Predator-Prey Evolution from an Eco-evolutionary Trade-off Model: The Role of Trait Differentiation.

We develop a novel eco-evolutionary modelling framework and demonstrate its efficacy by simulating the evolution of trait distributions in predator and prey populations. The eco-evolutionary modelling framework assumes that population traits have beta distributions and defines canonical equations for the dynamics of each total population size, the population's average trait value, and a measure of the population's trait differentiation. The trait differentiation is included in the modelling framework as a phenotype analogue, Q, of Wright's fixation index [Formula: see text], which is inversely related to the sum of the beta distribution shape parameters. The canonical equations may be used as templates to describe the evolution of population trait distributions in many ecosystems that are subject to stabilising selection. The solutions of the "population model" are compared with those of a "phenotype model" that simulates the growth of each phenotype as it interacts with every other phenotype under the same trade-offs. The models assume no sources of new phenotypic variance, such as mutation or gene flow. We examine a predator-prey system in which each population trades off growth against mortality: the prey optimises devoting resources to growth or defence against predation; and the predator trades off increasing its attack rate against increased mortality. Computer solutions with stabilising selection reveal very close agreement between the phenotype and population model results, which both predict that evolution operates to stabilise an initially oscillatory system. The population model reduces the number of equations required to simulate the eco-evolutionary system by several orders of magnitude, without losing verisimilitude for the overarching population properties. The population model also allows insights into the properties of the system that are not available from the equivalent phenotype model.

South polar skua (Catharacta maccormicki) as biovectors for long-range transport of persistent organic pollutants to Antarctica.

Migratory bird species may serve as vectors of contaminants to Antarctica through the local deposition of guano, egg abandonment, or mortality. To further investigate this chemical input pathway, we examined the contaminant burdens and profiles of the migratory South polar skua (Catharacta maccormicki) and compared them to the endemic Adélie penguin (Pygoscelis adeliae). A range of persistent organic pollutants were targeted in muscle and guano to facilitate differentiation of likely exposure pathways. A total of 56 of 65 targeted analytes were detected in both species, but there were clear profile and magnitude differences between the species. The South polar skua and Adélie penguin muscle tissue burdens were dominated by p,p'-dichlorodiphenyldichloroethylene (mean 5600 ng g-1 lw and 330 ng g-1 lw respectively) and hexachlorobenzene (mean 2500 ng g-1 lw and 570 ng g-1 lw respectively), a chemical profile characteristic of the Antarctic and Southern Ocean region. Species profile differences, indicative of exposure at different latitudes, were observed for polychlorinated biphenyls (PCBs), with lower chlorinated congeners and deca-chlorinated PCB-209 detected in South polar Skua, but not in Adélie penguins. Notably, the more recently used perfluoroalkyl substances and the brominated flame retardants, hexabromocyclododecane and tetrabromobisphenol A, were detected in both species. This finding suggests local exposure, given the predicted slow and limited long-range environmental transport capacity of these compounds to the eastern Antarctic sector.

The eco-evolutionary modelling of populations and their traits using a measure of trait differentiation.

We develop new equations for the eco-evolutionary dynamics of populations and their traits. These equations resolve the change in the phenotypic differentiation within a population, which better estimates how the variance of the trait distribution changes. We note that traits may be bounded, assume they may be described by beta distributions with small variances, and develop a coupled ordinary differential equation system to describe the dynamics of the total population, the mean trait value, and a measure of phenotype differentiation. The variance of the trait in the population is calculated from its mean and the population's phenotype differentiation. We consider an example of two competing plant populations to demonstrate the efficacy of the new approach. Each population may trade-off its growth rate against its susceptibility to direct competition from the other population. We create two models of this system: a population model based on our new eco-evolutionary equations; and a phenotype model, in which the growth or demise of each fraction of each population with a defined phenotype is simulated as it interacts with a shared limiting resource and its competing phenotypes and populations. Comparison of four simulation scenarios reveals excellent agreement between the predicted quantities from both models: total populations, the average trait values, the trait variances, and the degree of phenotypic differentiation within each population. In each of the four scenarios simulated, three of which are initially subject to competitive exclusion in the absence of evolution, the populations adapt to coexist. One population maximises growth and dominates, while the other minimises competitive losses. These simulations suggest that our new eco-evolutionary equations may provide an excellent approximation to phenotype changes in populations.

