Inclusion of trophic interactions increases the vulnerability of an alpine butterfly species to climate change.

Climate change is expected to have significant and complex impacts on ecological communities. In addition to direct effects of climate on species, there can also be indirect effects through an intermediary species, such as in host-plant interactions. Indirect effects are expected to be more pronounced in alpine environments because these ecosystems are sensitive to temperature changes and there are limited areas for migration of both species (i.e. closed systems), and because of simpler trophic interactions. We tested the hypothesis that climate change will reduce the range of an alpine butterfly (Parnassius smintheus) because of indirect effects through its host plant (Sedum sp.). To test for direct and indirect effects, we used the simulations of climate change to assess the distribution of P. smintheus with and without Sedum sp. We also compared the projected ranges of P. smintheus to four other butterfly species that are found in the alpine, but that are generalists feeding on many plant genera. We found that P. smintheus gained distributional area in climate-only models, but these gains were significantly reduced with the inclusion of Sedum sp. and in dry-climate scenarios which resulted in a reduction in net area. When compared to the more generalist butterfly species, P. smintheus exhibited the largest loss in suitable habitat. Our findings support the importance of including indirect effects in modelling species distributions in response to climate change. We highlight the potentially large and still neglected impacts climate change can have on the trophic structure of communities, which can lead to significant losses of biodiversity. In the future, communities will continue to favour species that are generalists as climate change induces asynchronies in the migration of species.

Demographic fluctuations lead to rapid and cyclic shifts in genetic structure among populations of an alpine butterfly, Parnassius smintheus.

Many populations, especially in insects, fluctuate in size, and periods of particularly low population size can have strong effects on genetic variation. Effects of demographic bottlenecks on genetic diversity of single populations are widely documented. Effects of bottlenecks on genetic structure among multiple interconnected populations are less studied, as are genetic changes across multiple cycles of demographic collapse and recovery. We take advantage of a long-term data set comprising demographic, genetic and movement data from a network of populations of the butterfly, Parnassius smintheus, to examine the effects of fluctuating population size on spatial genetic structure. We build on a previous study that documented increased genetic differentiation and loss of spatial genetic patterns (isolation by distance and by intervening forest cover) after a network-wide bottleneck event. Here, we show that genetic differentiation was reduced again and spatial patterns returned to the system extremely rapidly, within three years (i.e. generations). We also show that a second bottleneck had similar effects to the first, increasing differentiation and erasing spatial patterns. Thus, bottlenecks consistently drive random divergence of allele frequencies among populations in this system, but these effects are rapidly countered by gene flow during demographic recovery. Our results reveal a system in which the relative influence of genetic drift and gene flow continually shift as populations fluctuate in size, leading to cyclic changes in genetic structure. Our results also suggest caution in the interpretation of patterns of spatial genetic structure, and its association with landscape variables, when measured at only a single point in time.

Danish first aid books compliance with the new evidence-based non-resuscitative first aid guidelines.

BACKGROUND: The European Resuscitation Council (ERC) released new guidelines on resuscitation in 2015. For the first time, the guidelines included a separate chapter on first aid for laypersons. We analysed the current major Danish national first aid books to identify potential inconsistencies between the current books and the new evidence-based first aid guidelines. METHODS: We identified first aid books from all the first aid courses offered by major Danish suppliers. Based on the new ERC first aid guidelines, we developed a checklist of 26 items within 16 different categories to assess the content; this checklist was adapted following the principle of mutually exclusive and collectively exhaustive questioning. To assess the agreement between four raters, Fleiss' kappa test was used. Items that did not reach an acceptable kappa score were excluded. RESULTS: We evaluated 10 first aid books used for first aid courses and published between 2009 and 2015. The content of the books complied with the new in 38% of the answers. In 12 of the 26 items, there was less than 50% consistency. These items include proximal pressure points and elevation of extremities for the control of bleeding, use of cervical collars, treatment for an open chest wound, burn dressing, dental avulsion, passive leg raising, administration of bronchodilators, adrenaline, and aspirin. CONCLUSIONS: Danish course material showed significant inconsistencies with the new evidence-based first aid guidelines. The new knowledge from the evidence-based guidelines should be incorporated into revised and updated first aid course material.

Connectivity rescues genetic diversity after a demographic bottleneck in a butterfly population network.

