Evolutionary dynamics of dispersal and local adaptation in multi-resource landscapes.

Dispersal can enable access to resources in new locations. Consequently, traits that govern dispersal probability and dispersal distance may impact an individual's ability to acquire resources. However, spatial variation in the quality or quantity of resources may mediate potential adaptive benefits of novel dispersal traits. Ecological traits (i.e., those that determine how an individual processes resources) will also, by definition, affect how an individual interacts with the resource landscape. In a spatially heterogeneous environment, this creates potential for evolutionary feedbacks between dispersal-related traits and ecological traits. For example, dispersal may introduce individuals to novel resources, at which point there may be selection for local adaptation of ecological traits. Conversely, an individual's ability to utilize different resource types may determine how dispersal impacts fitness. Here, we develop an individual-based model to investigate co-evolution of dispersal and ecological traits in a landscape where multiple resources vary independently across space. We find that: (1) resource specialists can emerge and tend to evolve dispersal strategies suited to the structure of their preferred resource type and (2) generalists, when they emerge, tend to possess intermediate dispersal strategies. Lastly, we note that the effect of dispersal on the evolution of the ecological trait is weaker than vice versa and, as a result, appreciable heterogeneity in the abundance of resources across a landscape will likely obscure a signal of co-evolution.

Climate change winners and losers among North American bumblebees.

Mounting evidence suggests that climate change, agricultural intensification and disease are impacting bumblebee health and contributing to species' declines. Identifying how these factors impact insect communities at large spatial and temporal scales is difficult, partly because species may respond in different ways. Further, the necessary data must span large spatial and temporal scales, which usually means they comprise aggregated, presence-only records collected using numerous methods (e.g. diversity surveys, educational collections, citizen-science projects, standardized ecological surveys). Here, we use occupancy models, which explicitly correct for biases in the species observation process, to quantify the effect of changes in temperature, precipitation and floral resources on bumblebee site occupancy over the past 12 decades in North America. We find no evidence of genus-wide declines in site occupancy, but do find that occupancy is strongly related to temperature, and is only weakly related to precipitation or floral resources. We also find that more species are likely to be climate change 'losers' than 'winners' and that this effect is primarily associated with changing temperature. Importantly, all trends were highly species-specific, highlighting that genus or community-wide measures may not reflect diverse species-specific patterns that are critical in guiding allocation of conservation resources.

Interactive effects of multiscale diversification practices on farmland bird stress.

Farmland diversification practices (i.e., methods used to produce food sustainably by enhancing biodiversity in cropping systems) are sometimes considered beneficial to both agriculture and biodiversity, but most studies of these practices rely on species richness, diversity, or abundance as a proxy for habitat quality. Biodiversity assessments may miss early clues that populations are imperiled when species presence does not imply persistence. Physiological stress indicators may help identify low-quality habitats before population declines occur. We explored how avian stress indicators respond to on-farm management practices and surrounding seminatural area (1-km radius) across 21 California strawberry farms. We examined whether commonly used biodiversity metrics correlate with stress responses in wild birds. We used ∼1000 blood and feather samples and body mass and wing chord measurements, mostly from passerines, to test the effects of diversification practices on four physiological stress indicators: heterophil to lymphocyte ratios (H:L), body condition, hematocrit values, and feather growth rates of individual birds. We then tested the relationship between physiological stress indicators and species richness, abundance, occurrence, and diversity derived from 285 bird point count surveys. After accounting for other biological drivers, landscape context mediated the effect of local farm management on H:L and body condition. Local diversification practices were associated with reduced individual stress in intensive agricultural landscapes but increased it in landscapes surrounded by relatively more seminatural area. Feathers grew more slowly in landscapes dominated by strawberry production, suggesting that nutritional condition was lower here than in landscapes with more crop types and seminatural areas. We found scant evidence that species richness, abundance, occurrence, or diversity metrics were correlated with the individual's physiological stress, suggesting that reliance on these metrics may obscure the impacts of management on species persistence. Our findings underscore the importance of considering landscape context when designing local management strategies to promote wildlife conservation.