Coral reefs as a source of climate-active aerosols.

We review the evidence for bio-regulation by coral reefs of local climate through stress-induced emissions of aerosol precursors, such as dimethylsulfide. This is an issue that goes to the core of the coral ecosystem's ability to maintain homeostasis in the face of increasing climate change impacts and other anthropogenic pressures. We examine this through an analysis of data on aerosol emissions by corals of the Great Barrier Reef, Australia. We focus on the relationship with local stressors, such as surface irradiance levels and sea surface temperature, both before and after notable coral bleaching events. We conclude that coral reefs may be able to regulate their exposure to environmental stressors through modification of the optical properties of the atmosphere, however this ability may be impaired as climate change intensifies.

The emergence of new trophic levels in eco-evolutionary models with naturally-bounded traits.

Ecosystems and food webs are structured into trophic levels of who eats whom. Species that occupy higher trophic levels have less available energy and higher energetic costs than species at lower trophic levels. So why do higher trophic levels exist? What processes generate new trophic levels? We consider a heuristic eco-evolutionary model based on simple Lotka-Volterra equations, where the evolution of traits is described by a generalisation of Lande's equation. The transition from competition to predation in this simplest of models is a successful, safe strategy for a population, and suggests a propensity to develop new trophic levels may be an inherent property of ecosystems. Numerical simulations with a more complex eco-evolutionary model of interacting plant and herbivore populations display the emergence of a new trophic level as an alternative to continued competition. These simulations reveal that new trophic levels may arise naturally from ecosystems because a robust strategy for a population in the presence of a strong competitor that could dominate or potentially extinguish them, is to predate upon the competitor. The same properties that make the competitor strong make it an ideal prey, suggesting the rubric that it is better to eat a strong competitor than to continue competing.

Simulating Eco-evolutionary Processes in an Obligate Pollination Model with a Genetic Algorithm.

Pollination interactions are common, and their maintenance is critical for many food crops upon which human populations depend. Pollination is a mutualism interaction; together with predation and competition, mutualism makes up the triumvirate of fundamental interactions that control population dynamics. Here we examine pollination interactions (nectar reward for gamete transport service) using a simple heuristic model similar to the Lotka-Volterra models that have underpinned our understanding of predation and competition so effectively since the 1920s. We use a genetic algorithm to simulate the eco-evolutionary interactions of the plant and pollinator populations and examine the distributions of the parameter values and zero isoclines to infer the relative ubiquity of the various eco-evolutionary outcomes possible in the model. Our results suggest that trade-offs between costs and benefits for the pollinator may be a key component of obligate pollination systems in achieving adaptive success creating and stably occupying mutualist niches.

Obligate Mutualism in an Extended Consumer-Resource Framework.

The development of a theory to underpin the obligate mutualist interactions that appear to be ubiquitous in nature has not proceeded at the same pace as the development of theory to support competition and predation. A constraint may be that obligate mutualism appears unable to be presented in the simple linear models that have so successfully served as heuristics for the other interactions. A number of simple nonlinear models have been used to propose explanations of obligate mutualism, but these solutions are often predicated on careful choices of functional forms. We present a theory of obligate mutualism in an explicit mass-conserving framework using simple models that are robust to choices of functional forms.

Effects of ocean warming and coral bleaching on aerosol emissions in the Great Barrier Reef, Australia.

It is proposed that emissions of volatile sulfur compounds by coral reefs contribute to the formation of a biologically-derived feedback on sea surface temperature (SST) through the formation of marine biogenic aerosol (MBA). The direction and strength of this feedback remains uncertain and constitutes a fundamental constraint on predicting the ability of corals to cope with future ocean warming. We investigate the effects of elevated SST and irradiance on satellite-derived fine-mode aerosol optical depth (AOD) throughout the Great Barrier Reef, Australia (GBR) over an 18-year time period. AOD is positively correlated with SST and irradiance and increases two-fold during spring and summer with high frequency variability. As the influence of non-biogenic and distant aerosol sources are found to be negligible, the results support recent findings that the 2,300 km stretch of coral reefs can be a substantial source of biogenic aerosol and thus, influence local ocean albedo. Importantly however, a tipping point in the coral stress response is identified, whereby thermal stress reaches a point that exceeds the capacity of corals to influence local atmospheric properties. Beyond this point, corals may become more susceptible to permanent damage with increasing stress, with potential implications for mass coral bleaching events.

