Effects of environment and genotype on dispersal differ across departure, transfer and settlement in a butterfly metapopulation

Active dispersal is driven by extrinsic and intrinsic factors at the three stages of departure, transfer and settlement. Most empirical studies capture only one stage of this complex process, and knowledge of how much can be generalized from one stage to another remains unknown. Here we use genetic assignment tests to reconstruct dispersal across 5 years and 232 habitat patches of a Glanville fritillary butterfly (Melitaea cinxia) metapopulation. We link individual dispersal events to weather, landscape structure, size and quality of habitat patches, and individual genotype to identify the factors that influence the three stages of dispersal and post-settlement survival. We found that nearly all tested factors strongly affected departure probabilities, but that the same factors explained very little variation in realized dispersal distances. Surprisingly, we found no effect of dispersal distance on post-settlement survival. Rather, survival was influenced by weather conditions, quality of the natal habitat patch, and a strong interaction between genotype and occupancy status of the settled habitat patch, with more mobile genotypes having higher survival as colonists rather than as immigrants. Our work highlights the multi-causality of dispersal and that some dispersal costs can only be understood by considering extrinsic and intrinsic factors and their interaction across the entire dispersal process.Peer reviewe

Toward an integration of evolutionary biology and ecosystem

Abstract At present, the disciplines of evolutionary biology and ecosystem science are weakly integrated. As a result, we have a poor understanding of how the ecological and evolutionary processes that create, maintain, and change biological diversity affect the flux of energy and materials in global biogeochemical cycles. The goal of this article was to review several research fields at the interfaces between ecosystem science, community ecology and evolutionary biology, and suggest new ways to integrate evolutionary biology and ecosystem science. In particular, we focus on how phenotypic evolution by natural selection can influence ecosystem functions by affecting processes at the environmental, population and community scale of ecosystem organization. We develop an eco-evolutionary model to illustrate linkages between evolutionary change (e.g. phenotypic evolution of producer), ecological interactions (e.g. consumer grazing) and ecosystem processes (e.g. nutrient cycling). We conclude by proposing experiments to test the ecosystem consequences of evolutionary changes

Persistence of Pathogens with Short Infectious Periods in Seasonal Tick Populations: The Relative Importance of Three Transmission Routes

BACKGROUND: The flaviviruses causing tick-borne encephalitis (TBE) persist at low but consistent levels in tick populations, despite short infectious periods in their mammalian hosts and transmission periods constrained by distinctly seasonal tick life cycles. In addition to systemic and vertical transmission, cofeeding transmission has been proposed as an important route for the persistence of TBE-causing viruses. Because cofeeding transmission requires ticks to feed simultaneously, the timing of tick activity may be critical to pathogen persistence. Existing models of tick-borne diseases do not incorporate all transmission routes and tick seasonality. Our aim is to evaluate the influence of seasonality on the relative importance of different transmission routes by using a comprehensive mathematical model. METHODOLOGY/PRINCIPAL FINDINGS: We developed a stage-structured population model that includes tick seasonality and evaluated the relative importance of the transmission routes for pathogens with short infectious periods, in particular Powassan virus (POWV) and the related "deer tick virus," emergent encephalitis-causing flaviviruses in North America. We used the next generation matrix method to calculate the basic reproductive ratio and performed elasticity analyses. We confirmed that cofeeding transmission is critically important for such pathogens to persist in seasonal tick populations over the reasonable range of parameter values. At higher but still plausible rates of vertical transmission, our model suggests that vertical transmission can strongly enhance pathogen prevalence when it operates in combination with cofeeding transmission. CONCLUSIONS/SIGNIFICANCE: Our results demonstrate that the consistent prevalence of POWV observed in tick populations could be maintained by a combination of low vertical, intermediate cofeeding and high systemic transmission rates. When vertical transmission is weak, nymphal ticks support integral parts of the transmission cycle that are critical for maintaining the pathogen. We also extended the model to pathogens that cause chronic infections in hosts and found that cofeeding transmission could contribute to elevating prevalence even in these systems. Therefore, the common assumption that cofeeding transmission is not relevant in models of chronic host infection, such as Lyme disease, could lead to underestimating pathogen prevalence

The effects of functional response and host abundance fluctuations on genetic rescue in parasitoids with single-locus sex determination

