Novel host plant use by a specialist insect depends on geographic variation in both the host and herbivore species.

Understanding the circumstances under which insect herbivores will adopt a novel host plant is a longstanding question in basic and applied ecology. While geographic variation in host use can arise through differences in both herbivore preference and plant characteristics, there is a tendency to attribute geographic variation in host use to regional differences in herbivore preference alone. This is especially true for herbivores specialized to one or a few plant species. We compared how geographic variation in herbivore preference and host plant origin shape regional differences in host plant use by the specialized herbivore, Euphydryas phaeton. In parts of its range, E. phaeton uses only a native host, Chelone glabra, while in others, it also uses an introduced host, Plantago lanceolata. We offered female butterflies from each region the non-native host plant sourced from both regions and compared their oviposition behavior. The non-native host was almost universally rejected by butterflies in the region where only the native plant is used. In the region where butterflies use both hosts, females accepted non-native plants from their natal region twice as often as non-native plants from the other region where they are not used. Acceptance differed substantially among individual butterflies within regions but not among plants within regions. Thus, both individual preference and regional differences in both the insect and non-native host contributed to the geographic variation in different ways. These results highlight that, in addition to herbivore preference, regional differences in perceived plant suitability may be an important driver of diet breadth.

Why are monarch butterflies declining in the West? Understanding the importance of multiple correlated drivers.

Understanding the factors associated with declines of at-risk species is an important first step in setting management and recovery targets. This step can be challenging when multiple aspects of climate and land use are changing simultaneously, and any or all could be contributing to population declines. We analyzed population trends of monarch butterflies in western North America in relation to likely environmental drivers. Unlike the larger eastern monarch population, past analyses of western monarchs have only evaluated the importance of climate (i.e., not land use) factors as drivers of abundance. We used partial least squares regression (PLSR) to evaluate the potential importance of changes in land use and climate variables. Trends in western monarch abundance were more strongly associated with land use variables than climate variables. Conclusions about importance of climate and land use variables were robust to changes in PLSR model structure. However, individual variables were too collinear to unambiguously separate their effects. We compared these conclusions to the more widely used technique of multiple regression, followed by multi-model inference (MRMI). Naïve interpretation of MRMI results could be misleading, if collinearity were not taken into account. MRMI was also highly sensitive to variation in model construction. Our results suggest a two-pronged approach to monarch conservation, specifically, starting efforts now to restore habitat, while also using experiments to more clearly delineate separate effects of climate and land use factors. They also demonstrate the utility of PLSR, a technique that is growing in use but is still relatively under-appreciated in conservation biology.

On the relationship between body condition and parasite infection in wildlife: a review and meta-analysis.

Body condition metrics are widely used to infer animal health and to assess costs of parasite infection. Since parasites harm their hosts, ecologists might expect negative relationships between infection and condition in wildlife, but this assumption is challenged by studies showing positive or null condition-infection relationships. Here, we outline common condition metrics used by ecologists in studies of parasitism, and consider mechanisms that cause negative, positive, and null condition-infection relationships in wildlife systems. We then perform a meta-analysis of 553 condition-infection relationships from 187 peer-reviewed studies of animal hosts, analysing observational and experimental records separately, and noting whether authors measured binary infection status or intensity. Our analysis finds substantial heterogeneity in the strength and direction of condition-infection relationships, a small, negative average effect size that is stronger in experimental studies, and evidence for publication bias towards negative relationships. The strongest predictors of variation in study outcomes are host thermoregulation and the methods used to evaluate body condition. We recommend that studies aiming to assess parasite impacts on body condition should consider host-parasite biology, choose condition measures that can change during the course of infection, and employ longitudinal surveys or manipulate infection status when feasible.

Anthropogenic fragmentation of landscapes: mechanisms for eroding the specificity of plant-herbivore interactions.

Reduced ecological specialization is an emerging, general pattern of ecological networks in fragmented landscapes. In plant-herbivore interactions, reductions in dietary specialization of herbivore communities are consistently associated with fragmented landscapes, but the causes remain poorly understood. We propose several hypothetical bottom-up and top-down mechanisms that may reduce the specificity of plant-herbivore interactions. These include empirically plausible applications and extensions of theory based on reduced habitat patch size and isolation (considered jointly), and habitat edge effects. Bottom-up effects in small, isolated habitat patches may limit availability of suitable hostplants, a constraint that increases with dietary specialization. Poor hostplant quality due to inbreeding in such fragments may especially disadvantage dietary specialist herbivores even when their hostplants are present. Size and isolation of habitat patches may change patterns of predation of herbivores, but whether such putative changes are associated with herbivore dietary specialization should depend on the mobility, size, and diet breadth of predators. Bottom-up edge effects may favor dietary generalist herbivores, yet top-down edge effects may favor dietary specialists owing to reduced predation. An increasingly supported edge effect is trophic ricochets generated by large grazers/browsers, which remove key hostplant species of specialist herbivores. We present empirical evidence that greater deer browsing in small forest fragments disproportionately reduces specialist abundances in lepidopteran assemblages in northeastern USA. Despite indirect evidence for these mechanisms, they have received scant direct testing with experimental approaches at a landscape scale. Identifying their relative contributions to reduced specificity of plant-herbivore interactions in fragmented landscapes is an important research goal.