Algunas veces se considera a las prácticas de diversificación agrícola (es decir, los métodos usados para producir alimentos de manera sustentable mediante el enriquecimiento de la biodiversidad en los sistemas de cultivo) como benéficas para la agricultura y la biodiversidad, pero la mayoría de los estudios sobre estas prácticas dependen de la riqueza, abundancia o diversidad de especies como indicadores de la calidad del hábitat. Las valoraciones de la biodiversidad pueden ignorar las señales tempranas de una población en peligro cuando la presencia de la especie no implica persistencia. Los indicadores de estrés fisiológico pueden auxiliar en la identificación de hábitats de baja calidad antes de que ocurra la declinación poblacional. Analizamos las respuestas de los indicadores de estrés en aves al manejo en las granjas y áreas seminaturales circundantes (1 km de radio) de 21 cultivos de fresas en California. Evaluamos si las medidas comunes de biodiversidad se correlacionan con las respuestas al estrés de las aves silvestres. Usamos aproximadamente mil muestras de sangre y plumas y medidas de masa corporal y cuerda alar, la mayoría de paseriformes, para analizar los efectos de las prácticas de diversificación sobre cuatro indicadores de estrés fisiológico: la relación heterófilos/linfocitos (H:L), condición corporal; valores hematocritos; y la tasa de crecimiento de las plumas en aves individuales. Después probamos la relación entre los indicadores de estrés fisiológicos y la riqueza, abundancia, presencia y diversidad de especies tomadas de 285 conteos por puntos de aves. Después de considerar otros factores biológicos, el contexto del paisaje medió el efecto de la gestión de las granjas locales sobre la H:L y la condición corporal. Las prácticas locales de diversificación estuvieron asociadas con una reducción en el estrés individual en los paisajes con agricultura intensiva; sin embargo, el estrés aumentó en los paisajes rodeados por áreas relativamente más seminaturales. Encontramos poca evidencia que respalde que las medidas de riqueza, abundancia, presencia y diversidad de especies estuvieran correlacionadas con el estrés fisiológico de los individuos, lo que sugiere que depender de estas medidas puede nublar el impacto de la gestión sobre la persistencia de las especies. Nuestros descubrimientos apuntalan lo importante que es considerar el contexto del paisaje cuando se diseñan las estrategias de gestión local para promover la conservación de la fauna.

Can You Trust Who You See? The Evolution of Socially Cued Anticipatory Plasticity.

AbstractThe social environment can affect development and fitness. However, we do not know how selection acts on individuals that cue developmental pathways using features of the social environment. Socially cued anticipatory plasticity (SCAP) is a hypothetical strategy whereby juveniles use social cues to alter development to match their adult phenotype to the social environment that they expect to encounter. While intuitively appealing, the evolution of such plasticity is a puzzle, because the cue changes when individuals use it. Can socially cued plasticity evolve when such a feedback occurs? We use individual-based simulations to model evolution of SCAP in an environment that fluctuates between favoring each of two discrete phenotypes. We found that socially cued plasticity evolved, but only when strong selection acted on survival rather than on fecundity differences between adult phenotypes. In this case, the social cue reliably predicted which phenotype would be favored on maturation. Surprisingly, costs to plasticity increased the range of conditions under which it was adaptive. In the absence of costs, evolution led to a state where SCAP individuals could not effectively respond to environmental changes. Costs to plasticity lowered the proportion of the population that used SCAP, which in turn increased the reliability of the social cue and allowed individuals that used socially cued plasticity to switch between the favored phenotypes more consistently. Our results suggest that the evolution of adaptive plasticity in response to social cues may represent a larger class of problems in which evolution is hard to predict because of feedbacks among critical processes.

Proximity of restored hedgerows interacts with local floral diversity and species' traits to shape long-term pollinator metacommunity dynamics.