Demographic bottlenecks that occur when populations fluctuate in size erode genetic diversity, but that diversity can be recovered through immigration. Connectivity among populations and habitat patches in the landscape enhances immigration and should in turn facilitate recovery of genetic diversity after a sudden reduction in population size. For the conservation of genetic diversity, it may therefore be particularly important to maintain connectivity in the face of factors that increase demographic instability, such as climate change. However, a direct link between connectivity and recovery of genetic diversity after a demographic bottleneck has not been clearly demonstrated in an empirical system. Here, we show that connectivity of habitat patches in the landscape contributes to the maintenance of genetic diversity after a demographic bottleneck. We were able to monitor genetic diversity in a network of populations of the alpine butterfly, Parnassius smintheus, before, during, and after a severe reduction in population size that lasted two generations. We found that allelic diversity in the network declined after the demographic bottleneck but that less allelic diversity was lost from populations occupying habitat patches with higher connectivity. Furthermore, the effect of connectivity on allelic diversity was important during the demographic recovery phase. Our results demonstrate directly the ability of connectivity to mediate the rescue of genetic diversity in a natural system.

Landscape structure and the genetic effects of a population collapse.

Both landscape structure and population size fluctuations influence population genetics. While independent effects of these factors on genetic patterns and processes are well studied, a key challenge is to understand their interaction, as populations are simultaneously exposed to habitat fragmentation and climatic changes that increase variability in population size. In a population network of an alpine butterfly, abundance declined 60-100% in 2003 because of low over-winter survival. Across the network, mean microsatellite genetic diversity did not change. However, patch connectivity and local severity of the collapse interacted to determine allelic richness change within populations, indicating that patch connectivity can mediate genetic response to a demographic collapse. The collapse strongly affected spatial genetic structure, leading to a breakdown of isolation-by-distance and loss of landscape genetic pattern. Our study reveals important interactions between landscape structure and temporal demographic variability on the genetic diversity and genetic differentiation of populations. Projected future changes to both landscape and climate may lead to loss of genetic variability from the studied populations, and selection acting on adaptive variation will likely occur within the context of an increasing influence of genetic drift.

Trophic cascades: linking ungulates to shrub-dependent birds and butterflies.

1. Studies demonstrating trophic cascades through the loss of top-down regulatory processes in productive and biologically diverse terrestrial ecosystems are limited. 2. Elk Island National Park, Alberta and surrounding protected areas have a wide range of ungulate density due to the functional loss of top predators, management for high ungulate numbers and variable hunting pressure. This provides an ideal setting for studying the effects of hyper-abundant ungulates on vegetation and shrub-dependent bird and butterfly species. 3. To examine the cascading effects of high ungulate density, we quantified vegetation characteristics and abundances of yellow warbler Dendroica petechia and Canadian tiger swallowtail Papilio canadensis under different ungulate density in and around Elk Island National Park. 4. Using Structural Equation Models we found that ungulate density was inversely related to shrub cover, whereas shrub cover was positively related to yellow warbler abundance. In addition, chokecherry Prunus virginiana abundance was inversely related to browse impact but positively related to P. canadensis abundance. 5. These results demonstrate a cascade resulting from hyper-abundant ungulates on yellow warblers and Canadian tiger swallowtails through reductions in shrub cover and larval host plant density. The combined effect of the functional loss of top predators and management strategies that maintain high ungulate numbers can decouple top-down regulation of productive temperate ecosystems.

Variability in winter climate and winter extremes reduces population growth of an alpine butterfly.

We examined the long-term, 15-year pattern of population change in a network of 21 Rocky Mountain populations of Parnassius smintheus butterflies in response to climatic variation. We found that winter values of the broadscale climate variable, the Pacific Decadal Oscillation (PDO) index, were a strong predictor of annual population growth, much more so than were endogenous biotic factors related to population density. The relationship between PDO and population growth was nonlinear. Populations declined in years with extreme winter PDO values, when there were either extremely warm or extremely cold sea surface temperatures in the eastern Pacific relative to that in the western Pacific. Results suggest that more variable winters, and more frequent extremely cold or warm winters, will result in more frequent decline of these populations, a pattern exacerbated by the trend for increasingly variable winters seen over the past century.

Treeline proximity alters an alpine plant-herbivore interaction.