Losing a battle but winning the war: moving past preference-performance to understand native herbivore-novel host plant interactions.

Introduced plants can positively affect population viability by augmenting the diet of native herbivores, but can negatively affect populations if they are subpar or toxic resources. In organisms with complex life histories, such as insects specializing on host plants, the impacts of a novel host may differ across life stages, with divergent effects on population persistence. Most research on effects of novel hosts has focused on adult oviposition preference and larval performance, but adult preference may not optimize offspring performance, nor be indicative of host quality from a demographic perspective. We compared population growth rates of the Baltimore checkerspot butterfly, Euphydryas phaeton, on an introduced host, Plantago lanceolata (English plantain), and the native host Chelone glabra (white turtlehead). Contrary to the previous findings suggesting that P. lanceolata could be a population sink, we found higher population growth rates (λ) on the introduced than the native host, even though some component parameters of λ were higher on the native host. Our findings illustrate the importance of moving beyond preference-performance studies to integrate vital rates across all life stages for evaluating herbivore-host plant relationships. Single measures of preference or performance are not sufficient proxies for overall host quality nor do they provide insights into longer term consequences of novel host plant use. In our system, in particular, P. lanceolata may buffer checkerspot populations when the native host is limiting, but high growth rates could lead to crashes over longer time scales.

Individual variation changes dispersal distance and area requirements of a checkerspot butterfly.

Individual variation in movement can have important consequences for spatial population dynamics. For instance, individual variation increases leptokurtosis in dispersal distance, such that more individuals move very short and very long distances relative to a homogeneous population. We quantified individual variation in movement of the Baltimore checkerspot butterfly (Euphydryas phaeton) to investigate its importance for two conservation-related metrics: the expected dispersal distance and the critical minimum patch size, or the smallest area within which a population can persist based on loss due to emigration. All movement parameters showed among-individual variation, with the greatest variation in move lengths and time spent resting. Correlations in among-individual movement parameters indicated that some butterflies were generally more mobile than others. We incorporated empirically estimated movement and demographic parameters into two individual-based models (IBMs), one with homogeneity in movement among individuals, and one with heterogeneity in movement. As expected, individual variation in movement increased the leptokurtosis of lifetime movement distance; the maximum difference in distance moved was substantial (-850 m vs. -5800 m) and is likely of significance for conservation. Individual variation also affected the critical minimum patch size, but the difference (-0.7 ha vs. -0.5 ha) is unlikely to be ecologically significant. Notably, populations with individual variation had higher growth rates in small patches and lower growth rates in large patches, a logical consequence of increased leptokurtosis. Individual variation in movement is fairly straightforward to quantify using mixed effects models and is important for spatial population dynamics, thus we encourage its inclusion in studies of other systems.

Minimum area requirements for an at-risk butterfly based on movement and demography.

Determining the minimum area required to sustain populations has a long history in theoretical and conservation biology. Correlative approaches are often used to estimate minimum area requirements (MARs) based on relationships between area and the population size required for persistence or between species' traits and distribution patterns across landscapes. Mechanistic approaches to estimating MAR facilitate prediction across space and time but are few. We used a mechanistic MAR model to determine the critical minimum patch size (CMP) for the Baltimore checkerspot butterfly (Euphydryas phaeton), a locally abundant species in decline along its southern range, and sister to several federally listed species. Our CMP is based on principles of diffusion, where individuals in smaller patches encounter edges and leave with higher probability than those in larger patches, potentially before reproducing. We estimated a CMP for the Baltimore checkerspot of 0.7-1.5 ha, in accordance with trait-based MAR estimates. The diffusion rate on which we based this CMP was broadly similar when estimated at the landscape scale (comparing flight path vs. capture-mark-recapture data), and the estimated population growth rate was consistent with observed site trends. Our mechanistic approach to estimating MAR is appropriate for species whose movement follows a correlated random walk and may be useful where landscape-scale distributions are difficult to assess, but demographic and movement data are obtainable from a single site or the literature. Just as simple estimates of lambda are often used to assess population viability, the principles of diffusion and CMP could provide a starting place for estimating MAR for conservation.

Taxonomic, phylogenetic, and trait Beta diversity in South American hummingbirds.

Comparison of the taxonomic, phylogenetic, and trait dimensions of beta diversity may uncover the mechanisms that generate and maintain biodiversity, such as geographic isolation, environmental filtering, and convergent adaptation. We developed an approach to predict the relationship between environmental and geographic distance and the dimensions of beta diversity. We tested these predictions using hummingbird assemblages in the northern Andes. We expected taxonomic beta diversity to result from recent geographic barriers limiting dispersal, and we found that cost distance, which includes barriers, was a better predictor than Euclidean distance. We expected phylogenetic beta diversity to result from historical connectivity and found that differences in elevation were the best predictors of phylogenetic beta diversity. We expected high trait beta diversity to result from local adaptation to differing environments and found that differences in elevation were correlated with trait beta diversity. When combining beta diversity dimensions, we observe that high beta diversity in all dimensions results from adaption to different environments between isolated assemblages. Comparisons with high taxonomic, low phylogenetic, and low trait beta diversity occurred among lowland assemblages separated by the Andes, suggesting that geographic barriers have recently isolated lineages in similar environments. We provide insight into mechanisms governing hummingbird biodiversity patterns and provide a framework that is broadly applicable to other taxonomic groups.