Coral reef aerosol emissions in response to irradiance stress in the Great Barrier Reef, Australia.

We investigate the correlation between stress-related compounds produced by corals of the Great Barrier Reef (GBR) and local atmospheric properties-an issue that goes to the core of the coral ecosystem's ability to survive climate change. We relate the variability in a satellite decadal time series of fine-mode aerosol optical depth (AOD) to a coral stress metric, formulated as a function of irradiance, water clarity, and tide, at Heron Island in the southern GBR. We found that AOD was correlated with the coral stress metric, and the correlation increased at low wind speeds, when horizontal advection of air masses was low and the production of non-biogenic aerosols was minimal. We posit that coral reefs may be able to protect themselves from irradiance stress during calm weather by affecting the optical properties of the atmosphere and local incident solar radiation.

Uptake and Depuration Kinetics Influence Microplastic Bioaccumulation and Toxicity in Antarctic Krill ( Euphausia superba).

The discarding of plastic products has led to the ubiquitous occurrence of microplastic particles in the marine environment. The uptake and depuration kinetics of ingested microplastics for many marine species still remain unknown despite its importance for understanding bioaccumulation potential to higher trophic level consumers. In this study, Antarctic krill ( Euphausia superba) were exposed to polyethylene microplastics to quantify acute toxicity and ingestion kinetics, providing insight into the bioaccumulation potential of microplastics at the first-order consumer level. In the 10 day acute toxicity assay, no mortality or dose-dependent weight loss occurred in exposed krill, at any of the exposure concentrations (0, 10, 20, 40, or 80% plastic diet). Krill exposed to a 20% plastic diet for 24 h displayed fast uptake (22 ng mg-1 h-1) and depuration (0.22 h-1) rates, but plastic uptake did not reach steady state. Efficient elimination also resulted in no bioaccumulation over an extended 25 day assay, with most individuals completely eliminating their microplastic burden in less than 5 days post exposure. Our results support recent findings of limited acute toxicity of ingested microplastics at this trophic level, and suggest sublethal chronic end points should be the focus of further ecotoxicological investigation.

Signals from the south; humpback whales carry messages of Antarctic sea-ice ecosystem variability.

Southern hemisphere humpback whales (Megaptera novaeangliae) rely on summer prey abundance of Antarctic krill (Euphausia superba) to fuel one of the longest-known mammalian migrations on the planet. It is hypothesized that this species, already adapted to endure metabolic extremes, will be one of the first Antarctic consumers to show measurable physiological change in response to fluctuating prey availability in a changing climate; and as such, a powerful sentinel candidate for the Antarctic sea-ice ecosystem. Here, we targeted the sentinel parameters of humpback whale adiposity and diet, using novel, as well as established, chemical and biochemical markers, and assembled a time trend spanning 8 years. We show the synchronous, inter-annual oscillation of two measures of humpback whale adiposity with Southern Ocean environmental variables and climate indices. Furthermore, bulk stable isotope signatures provide clear indication of dietary compensation strategies, or a lower trophic level isotopic change, following years indicated as leaner years for the whales. The observed synchronicity of humpback whale adiposity and dietary markers, with climate patterns in the Southern Ocean, lends strength to the role of humpback whales as powerful Antarctic sea-ice ecosystem sentinels. The work carries significant potential to reform current ecosystem surveillance in the Antarctic region.

Persistent Organic Pollutants in the East Antarctic Atmosphere: Inter-Annual Observations from 2010 to 2015 Using High-Flow-Through Passive Sampling.

In the first multiyear sampling effort for POPs in the eastern Antarctic atmosphere, 32 PCBs and 38 organochlorine pesticides were targeted in air collected with a high-flow-through passive sampler. Agricultural chemicals were found to dominate atmospheric profiles, in particular HCB and endosulfan-I, with average concentrations of 12 600 and 550 fg/m3, respectively. HCB showed higher concentrations in the austral summer, indicative of local, temperature-dependent volatilisation, while endosulfan-I appeared to show fresh, late-austral-summer input followed by temporally decreasing levels throughout the year. The current-use herbicide, trifluralin, and the legacy pesticides mirex and toxaphene, were detected in Antarctic air for the first time. Trifluralin was observed at low but increasing levels over the five-year period. Its detection in the Antarctic atmosphere provides evidence of its persistence and long-range environmental transport capability. While a time frame of five years exceeds the duration of most Antarctic air monitoring efforts, it is projected that continuous monitoring at the decadal scale is required to detect an annual 10% change in atmospheric concentrations of key analytes. This finding emphasizes the importance of continuous, long-term monitoring efforts in polar regions, that serve a special role as sentinel environments of hemispheric chemical usage trends.