Rising treeline threatens the size and contiguity of alpine meadows worldwide. As trees encroach into previously open habitat, the movement and population dynamics of above-treeline alpine species may be disrupted. This process is well documented in studies of the Rocky Mountain apollo butterfly (Parnassius smintheus). However, subtler consequences of treeline rise remain poorly understood. In this study, we examine whether treeline proximity affects feeding behaviour of P. smintheus larvae, due to altered habitat affecting the distribution and availability of their host plant, lance-leaved stonecrop (Sedum lanceolatum). Understanding differential larval exploitation of food resources in relation to the treeline is an important step in predicting the consequences of continued treeline rise. Parnassius smintheus larvae feed more intensively on S. lanceolatum away from the treeline despite the relative paucity of hosts in these areas, and despite higher fitness penalties associated with the plant's herbivory-induced chemical defenses. Sedum lanceolatum growing near the treeline is less attractive, and therefore represents a less significant resource for P. smintheus larvae than its abundance might imply. If treeline rise continues, we suggest that this pattern of altered resource exploitation may represent a mechanism by which larvae are adversely affected even while adult movement among and within meadows appears sufficient for maintaining population health, and total host availability seems ample.

Local extinction synchronizes population dynamics in spatial networks.

Spatial population theory predicts that synchrony in the dynamics of local populations should decrease as dispersal among populations decreases. Thus, it would be expected that the extinction of local populations and the attendant loss of immigrants to surrounding populations would reduce synchrony. We tested this hypothesis through a large-scale experiment, simulation of the experimental system and general models. Experimental removal of two adjacent subpopulations of the Rocky Mountain Apollo butterfly, Parnassius smintheus within a network consisting of 15 other local populations resulted in a decrease in immigration to surrounding populations that was proportional to their connectivity to the removal populations. These populations also showed a significant increase in synchrony during population removal. The spatial extent of the synchrony showed good agreement with the predicted loss of immigrants owing to the removals. Simulation of the Parnassius system showed a similar short-term result and also indicated that permanent loss of populations produces structural changes increasing synchrony. General models indicate that an increase in synchrony following extinction occurs when populations undergoing extinction have different carrying capacities than surrounding populations. The result is not owing to biased migration per se, but rather is because of the number of immigrants relative to the carrying capacity. Synchrony following extinction should be most common for patchy populations, but can occur in any situation where carrying capacities differ. Overall, our results indicate that local extinction can create a positive feedback for extinction risk, increasing the probability of extinction for population networks by synchronizing their dynamics.

Development of a combined sex pheromone-based monitoring system for Malacosoma disstria (Lepidoptera: Lasoicampidae) and Choristoneura conflictana (Lepidoptera: Tortricidae).

Sympatric insect species that do not share sex pheromone components but have a common host and overlapping adult flight periods are potential targets for the development of a combined sex pheromone-based monitoring tool. A system using a single synthetic pheromone blend in a single lure to bait a single trap to monitor two pests simultaneously represents a novel approach. In this study, a combined pheromone-based monitoring system was developed for two lepidopterous defoliators of trembling aspen Populus tremuloides Michenaux in western Canada, Malacosoma disstria Hübner (Lepidoptera: Lasoicampidae) and Choristoneura conflictana (Walker) (Lepidoptera: Tortricidae). Efficacy and longevity of a lure containing both species' pheromones were tested. Immature stages of each species were sampled to evaluate the ability of pheromone traps baited with the combined lure to predict population density. The combined lure was as attractive to M. disstria and C. conflictana males as were traps baited with each species' pheromone alone. Lure age had no effect on attraction of male C. conflictana to the combined lure but had a negative effect on attraction of M. disstria. The number of male moths captured in traps baited with the combined lure was related to immature counts for both species. Pupal counts of M. disstria and larval counts of C. conflictana provided the best relationships with male captures. The combined lure does not attract M. disstria males in direct proportion to population density, because trap catch was comparatively low at high-density M. disstria sites.

The impact of parasitoid emergence time on host-parasitoid population dynamics.