Many parasitoids have single-locus complementary sex determination (sl-CSD), which produces sterile or inviable males when homozygous at the sex determining locus. A previous study theoretically showed that small populations have elevated risks of extinction due to the positive feedback between inbreeding and small population size, referred to as the diploid male vortex. A few modeling studies have suggested that the diploid male vortex may not be as common because balancing selection at sex determining loci tends to maintain high allelic diversity in spatially structured populations. However, the generality of the conclusion is yet uncertain, as they were drawn either from models developed for particular systems or from a general-purpose competition model. To attest the conclusion, we study several well-studied host-parasitoid models that incorporate functional response specifying the number of attacked hosts given a host density and derive the conditions for a diploid male vortex in a single population. Then, we develop spatially structured individual-based versions of the models to include female behavior, diploid male fertility, and temporal fluctuations. The results show that producing a handful of successful offspring per female parasitoid could enable parasitoid persistence when a typical number of CSD alleles are present. The effect of functional response depends on the levels of fluctuations in host abundance, and inviable or partially fertile diploid males and a small increase in dispersal can alleviate the risk of a diploid male vortex. Our work supports the generality of effective genetic rescue in spatially connected parasitoid populations with sl-CSD. However, under more variable climate, the efficacy of the CSD mechanism may substantially decline.Peer reviewe

Maintaining pathogens with short infectivity in seasonally structured tick populations: relative importance of three transmission pathways

Infectivity periods of different tick-borne pathogens in host species can vary widely, and the variation affects how the pathogens are maintained in tick populations. In addition to systemic and vertical transmission, cofeeding transmission has been proposed as an important route for the persistence of pathogens with short infectivity (e.g., tick-borne encephalitis causing viruses, TBEv). Because cofeeding transmission requires ticks to feed simultaneously, the temporal dynamics of tick populations become important. Existing models of tick-borne diseases do not fully incorporate all three transmission pathways (systemic, vertical, and cofeeding transmission) and tick seasonality. We developed a comprehensive stage-structured population model that includes seasonality and evaluated the relative importance of the three transmission pathways for pathogens with short infectivity. We used the next generation matrix method to calculate R0 and performed elasticity analyses for complex disease systems. We found that cofeeding transmission is a critically important route for such pathogens to persist in seasonal tick populations over the reasonable range of parameter values. At higher but still plausible rates of vertical transmission, our model suggests that vertical transmission can be a strong enhancer of pathogen prevalence when it operates in combination with cofeeding transmission. We discuss potential mechanisms behind consistent but low prevalence of TBEv observed in tick populations in the field.Master of ScienceMastersMathematics and StatisticsWearing, HelenSteinberg, StanlyPedro, Embi

Evolutionary consequences of ecological interactions

Eco-evolutionary dynamics integrates the reciprocal interactions betweenecology and evolution. These two branches of biology traditionally assumethe other as static for simplicity. However, increasing evidence shows thatthis simplification may not always hold because ecology and evolution canoperate in similar timescales. This thesis theoretically explores how thereciprocal interactions may influence ecological and evolutionary outcomesin four different eco-evolutionary contexts.Many species of non-social animals live in groups. Aggregating ingroups often has both benefits and costs that depend on group size. Thanksto the benefits of aggregation, population growth likely depends positivelyon population density when it is small. This phenomenon, the Allee effect,has been hypothesized to explain the evolution of aggregation behavior. Ifind that the Allee effect alone does not lead to the evolution whenpopulation dynamics is explicitly accounted for. Some other mechanisms,such as frequent needs for colonizing new patches or anti-aggregation,should be invoked to explain why aggregation behavior could evolve.Phenotypic plasticity is the ability of a genotype to express distinctphenotypes when exposed to different environments. Although it is oftenshown to be adaptive and not costly, highly plastic organisms are rare. Paststudies demonstrated some potential reasons. I test another possibility; costsmay arise from sexual selection because highly plastic individuals may beless preferred as a mate. I show that, even in the absence of the direct cost ofplasticity, the level of plasticity remained low at intermediate strengths ofassortative mating. This pattern is robust across wide ranges of parametervalues.Ecological speciation occurs when ecologically divergent selectionbetween environments causes reproductive isolation between divergingsubpopulations. Several verbal models of ecological speciation emphasizethe roles of phenotypic plasticity in promoting speciation. The complexprocesses involved in speciation, however, are difficult to be evaluated byverbal accounts. I quantitatively test the proposed idea in a mechanisticmodel of ecological speciation in the presence and absence of plasticity. Ifind conditions under which plasticity can promote or hinder ecologicalspeciation. Plasticity facilitates speciation by producing a gap in thedistributions of expressed phenotypes, which serves as a barrier to gene flowin an assortatively mating population.Ecosystem ecology and evolutionary biology are the least integratedfields in ecology and evolution. Natural selection operating at the individuallevels on traits governing ecosystem functions may affect ecosystemproperties, which may feedback to individuals. I reviewed this idea anddemonstrate the feedback loop by using a simple consumer-resource model

Disturbance and landscape history as a reference for evaluating forest management effects at a regional scale : examples from the Coast Range of Oregon, USA