Disconnected habitat fragments are poor at supporting population and community persistence; restoration ecologists, therefore, advocate for the establishment of habitat networks across landscapes. Few empirical studies, however, have considered how networks of restored habitat patches affect metacommunity dynamics. Here, using a 10-year study on restored hedgerows and unrestored field margins within an intensive agricultural landscape, we integrate occupancy modelling with network theory to examine the interaction between local and landscape characteristics, habitat selection and dispersal in shaping pollinator metacommunity dynamics. We show that surrounding hedgerows and remnant habitat patches interact with the local floral diversity, bee diet breadth and bee body size to influence site occupancy, via colonisation and persistence dynamics. Florally diverse sites and generalist, small-bodied species are most important for maintaining metacommunity connectivity. By providing the first in-depth assessment of how a network of restored habitat influences long-term population dynamics, we confirm the conservation benefit of hedgerows for pollinator populations and demonstrate the importance of restoring and maintaining habitat networks within an inhospitable matrix.

Temporally varying disruptive selection in the medium ground finch (Geospiza fortis).

Disruptive natural selection within populations exploiting different resources is considered to be a major driver of adaptive radiation and the production of biodiversity. Fitness functions, which describe the relationships between trait variation and fitness, can help to illuminate how this disruptive selection leads to population differentiation. However, a single fitness function represents only a particular selection regime over a single specified time period (often a single season or a year), and therefore might not capture longer-term dynamics. Here, we build a series of annual fitness functions that quantify the relationships between phenotype and apparent survival. These functions are based on a 9-year mark-recapture dataset of over 600 medium ground finches (Geospiza fortis) within a population bimodal for beak size. We then relate changes in the shape of these functions to climate variables. We find that disruptive selection between small and large beak morphotypes, as reported previously for 2 years, is present throughout the study period, but that the intensity of this selection varies in association with the harshness of environment. In particular, we find that disruptive selection was strongest when precipitation was high during the dry season of the previous year. Our results shed light on climatic factors associated with disruptive selection in Darwin's finches, and highlight the role of temporally varying fitness functions in modulating the extent of population differentiation.

Sexual selection enables long-term coexistence despite ecological equivalence.

Empirical data indicate that sexual preferences are critical for maintaining species boundaries, yet theoretical work has suggested that, on their own, they can have only a minimal role in maintaining biodiversity. This is because long-term coexistence within overlapping ranges is thought to be unlikely in the absence of ecological differentiation. Here we challenge this widely held view by generalizing a standard model of sexual selection to include two ubiquitous features of populations with sexual selection: spatial variation in local carrying capacity, and mate-search costs in females. We show that, when these two features are combined, sexual preferences can single-handedly maintain coexistence, even when spatial variation in local carrying capacity is so slight that it might go unnoticed empirically. This theoretical study demonstrates that sexual selection alone can promote the long-term coexistence of ecologically equivalent species with overlapping ranges, and it thus provides a novel explanation for the maintenance of species diversity.

Phylogenetic occupancy models integrate imperfect detection and phylogenetic signal to analyze community structure.

Biological communities are structured phylogenetically-closely related species are typically more likely to be found at the same sites. This may be, in part, because they respond similarly to environmental gradients. Accurately surveying biological communities is, however, made difficult by the fact that detection of species is not perfect. In recent years, numerous statistical methods have been developed that aim to overcome deficiencies in the species detection process. However, these methods do not allow investigators to assess phylogenetic community structure. Here, we introduce the phylogenetic occupancy model (POM), which accounts for imperfect species detection while assessing phylogenetic patterns in community structure. Using simulated data sets we show that the POM grants less biased estimates of phylogenetic structure than models without imperfect detection, and can correctly ascertain the effects of species traits on community composition while accounting for evolutionary non-independence of taxa. Integrating phylogenetic methods into widely used occupancy models will help clarify how evolutionary history influences modern day communities.

Climate change and habitat conversion favour the same species.

Land-use change and climate change are driving a global biodiversity crisis. Yet, how species' responses to climate change are correlated with their responses to land-use change is poorly understood. Here, we assess the linkages between climate and land-use change on birds in Neotropical forest and agriculture. Across > 300 species, we show that affiliation with drier climates is associated with an ability to persist in and colonise agriculture. Further, species shift their habitat use along a precipitation gradient: species prefer forest in drier regions, but use agriculture more in wetter zones. Finally, forest-dependent species that avoid agriculture are most likely to experience decreases in habitable range size if current drying trends in the Neotropics continue as predicted. This linkage suggests a synergy between the primary drivers of biodiversity loss. Because they favour the same species, climate and land-use change will likely homogenise biodiversity more severely than otherwise anticipated.