Faster movement in nonhabitat matrix promotes range shifts in heterogeneous landscapes.

Ecologists often assume that range expansion will be fastest in landscapes composed entirely of the highest-quality habitat. Theoretical models, however, show that range expansion depends on both habitat quality and habitat-specific movement rates. Using data from 78 species in 70 studies, we find that animals typically have faster movement through lower-quality environments (73% of published cases). Therefore, if we want to manage landscapes for range expansion, there is a trade-off between promoting movement with nonhostile matrix, and promoting population growth with high-quality habitat. We illustrate how this trade-off plays out with the use of an exemplar species, the Baltimore checkerspot butterfly. For this species, we calculate that the expected rate of range expansion is fastest in landscapes with ~15% high-quality habitat. Behavioral responses to nonhabitat matrix have often been documented in animal populations, but rarely included in empirical predictions of range expansion. Considering movement behavior could change land-planning priorities from focus on high-quality habitat only to integrating high- and low-quality land cover types, and evaluating the costs and benefits of different matrix land covers for range expansion.

Consequences of resource supplementation for disease risk in a partially migratory population.

Anthropogenic landscape features such as urban parks and gardens, landfills and farmlands can provide novel, seasonally reliable food sources that impact wildlife ecology and distributions. In historically migratory species, food subsidies can cause individuals to forgo migration and form partially migratory or entirely sedentary populations, eroding a crucial benefit of migration: pathogen avoidance through seasonal abandonment of transmission sites and mortality of infected individuals during migration. Since many migratory taxa are declining, and wildlife populations in urban areas can harbour zoonotic pathogens, understanding the mechanisms by which anthropogenic resource subsidies influence infection dynamics and the persistence of migration is important for wildlife conservation and public health. We developed a mathematical model for a partially migratory population and a vector-borne pathogen transmitted at a shared breeding ground, where food subsidies increase the nonbreeding survival of residents. We found that higher resident nonbreeding survival increased infection prevalence in residents and migrants, and lowered the fraction of the population that migrated. The persistence of migration may be especially threatened if residency permits emergence of more virulent pathogens, if resource subsidies reduce costs of infection for residents, and if infection reduces individual migratory propensity.This article is part of the theme issue 'Anthropogenic resource subsidies and host-parasite dynamics in wildlife'.

Modeling vector-borne disease risk in migratory animals under climate change.

Recent theory suggests that animals that migrate to breed at higher latitudes may benefit from reduced pressure from natural enemies, including pathogens ("migratory escape"), and that migration itself weeds out infected individuals and lowers infection prevalence ("migratory culling"). The distribution and activity period of arthropod disease vectors in temperate regions is expected to respond rapidly to climate change, which could reduce the potential for migratory escape. However, climate change could have the opposite effect of reducing transmission if differential responses in the phenology and distribution of migrants and disease vectors reduce their overlap in space and time. Here we outline a simple modeling framework for exploring the influence of climate change on vector-borne disease dynamics in a migratory host. We investigate two scenarios under which pathogen transmission dynamics might be mediated by climate change: (1) vectors respond more rapidly than migrants to advancing phenology at temperate breeding sites, causing peak susceptible host density and vector emergence to diverge ("migratory mismatch") and (2) reduced migratory propensity allows increased nonbreeding survival of infected hosts and larger breeding-site epidemics (loss of migratory culling, here referred to as "sedentary amplification"). Our results highlight the need for continued surveillance of climate-induced changes to migratory behavior and vector activity to predict pathogen prevalence and its impacts on migratory animals.

Seasonal mortality patterns in non-human primates: implications for variation in selection pressures across environments.

Examining seasonal mortality patterns can yield insights into the drivers of mortality and thus potential selection pressures acting on individuals in different environments. We compiled adult and juvenile mortality data from nine wild non-human primate taxa to investigate the role of seasonality in patterns of mortality and address the following questions: Is mortality highly seasonal across species? Does greater environmental seasonality lead to more seasonal mortality patterns? If mortality is seasonal, is it higher during wet seasons or during periods of food scarcity? and Do folivores show less seasonal mortality than frugivores? We found seasonal mortality patterns in five of nine taxa, and mortality was more often tied to wet seasons than food-scarce periods, a relationship that may be driven by disease. Controlling for phylogeny, we found a positive relationship between the degree of environmental seasonality and mortality, with folivores exhibiting more seasonal mortality than frugivores. These results suggest that mortality patterns are influenced both by diet and degree of environmental seasonality. Applied to a wider array of taxa, analyses of seasonal mortality patterns may aid understanding of life-history evolution and selection pressures acting across a broad spectrum of environments and spatial and temporal scales.