We investigate the effect of parasitoid phenology on host-parasitoid population cycles. Recent experimental research has shown that parasitized hosts can continue to interact with their unparasitized counterparts through competition. Parasitoid phenology, in particular the timing of emergence from the host, determines the duration of this competition. We construct a discrete-time host-parasitoid model in which within-generation dynamics associated with parasitoid timing is explicitly incorporated. We found that late-emerging parasitoids induce less severe, but more frequent, host outbreaks, independent of the choice of competition model. The competition experienced by the parasitized host reduces the parasitoids' numerical response to changes in host numbers, preventing the 'boom-bust' dynamics associated with more efficient parasitoids. We tested our findings against experimental data for the forest tent caterpillar (Malacosoma disstria Hübner) system, where a large number of consecutive years at a high host density is synonymous with severe forest damage.

Reproductive asynchrony in natural butterfly populations and its consequences for female matelessness.

1. Reproductive asynchrony, where individuals in a population are short-lived relative to the population-level reproductive period, has been identified recently as a theoretical mechanism of the Allee effect that could operate in diverse plant and insect species. The degree to which this effect impinges on the growth potential of natural populations is not yet well understood. 2. Building on previous models of reproductive timing, we develop a general framework that allows a detailed, quantitative examination of the reproductive potential lost to asynchrony in small natural populations. 3. Our framework includes a range of biologically plausible submodels that allow details of mating biology of different species to be incorporated into the basic reproductive timing model. 4. We tailor the parameter estimation methods of the full model (basic model plus mating biology submodels) to take full advantage of data from detailed field studies of two species of Parnassius butterflies whose mating status may be assessed easily in the field. 5. We demonstrate that for both species, a substantial portion of the female population (6.5-18.6%) is expected to die unmated. These analyses provide the first direct, quantitative evidence of female reproductive failure due to asynchrony in small natural populations, and suggest that reproductive asynchrony exerts a strong and largely unappreciated influence on the population dynamics of these butterflies and other species with similarly asynchronous reproductive phenology.

Encroaching forests decouple alpine butterfly population dynamics.

Over the past 50 years, the rising tree line along Jumpingpound Ridge in the Rocky Mountains of Alberta, Canada, has reduced the area of alpine meadows and isolated populations that reside within them. By analyzing an 11-year data set of butterfly population sizes for 17 subpopulations along the ridge, we show that forest habitat separating alpine meadows decouples the dynamics of populations of the alpine butterfly Parnassius smintheus. Although the distance between populations is often negatively correlated with synchrony of dynamics, here we show that distance through forest, not Euclidean distance, determines the degree of synchrony. This effect is consistent with previous results demonstrating that encroaching forest reduces dispersal among populations and reduces gene flow. Decoupling dynamics produces more smaller independent populations, each with greater risk of local extinction, but decoupling may produce a lower risk of regional extinction in this capricious environment.

Genetic differentiation and gene flow among populations of the alpine butterfly, Parnassius smintheus, vary with landscape connectivity.

Levels of gene flow among populations vary both inter- and intraspecifically, and understanding the ecological bases of variation in levels of gene flow represents an important link between the ecological and evolutionary dynamics of populations. The effects of habitat spatial structure on gene flow have received considerable attention; however, most studies have been conducted at a single spatial scale and without background data on how individual movement is affected by landscape features. We examined the influence of habitat connectivity on inferred levels of gene flow in a high-altitude, meadow-dwelling butterfly, Parnassius smintheus. For this species, we had background data on the effects of landscape structure on both individual movement and on small-scale population genetic differentiation. We compared genetic differentiation and patterns of isolation by distance, based on variation at seven microsatellite loci, among three regions representing two levels of connectivity of high-altitude, nonforested habitats. We found that reduced connectivity of habitats, resulting from more forest cover at high altitudes, was associated with greater genetic differentiation among populations (higher estimated FST), a breakdown of isolation by distance, and overall lower levels of inferred gene flow. These observed differences were consistent with expectations based on our knowledge of the movement behaviour of this species and on previous population genetic analyses conducted at the smaller spatial scale. Our results indicate that the role of gene flow may vary among groups of populations depending on the interplay between individual movement and the structure of the surrounding landscape.

Among- and within-patch components of genetic diversity respond at different rates to habitat fragmentation: an empirical demonstration.

