigate the differential responses of populations to changing climatic conditions, we exposed poleward peripheral and central populations of two Lepidoptera to reciprocal, common-garden climatic conditions and compared their whole-transcriptome expression. We found evidence of simple population differentiation in both species, and in the species with previously identified population structure and phenotypic local adaptation, we found several hundred genes that responded in a synchronized and localized fashion. These genes were primarily involved in energy metabolism and oxidative stress, and expression levels were most divergent between populations in the same environment in which we previously detected divergence for metabolism. We found no localized genes in the species with less population structure and for which no local adaptation was previously detected. These results challenge the assumption that species are functionally similar across their ranges and poleward peripheral populations are preadapted to warmer conditions. Rather, some taxa deserve population-level consideration when predicting the effects of climate change because they respond in genetically based, distinctive ways to changing conditions.

Effects of local adaptation and interspecific competition on species' responses to climate change.

Local adaptation and species interactions have been shown to affect geographic ranges; therefore, we need models of climate impact that include both factors. To identify possible dynamics of species when including these factors, we ran simulations of two competing species using an individual-based, coupled map-lattice model using a linear climatic gradient that varies across latitude and is warmed over time. Reproductive success is governed by an individual's adaptation to local climate as well as its location relative to global constraints. In exploratory experiments varying the strength of adaptation and competition, competition reduces genetic diversity and slows range change, although the two species can coexist in the absence of climate change and shift in the absence of competitors. We also found that one species can drive the other to extinction, sometimes long after climate change ends. Weak selection on local adaptation and poor dispersal ability also caused surfing of cooler-adapted phenotypes from the expanding margin backwards, causing loss of warmer-adapted phenotypes. Finally, geographic ranges can become disjointed, losing centrally-adapted genotypes. These initial results suggest that the interplay between local adaptation and interspecific competition can significantly influence species' responses to climate change, in a way that demands future research.

Does enemy loss cause release? A biogeographical comparison of parasitoid effects on an introduced insect.

The loss of natural enemies is a key feature of species introductions and is assumed to facilitate the increased success of species in new locales (enemy release hypothesis; ERH). The ERH is rarely tested experimentally, however, and is often assumed from observations of enemy loss. We provide a rigorous test of the link between enemy loss and enemy release by conducting observational surveys and an in situ parasitoid exclusion experiment in multiple locations in the native and introduced ranges of a gall-forming insect, Neuroterus saltatorius, which was introduced poleward, within North America. Observational surveys revealed that the gall-former experienced increased demographic success and lower parasitoid attack in the introduced range. Also, a different composition of parasitoids attacked the gall-former in the introduced range. These observational results show that enemies were lost and provide support for the ERH. Experimental results, however, revealed that, while some enemy release occurred, it was not the sole driver of demographic success. This was because background mortality in the absence of enemies was higher in the native range than in the introduced range, suggesting that factors other than parasitoids limit the species in its native range and contribute to its success in its introduced range. Our study demonstrates the importance of measuring the effect of enemies in the context of other community interactions in both ranges to understand what factors cause the increased demographic success of introduced species. This case also highlights that species can experience very different dynamics when introduced into ecologically similar communities.

Thermal variability increases the impact of autumnal warming and drives metabolic depression in an overwintering butterfly.

Increases in thermal variability elevate metabolic rate due to Jensen's inequality, and increased metabolic rate decreases the fitness of dormant ectotherms by increasing consumption of stored energy reserves. Theory predicts that ectotherms should respond to increased thermal variability by lowering the thermal sensitivity of metabolism, which will reduce the impact of the warm portion of thermal variability. We examined the thermal sensitivity of metabolic rate of overwintering Erynnis propertius (Lepidoptera: Hesperiidae) larvae from a stable or variable environment reared in the laboratory in a reciprocal common garden design, and used these data to model energy use during the winters of 1973-2010 using meteorological data to predict the energetic outcomes of metabolic compensation and phenological shifts. Larvae that experienced variable temperatures had decreased thermal sensitivity of metabolic rate, and were larger than those reared at stable temperatures, which could partially compensate for the increased energetic demands. Even with depressed thermal sensitivity, the variable environment was more energy-demanding than the stable, with the majority of this demand occurring in autumn. Autumn phenology changes thus had disproportionate influence on energy consumption in variable environments, and variable-reared larvae were most susceptible to overwinter energy drain. Therefore the energetic impacts of the timing of entry into winter dormancy will strongly influence ectotherm fitness in northern temperate environments. We conclude that thermal variability drives the expression of metabolic suppression in this species; that phenological shifts will have a greater impact on ectotherms in variable thermal environments; and that E. propertius will be more sensitive to shifts in phenology in autumn than in spring. This suggests that increases in overwinter thermal variability and/or extended, warm autumns, will negatively impact all non-feeding dormant ectotherms which lack the ability to suppress their overwinter metabolic thermal sensitivity.

The influence of species interactions on geographic range change under climate change.

The fossil record tells us that many species shifted their geographic distributions during historic climate changes, but this record does not portray the complete picture of future range change in response to climate change. In particular, it does not provide information on how species interactions will affect range shifts. Therefore, we also need modern research to generate understanding of range change. This paper focuses on the role that species interactions play in promoting or preventing geographic ranges shifts under current and future climate change, and we illustrate key points using empirical case studies from an integrated study system. Case studies can have limited generalizability, but they are critical to defining possible outcomes under climate change. Our case studies emphasize host limitation that could reduce range shifts and enemy release that could facilitate range expansion. We also need improvements in modeling that explicitly consider species interactions, and this modeling can be informed by empirical research. Finally, we discuss how species interactions have implications for range management by people.