On-farm habitat restoration counters biotic homogenization in intensively managed agriculture.

To slow the rate of global species loss, it is imperative to understand how to restore and maintain native biodiversity in agricultural landscapes. Currently, agriculture is associated with lower spatial heterogeneity and turnover in community composition (ß-diversity). While some techniques are known to enhance α-diversity, it is unclear whether habitat restoration can re-establish ß-diversity. Using a long-term pollinator dataset, comprising ∼9,800 specimens collected from the intensively managed agricultural landscape of the Central Valley of California, we show that on-farm habitat restoration in the form of native plant 'hedgerows', when replicated across a landscape, can boost ß-diversity by approximately 14% relative to unrestored field margins, to levels similar to some natural communities. Hedgerows restore ß-diversity by promoting the assembly of phenotypically diverse communities. Intensively managed agriculture imposes a strong ecological filter that negatively affects several important dimensions of community trait diversity, distribution, and uniqueness. However, by helping to restore phenotypically diverse pollinator communities, small-scale restorations such as hedgerows provide a valuable tool for conserving biodiversity and promoting ecosystem services.

Pyrodiversity begets plant-pollinator community diversity.

Fire has a major impact on the structure and function of many ecosystems globally. Pyrodiversity, the diversity of fires within a region (where diversity is based on fire characteristics such as extent, severity, and frequency), has been hypothesized to promote biodiversity, but changing climate and land management practices have eroded pyrodiversity. To assess whether changes in pyrodiversity will have impacts on ecological communities, we must first understand the mechanisms that might enable pyrodiversity to sustain biodiversity, and how such changes might interact with other disturbances such as drought. Focusing on plant-pollinator communities in mixed-conifer forest with frequent fire in Yosemite National Park, California, we examine how pyrodiversity, combined with drought intensity, influences those communities. We find that pyrodiversity is positively related to the richness of the pollinators, flowering plants, and plant-pollinator interactions. On average, a 5% increase in pyrodiversity led to the gain of approximately one pollinator and one flowering plant species and nearly two interactions. We also find that a diversity of fire characteristics contributes to the spatial heterogeneity (ß-diversity) of plant and pollinator communities. Lastly, we find evidence that fire diversity buffers pollinator communities against the effects of drought-induced floral resource scarcity. Fire diversity is thus important for the maintenance of flowering plant and pollinator diversity and predicted shifts in fire regimes to include less pyrodiversity compounded with increasing drought occurrence will negatively influence the richness of these communities in this and other forested ecosystems. In addition, lower heterogeneity of fire severity may act to reduce spatial turnover of plant-pollinator communities. The heterogeneity of community composition is a primary determinant of the total species diversity present in a landscape, and thus, lower pyrodiversity may negatively affect the richness of plant-pollinator communities across large spatial scales.

Diversification practices reduce organic to conventional yield gap.

Agriculture today places great strains on biodiversity, soils, water and the atmosphere, and these strains will be exacerbated if current trends in population growth, meat and energy consumption, and food waste continue. Thus, farming systems that are both highly productive and minimize environmental harms are critically needed. How organic agriculture may contribute to world food production has been subject to vigorous debate over the past decade. Here, we revisit this topic comparing organic and conventional yields with a new meta-dataset three times larger than previously used (115 studies containing more than 1000 observations) and a new hierarchical analytical framework that can better account for the heterogeneity and structure in the data. We find organic yields are only 19.2% (±3.7%) lower than conventional yields, a smaller yield gap than previous estimates. More importantly, we find entirely different effects of crop types and management practices on the yield gap compared with previous studies. For example, we found no significant differences in yields for leguminous versus non-leguminous crops, perennials versus annuals or developed versus developing countries. Instead, we found the novel result that two agricultural diversification practices, multi-cropping and crop rotations, substantially reduce the yield gap (to 9 ± 4% and 8 ± 5%, respectively) when the methods were applied in only organic systems. These promising results, based on robust analysis of a larger meta-dataset, suggest that appropriate investment in agroecological research to improve organic management systems could greatly reduce or eliminate the yield gap for some crops or regions.