Habitat fragmentation is a ubiquitous by-product of human activities that can alter the genetic structure of natural populations, with potentially deleterious effects on population persistence and evolutionary potential. When habitat fragmentation results in the subdivision of a population, random genetic drift then leads to the erosion of genetic diversity from within the resulting subpopulation, random genetic drift then leads to the erosion of genetic diversity from within the resulting subpopulations and greater genetic divergence among them. Theoretical and simulation analyses predict that these two main genetic effects of fragmentation, greater differentiation among resulting subpopulation and reduced genetic diversity within them, will proceed at very different rates. Despite important implications for the interpretation of genetics data from fragmented populations, empirical evidence for this phenomenon has been lacking. In this analysis, we carry out an empirical study in population of an alpine meadow-dwelling butterfly, which have become fragmented increasing forest cover over five decades. We show that genetic differentiation among subpopulations (G(ST)) is most highly correlated with contemporary forest cover, while genetics diversity within subpopulation (expected heterozygosity) is better correlated with the spatial pattern of forest cover 40 years in the past. Thus, where habitat fragmentation has occurred in recent decades, genetic differentiation among subpopulation can be near equilibrium while contemporary measures of within subpopulation diversity may substantially overestimate the equilibrium values that will eventually be attained.

The influence of matrix habitat on Aphthona flea beetle immigration to leafy spurge patches.

Variation in movement ability by insects among different non-habitat (matrix) types may have important implications for both metapopulation dynamics and weed biocontrol practices. We used a mark-recapture experiment to explore the effects of two different matrix habitats (grass vs shrub) on the ability of two species of Aphthona (Chrysomelidae: Coleoptera) flea beetle to immigrate to patches of the invasive weed, leafy spurge. Using generalized linear models, we compared effects of the matrix habitat types, species and sex on observed immigration probabilities. Our analyses demonstrated that one species (A. nigriscutis) had a much higher immigration probability when moving through a grass-dominated matrix than a shrub-dominated matrix whereas immigration probabilities for the second species (A. lacertosa) were similar in both matrix habitats but significantly lower overall than for A. nigriscutis. Furthermore, A. nigriscutis females were more likely to immigrate to spurge patches embedded in a grass matrix than in shrub, whereas the opposite occurred for males. Our results suggest that metapopulation dynamics may be strongly affected by the type(s) of matrix habitat present on a landscape. These effects also suggest that release strategies for weed biocontrol should be tailored according to the structure of the landscape into which releases are planned. In addition, even closely related species can have significantly different movement abilities which will also affect release strategies.

Large-scale forest fragmentation increases the duration of tent caterpillar outbreak.

I examined historical data (1950-1984) on the duration of outbreaks of the forest tent caterpillar (Malacosoma disstria) in northern Ontario, Canada. Outbreak duration was compared to host tree species dominance and forest structure over large areas of boreal forest partially cleared for agriculture. Abundance of the principal host tree species Populus tremuloides had no consistent effect on duration of outbreak within forest districts, and was negatively correlated with duration of outbreaks among the eight forest districts examined. The amount of forest edge per km2 was the best, and most consistent, predictor of the duration of tent caterpillar outbreaks both within individual forest districts and among forest districts. Because forest tent caterpillar populations are driven largely by the impact of parasitoids and pathogens, results here suggest that large-scale increase in forest fragmentation affects the interaction between these natural enemies and forest tent caterpillar. Increased clearing and fragmentation of boreal forests, by agriculture and forestry, may be exacerbating outbreaks of this forest defoliator.

Foraging pattern of pine siskins and its influence on winter moth survival in an apple orchard.

Foraging by migratory pine siskins in an apple orchard infested with varying densities of winter moth was observed, and winter moth mortality in the presence and absence of birds was recorded. Time spent foraging in a tree and number of birds foraging per tree was positively related to larval density but number of larvae removed per leaf cluster or per unit time was not. Level of defoliation was a better predictor of the number of clusters searched per tree than was prey density. Despite poor predictability in allocation of search effort with respect to prey density, siskins acted as a source of strong compensatory mortality on the winter moth population.

Melanism and diel activity of alpine Colias (Lepidoptera: Pieridae).

The hypothesis that increased melanism provides a benefit in prolonging diel activity through more efficient absorption of solar radiation was tested in the field on a population of alpine Colias sulphur butterflies. A marked increase in the duration of flight and feeding behaviour existed for melanistic individuals when compared to lighter individuals under cool temperatures and low intensity solar radiation. More melanistic butterflies moved longer distance per day, and emigrated from the population at a faster rate. At high temperature and high radiant load lighter coloured individuals appeared more active. This is the first field demonstration of the advantage of melanism for increasing activity of ectotherms in cold environments.