Moving forward in global-change ecology: capitalizing on natural variability.

Natural resources managers are being asked to follow practices that accommodate for the impact of climate change on the ecosystems they manage, while global-ecosystems modelers aim to forecast future responses under different climate scenarios. However, the lack of scientific knowledge about short-term ecosystem responses to climate change has made it difficult to define set conservation practices or to realistically inform ecosystem models. Until recently, the main goal for ecologists was to study the composition and structure of communities and their implications for ecosystem function, but due to the probable magnitude and irreversibility of climate-change effects (species extinctions and loss of ecosystem function), a shorter term focus on responses of ecosystems to climate change is needed. We highlight several underutilized approaches for studying the ecological consequences of climate change that capitalize on the natural variability of the climate system at different temporal and spatial scales. For example, studying organismal responses to extreme climatic events can inform about the resilience of populations to global warming and contribute to the assessment of local extinctions. Translocation experiments and gene expression are particular useful to quantitate a species' acclimation potential to global warming. And studies along environmental gradients can guide habitat restoration and protection programs by identifying vulnerable species and sites. These approaches identify the processes and mechanisms underlying species acclimation to changing conditions, combine different analytical approaches, and can be used to improve forecasts of the short-term impacts of climate change and thus inform conservation practices and ecosystem models in a meaningful way.

Impact of an invasive oak gall wasp on a native butterfly: a test of plant-mediated competition.

Phytophagous insects commonly interact through shared host plants. These interactions, however, do not occur in accordance with traditional paradigms of competition, and competition in phytophagous insects is still being defined. It remains unclear, for example, if particular guilds of insects are superior competitors or important players in structuring insect communities. Gall-forming insects are likely candidates for such superior competitors because of their ability to manipulate host plants, but their role as competitors is understudied. We investigate the effect of invasive populations of an oak gall wasp, Neuroterus saltatorius, on a native specialist butterfly, Erynnis propertius, as mediated by their shared host plant, Quercus garryana. This gall wasp occurs at high densities in its introduced range, where we stocked enclosures with caterpillars on trees that varied in gall wasp density. Biomass production of butterflies was lower in enclosures on high-density than on low-density trees because overwintering caterpillars were smaller, and fewer of them eclosed into adults the following spring. To see if the gall wasp induced changes in foliar quality, we measured host plant quality before and after gall induction on 30 trees each at two sites. We found a positive relationship between gall wasp density and the percentage change in foliar C:N, a negative relationship between gall wasp density and the percentage change in foliar water at one site, and no relationship between the percentage change in protein-binding capacity (i.e., phenolics) and gall-wasp density. Additionally, there was a negative relationship between foliar quality and butterfly performance. Our results provide evidence for a plant-mediated impact of an invasive oak gall wasp on a native butterfly and suggest that gall wasps could act as superior competitors, especially when they occur at high densities.

Population-level transcriptome sequencing of nonmodel organisms Erynnis propertius and Papilio zelicaon.

BACKGROUND: Several recent studies have demonstrated the use of Roche 454 sequencing technology for de novo transcriptome analysis. Low error rates and high coverage also allow for effective SNP discovery and genetic diversity estimates. However, genetically diverse datasets, such as those sourced from natural populations, pose challenges for assembly programs and subsequent analysis. Further, estimating the effectiveness of transcript discovery using Roche 454 transcriptome data is still a difficult task. RESULTS: Using the Roche 454 FLX Titanium platform, we sequenced and assembled larval transcriptomes for two butterfly species: the Propertius duskywing, Erynnis propertius (Lepidoptera: Hesperiidae) and the Anise swallowtail, Papilio zelicaon (Lepidoptera: Papilionidae). The Expressed Sequence Tags (ESTs) generated represent a diverse sample drawn from multiple populations, developmental stages, and stress treatments. Despite this diversity, > 95% of the ESTs assembled into long (> 714 bp on average) and highly covered (> 9.6x on average) contigs. To estimate the effectiveness of transcript discovery, we compared the number of bases in the hit region of unigenes (contigs and singletons) to the length of the best match silkworm (Bombyx mori) protein--this "ortholog hit ratio" gives a close estimate on the amount of the transcript discovered relative to a model lepidopteran genome. For each species, we tested two assembly programs and two parameter sets; although CAP3 is commonly used for such data, the assemblies produced by Celera Assembler with modified parameters were chosen over those produced by CAP3 based on contig and singleton counts as well as ortholog hit ratio analysis. In the final assemblies, 1,413 E. propertius and 1,940 P. zelicaon unigenes had a ratio > 0.8; 2,866 E. propertius and 4,015 P. zelicaon unigenes had a ratio > 0.5. CONCLUSIONS: Ultimately, these assemblies and SNP data will be used to generate microarrays for ecoinformatics examining climate change tolerance of different natural populations. These studies will benefit from high quality assemblies with few singletons (less than 26% of bases for each assembled transcriptome are present in unassembled singleton ESTs) and effective transcript discovery (over 6,500 of our putative orthologs cover at least 50% of the corresponding model silkworm gene).