Habitat restoration promotes pollinator persistence and colonization in intensively managed agriculture.

Widespread evidence of pollinator declines has led to policies supporting habitat restoration including in agricultural landscapes. Yet, little is yet known about the effectiveness of these restoration techniques for promoting stable populations and communities of pollinators, especially in intensively managed agricultural landscapes. Introducing floral resources, such as flowering hedgerows, to enhance intensively cultivated agricultural landscapes is known to increase the abundances of native insect pollinators in and around restored areas. Whether this is a result of local short-term concentration at flowers or indicative of true increases in the persistence and species richness of these communities remains unclear. It is also unknown whether this practice supports species of conservation concern (e.g., those with more specialized dietary requirements). Analyzing occupancies of native bees and syrphid flies from 330 surveys across 15 sites over eight years, we found that hedgerow restoration promotes rates of between-season persistence and colonization as compared with unrestored field edges. Enhanced persistence and colonization, in turn, led to the formation of more species-rich communities. We also find that hedgerows benefit floral resource specialists more than generalists, emphasizing the value of this restoration technique for conservation in agricultural landscapes.

Allee effects and species co-existence in an environment where resource abundance varies.

Explaining patterns of diversity has long been a central focus in ecology. One of the most challenging problems has been to understand how species occupying similar ecological niches can co-exist because, with limited resources, demographic stochasticity is expected to lead to the eventual extinction of all but one of them. The Allee effect has been widely studied for its impact on the extinction risk of rare species. Its potential role in promoting co-existence has received less attention. Here, we present a model in which two species compete for a single resource across a continuous landscape. We show that Allee effects can promote their co-existence when a simple condition is met: resources are distributed unevenly across space. Furthermore, the Allee effect can stabilize co-existence despite the reduction in population density and consequent increase in demographic stochasticity that it causes. The Allee effect might, therefore, be an important force maintaining diverse communities.

The evolution of mutation rate in an antagonistic coevolutionary model with maternal transmission of parasites.

By constantly selecting for novel genotypes, coevolution between hosts and parasites can favour elevated mutation rates. Models of this process typically assume random encounters. However, offspring are often more likely to encounter their mother's parasites. Because parents and offspring are genetically similar, they may be susceptible to the same parasite strains and thus, in hosts, maternal transmission should select for mechanisms that decrease intergenerational genetic similarity. In parasites, however, maternal transmission should select for genetic similarity. We develop and analyse a model of host and parasite mutation rate evolution when parasites are maternally inherited. In hosts, we find that maternal transmission has two opposing effects. First, it eliminates coevolutionary cycles that previous work shows select for higher mutation. Second, it independently selects for higher mutation rates, because offspring that differ from their mothers are more likely to avoid infection. In parasites, however, the two effects of maternal transmission act in the same direction. As for hosts, maternal transmission eliminates coevolutionary cycles, thereby reducing selection for increased mutation. Unlike for hosts, however, maternal transmission additionally selects against higher mutation by favouring parasite offspring that are the same as their mothers.

Positive impact of postfire environment on bumble bees not explained by habitat variables in a remote forested ecosystem.

Bumble bees are important pollinators in temperate forested regions where fire is a driving force for habitat change, and thus understanding how these insects respond to fire is critical. Previous work has shown bees are often positively affected by the postfire environment, with burned sites supporting greater bee abundance and diversity, and increased floral resources. The extent to which fire impacts variation in bumblebee site occupancy is not well-understood, especially in higher latitude regions with dense, primarily coniferous forests. Occupancy models are powerful tools for biodiversity analyses, as they separately estimate occupancy probability (likelihood that a species is present at a particular location) and detection probability (likelihood of observing a species when it is present). Using these models, we tested whether bumblebee site occupancy is higher in burned locations as a result of the increase in canopy openness, floral species richness, and floral abundance. We quantified the impact of fire, and associated habitat changes, on bumblebee species' occupancy in an area with high wildfire frequency in British Columbia, Canada. The burn status of a site was the only significant predictor for determining bumblebee occurrence (with burned sites having higher occupancy); floral resource availability and canopy openness only impacted detection probability (roughly, sample bias). These findings highlight the importance of controlling for the influence of habitat on species detection in pollinator studies and suggest that fire in this system changes the habitat for bumble bees in positive ways that extend beyond our measurements of differences in floral resources and canopy cover.

