Conceptualizing ecosystem services using social-ecological networks.

Social-ecological networks (SENs) represent the complex relationships between ecological and social systems and are a useful tool for analyzing and managing ecosystem services. However, mainstreaming the application of SENs in ecosystem service research has been hindered by a lack of clarity about how to match research questions to ecosystem service conceptualizations in SEN (i.e., as nodes, links, attributes, or emergent properties). Building from different disciplines, we propose a typology to represent ecosystem service in SENs and identify opportunities and challenges of using SENs in ecosystem service research. Our typology provides guidance for this growing field to improve research design and increase the breadth of questions that can be addressed with SEN to understand human-nature interdependencies in a changing world.

Efficient pedigree recording for fast population genetics simulation.

In this paper we describe how to efficiently record the entire genetic history of a population in forwards-time, individual-based population genetics simulations with arbitrary breeding models, population structure and demography. This approach dramatically reduces the computational burden of tracking individual genomes by allowing us to simulate only those loci that may affect reproduction (those having non-neutral variants). The genetic history of the population is recorded as a succinct tree sequence as introduced in the software package msprime, on which neutral mutations can be quickly placed afterwards. Recording the results of each breeding event requires storage that grows linearly with time, but there is a great deal of redundancy in this information. We solve this storage problem by providing an algorithm to quickly 'simplify' a tree sequence by removing this irrelevant history for a given set of genomes. By periodically simplifying the history with respect to the extant population, we show that the total storage space required is modest and overall large efficiency gains can be made over classical forward-time simulations. We implement a general-purpose framework for recording and simplifying genealogical data, which can be used to make simulations of any population model more efficient. We modify two popular forwards-time simulation frameworks to use this new approach and observe efficiency gains in large, whole-genome simulations of one to two orders of magnitude. In addition to speed, our method for recording pedigrees has several advantages: (1) All marginal genealogies of the simulated individuals are recorded, rather than just genotypes. (2) A population of N individuals with M polymorphic sites can be stored in O(N log N + M) space, making it feasible to store a simulation's entire final generation as well as its history. (3) A simulation can easily be initialized with a more efficient coalescent simulation of deep history. The software for recording and processing tree sequences is named tskit.

Stochastic Evolutionary Demography under a Fluctuating Optimum Phenotype.

Many natural populations exhibit temporal fluctuations in abundance that are consistent with external forcing by a randomly changing environment. As fitness emerges from an interaction between the phenotype and the environment, such demographic fluctuations probably include a substantial contribution from fluctuating phenotypic selection. We study the stochastic population dynamics of a population exposed to random (plus possibly directional) changes in the optimum phenotype for a quantitative trait that evolves in response to this moving optimum. We derive simple analytical predictions for the distribution of log population size over time both transiently and at stationarity under Gompertz density regulation. These predictions are well matched by population- and individual-based simulations. The log population size is approximately reverse gamma distributed, with a negative skew causing an excess of low relative to high population sizes, thus increasing extinction risk relative to a symmetric (e.g., normal) distribution with the same mean and variance. Our analysis reveals how the mean and variance of log population size change with the variance and autocorrelation of deviations of the evolving mean phenotype from the optimum. We apply our results to the analysis of evolutionary rescue in a stochastic environment and show that random fluctuations in the optimum can substantially increase extinction risk by both reducing the expected growth rate and increasing the variance of population size by several orders of magnitude.

Predicting evolutionary rescue via evolving plasticity in stochastic environments.

Phenotypic plasticity and its evolution may help evolutionary rescue in a novel and stressful environment, especially if environmental novelty reveals cryptic genetic variation that enables the evolution of increased plasticity. However, the environmental stochasticity ubiquitous in natural systems may alter these predictions, because high plasticity may amplify phenotype-environment mismatches. Although previous studies have highlighted this potential detrimental effect of plasticity in stochastic environments, they have not investigated how it affects extinction risk in the context of evolutionary rescue and with evolving plasticity. We investigate this question here by integrating stochastic demography with quantitative genetic theory in a model with simultaneous change in the mean and predictability (temporal autocorrelation) of the environment. We develop an approximate prediction of long-term persistence under the new pattern of environmental fluctuations, and compare it with numerical simulations for short- and long-term extinction risk. We find that reduced predictability increases extinction risk and reduces persistence because it increases stochastic load during rescue. This understanding of how stochastic demography, phenotypic plasticity, and evolution interact when evolution acts on cryptic genetic variation revealed in a novel environment can inform expectations for invasions, extinctions, or the emergence of chemical resistance in pests.

Allee effect from parasite spill-back.

The exchange of native pathogens between wild and domesticated animals can lead to novel disease threats to wildlife. However, the dynamics of wild host-parasite systems exposed to a reservoir of domesticated hosts are not well understood. A simple mathematical model reveals that the spill-back of native parasites from domestic to wild hosts may cause a demographic Allee effect in the wild host population. A second model is tailored to the particulars of pink salmon (Oncorhynchus gorbuscha) and salmon lice (Lepeophtheirus salmonis), for which parasite spill-back is a conservation and fishery concern. In both models, parasite spill-back weakens the coupling of parasite and wild host abundance-particularly at low host abundance-causing parasites per host to increase as a wild host population declines. These findings show that parasites shared across host populations have effects analogous to those of generalist predators and can similarly cause an unstable equilibrium in a focal host population that separates persistence and extirpation. Allee effects in wildlife arising from parasite spill-back are likely to be most pronounced in systems where the magnitude of transmission from domestic to wild host populations is high because of high parasite abundance in domestic hosts, prolonged sympatry of domestic and wild hosts, a high transmission coefficient for parasites, long-lived parasite larvae, and proximity of domesticated populations to wildlife migration corridors.