History is an invaluable source of information to understand and evaluate management influences on contemporary ecosystems and landscapes. The first two chapters (Chapters 2 and 3) explored the concept of historical range of variability (HRV) in landscape structure and stand structure using a stochastic fire simulation model to simulate presettlement (before 1850) landscapes of the Oregon Coast Range. HRV has been defined as the bounded variability of a system within constraints imposed by larger-scale phenomena (e.g. climate, topography) and without significant modern human influence. HRV of landscapes has been proposed as a guide for biodiversity conservation in the past decade. In Chapter 2, I estimated HRV of a regional landscape and evaluated the similarity of current and alternative future landscapes under two land management scenarios to the conditions within the HRV. The simulation results indicated that historical landscapes of the region were dynamic, composed of patches of various sizes and age classes ranging from 0 to> 800 years as well as numerous small unburned island patches. The current landscape was outside the HRV. The landscape did not return to the HRV in 100 years under either scenario largely because of lack of old-growth forests and overabundance of young forests. This study showed that the HRV can provide a reference condition for concrete, quantitative evaluations of landscape conditions and alternative management scenarios if sufficient data exist for estimating HRV. Departure from HRV can serve as an indicator of landscape conditions, but results depend on scale and quantification of landscape heterogeneity. In Chapter 3, I investigated the HRV in live and dead biomass and examined variability in disturbance history and forest stand development. I calculated biomass as a function of disturbance history. The HRV of live and dead wood biomass distributions revealed that the majority of the landscape historically contained > 500 Mg/ha of live wood and 50-200 Mg/ha of dead wood. The current dead wood condition is outside HRV. There was a wide variation in dead wood biomass because of variations in disturbance history. This study suggests that natural disturbance regimes and stand development are characterized by much larger variation than is typically portrayed or appreciated. The HRV approaches to evaluating landscape conditions need to include both landscape and stand characteristics to better represent ecological differences between managed and unmanaged landscapes. In Chapter 4, I used remotely sensed data and historical vegetation data in a GIS to examine changes occurred in vegetation cover since settlement in two major valleys, the Coquille and Tillamook, in the region. I used existing historical vegetation maps of the two valleys and collected historical vegetation data from the General Land Office (GLO) survey records. I characterized current vegetation conditions using an unsupervised classification of satellite images. Historically, the Coquille Valley was dominated by hardwood trees and the Tillamook was by conifers. Valley bottoms in both areas differed in vegetation from nearby uplands. Tree-covered areas have declined substantially in both valleys as a result of agriculture and development. The historical data offered reference conditions for assessment of changes in biodiversity that have occurred in these unique habitats. This thesis illustrates the benefit of using historical landscape information for better understanding of human influence on the landscape. Historical data often have many assumptions and limitations, but ecological impacts of landscape changes on native biota can be better understood by comparisons with historical conditions

Increased fluctuation in a butterfly metapopulation leads to diploid males and decline of a hyperparasitoid

Climate change can increase spatial synchrony of population dynamics, leading to large-scale fluctuation that destabilizes communities. High trophic level species such as parasitoids are disproportionally affected because they depend on unstable resources. Most parasitoid wasps have complementary sex determination, producing sterile males when inbred, which can theoretically lead to population extinction via the diploid male vortex (DMV). We examined this process empirically using a hyperparasitoid population inhabiting a spatially structured host population in a large fragmented landscape. Over four years of high host butterfly metapopulation fluctuation, diploid male production by the wasp increased, and effective population size declined precipitously. Our multitrophic spatially structured model shows that host population fluctuation can cause local extinctions of the hyperparasitoid because of the DMV. However, regionally it persists because spatial structure allows for efficient local genetic rescue via balancing selection for rare alleles carried by immigrants. This is, to our knowledge, the first empirically based study of the possibility of the DMV in a natural host–parasitoid system.Peer reviewe

A modified niche model for generating food webs with stage-structured consumers: The stabilizing effects of life-history stages on complex food webs

1. Almost all organisms grow in size during their lifetime and switch diets, trophic positions, and interacting partners as they grow. Such ontogenetic development introduces life-history stages and flows of biomass between the stages through growth and reproduction. However, current research on complex food webs rarely considers life-history stages. The few previously proposed methods do not take full advantage of the existing food web structural models that can produce realistic food web topologies. 2. We extended the niche model by Williams &amp; Martinez (2000) to generate food webs that included trophic species with a life-history stage structure. Our method aggregated trophic species based on niche overlap to form a life-history structured population; therefore, it largely preserved the topological structure of food webs generated by the niche model. We applied the theory of allometric predator-prey body mass ratio and parameterized an allometric bioenergetic model augmented with biomass flow between stages via growth and reproduction to study the effects of a stage structure on the stability of food webs. 3. When life-history stages were linked via growth and reproduction, fewer food webs persisted while persisting food webs tended to retain more trophic species. Topological differences between persisting linked and unlinked food webs were small to modest. Temporal variability of biomass dynamics and slopes of biomass spectra were lower in the linked food webs than the unlinked ones, suggesting that a life-history stage structure enhanced stability of complex food webs. 4. Our results suggest a positive relationship between the complexity and stability of complex food webs. A life-history stage structure in food webs may play important roles in dynamics of and diversity in food webs.</jats:p