Co-evolution of dormancy and dispersal in spatially autocorrelated landscapes.

The evolution of dispersal can be driven by spatial processes, such as landscape structure, and temporal processes, such as disturbance. Dormancy, or dispersal in time, is generally thought to evolve in response to temporal processes. In spite of broad empirical and theoretical evidence of trade-offs between dispersal and dormancy, we lack evidence that spatial structure can drive the evolution of dormancy. Here, we develop a simulation-based model of the joint evolution of dispersal and dormancy in spatially heterogeneous landscapes. We show that dormancy and dispersal are each favored under different landscape conditions, but not simultaneously under any of the conditions we tested. We further show that, when dispersal distances are short, dormancy can evolve directly in response to landscape structure. In this case, selection is primarily driven by benefits associated with avoiding kin competition. Our results are similar in both highly simplified and realistically complex landscapes.

Epigenetic induction may speed up or slow down speciation with gene flow.

Speciation is less likely to occur when there is gene flow between nascent species. Natural selection can oppose gene flow and promote speciation if there is variation in ecological conditions among the nascent species' locations. Previous theory on ecological speciation with gene flow has focused primarily on the role of genetic variation in ecological traits, largely neglecting the role of nongenetic inheritance or transgenerational plasticity. Here, we build and analyze models incorporating both genetic and epigenetic inheritance, the latter representing a form of nongenetic inheritance. We investigate the rate of speciation for a population that inhabits two patches connected by migration, and find that adaptively biased epigenetic induction can speed up or slow down speciation, depending on the form of the map from genotype and epigenotype to phenotype. While adaptively relevant epigenetic variation can speed up speciation by reducing the fitness of migrants and hybrids, it can also slow down speciation. This latter effect occurs when the epialleles are able to achieve adaptation faster than the genetic alleles, thereby weakening selection on the latter.

Ploidy and the evolution of parasitism.

Levels of parasitism are continuously distributed in nature. Models of host-parasite coevolution, however, typically assume that species can be easily characterized as either parasitic or non-parasitic. Consequently, it is poorly understood which factors influence the evolution of parasitism itself. We investigate how ploidy level and the genetic mechanisms underlying infection influence evolution along the continuum of parasitism levels. In order for parasitism to evolve, selective benefits to the successful invasion of hosts must outweigh the losses when encountering resistant hosts. However, we find that exactly where this threshold occurs depends not only on the strength of selection, but also on the genetic model of interaction, the ploidy level in each species, and the nature of the costs to virulence and resistance. With computer simulations, we are able to incorporate more realistic dynamics at the loci underlying species interactions and to extend our analyses in a number of directions, including finite population sizes, multiple alleles and different generation times.

Integrating high-speed videos in capture-mark-recapture studies of insects.

Capture-mark-recapture (CMR) studies have been used extensively in ecology and evolution. While it is feasible to apply CMR in some animals, it is considerably more challenging in small fast-moving species such as insects. In these groups, low recapture rates can bias estimates of demographic parameters, thereby handicapping effective analysis and management of wild populations. Here, we use high-speed videos (HSV) to capture two large dragonfly species, Anax junius and Rhionaeschna multicolor, that rarely land and, thus, are particularly challenging for CMR studies. We test whether HSV, compared to conventional "eye" observations, increases the "resighting" rates and, consequently, improves estimates of both survival rates and the effects of demographic covariates on survival. We show that the use of HSV increases the number of resights by 64% in A. junius and 48% in R. multicolor. HSV improved our estimates of resighting and survival probability which were either under- or overestimated with the conventional observations. Including HSV improved credible intervals for resighting rate and survival probability by 190% and 130% in A. junius and R. multicolor, respectively. Hence, it has the potential to open the door to a wide range of research possibilities on species that are traditionally difficult to monitor with distance sampling, including within insects and birds.