Spring Melt and the Redistribution of Organochlorine Pesticides in the Sea-Ice Environment: A Comparative Study between Arctic and Antarctic Regions.

Complementary sampling of air, snow, sea-ice, and seawater for a range of organochlorine pesticides (OCPs) was undertaken through the early stages of respective spring sea-ice melting at coastal sites in northeast Greenland and eastern Antarctica to investigate OCP concentrations and redistribution during this time. Mean concentrations in seawater, sea-ice and snow were generally greater at the Arctic site. For example, α-HCH was found to have the largest concentrations of all analytes in Arctic seawater and sea-ice meltwater samples (224-253 and 34.7-48.2 pg·L-1 respectively compared to 1.0-1.3 and <0.63 pg·L-1 respectively for Antarctic samples). Differences in atmospheric samples were generally not as pronounced however. Findings suggest that sea-ice OCP burdens originate from both snow and seawater. The distribution profile between seawater and sea-ice showed a compound-dependency for Arctic samples not evident with those from the Antarctic, possibly due to full submersion of sea-ice at the former. Seasonal sea-ice melt processes may alter the exchange rates of selected OCPs between air and seawater, but are not expected to reverse their direction, which fugacity modeling indicates is volatilisation in the Arctic and net deposition in the Antarctic. These predictions are consistent with the limited current observations.

Air-Seawater Exchange of Organochlorine Pesticides in the Southern Ocean between Australia and Antarctica.

This study contributes new data on the spatial variability of persistent organic pollutants in the Indian-Pacific sector of the Southern Ocean and represents the first empirical data obtained from this region in 25 years. Paired high-volume atmospheric and seawater samples were collected along a transect between Australia and Antarctica to investigate the latitudinal dependence of the occurrence and distribution of legacy organochlorine pesticides (OCPs) and the current use pesticide chlorpyrifos in the Southern Ocean. Dissolved ΣHCH and dieldrin concentrations decreased linearly with increasing latitude from 7.7 to 3.0 and from 1.0 to 0.6 pg·L(-1), respectively. There was no consistent trend observed in the latitudinal profile of atmospheric samples; however, some compounds (such as dieldrin) showed reduced concentrations from 7.5-3.4 to 2.7-0.65 pg·m(-3) at the highest latitudes south of the Polar Front. Chlorpyrifos was found in samples from this area for the first time. Estimated air-seawater fugacity ratios and fluxes indicate a current net deposition between -3600 and -900, -6400 and -400, and -1400 and -200 (pg·m(-2)·d(-1)) for γ-HCH, dieldrin, and chlorpyrifos, respectively. These findings suggest that, under current climatic conditions, the Southern Ocean reservoir in the Indian-Pacific sector serves as an environmental sink rather than a source of OCPs to the atmosphere.

On the formulation of environmental fugacity models and their numerical solutions.

Multimedia models based on chemical fugacity, solved numerically, play an important role in investigating and quantifying the environmental fate of chemicals such as persistent organic pollutants. These models have been used extensively in studying the local and global distribution of chemicals in the environment. The present study describes potential sources of error that may arise from the formulation and numerical solution of environmental fugacity models. The authors derive a general fugacity equation for the rate of change of mass in an arbitrary volume (e.g., an environmental phase). Deriving this general equation makes clear several assumptions that are often not articulated but can be important for successfully applying multimedia fugacity models. It shows that the homogeneity of fugacity and fugacity capacity in a volume (the homogeneity assumption) is fundamental to formulating discretized fugacity models. It also shows that when using the fugacity rather than mass as the state-variable, correction terms may be necessary to accommodate environmental factors such as varying phase temperatures and volume. Neglecting these can lead to conservation errors. The authors illustrate the manifestation of these errors using heuristic multimedia fugacity models. The authors also show that there are easily avoided errors that can arise in mass state-variable models if variables are not updated appropriately in the numerical integration scheme. Environ Toxicol Chem 2016;35:2182-2191. © 2016 SETAC.

An Antarctic research station as a source of brominated and perfluorinated persistent organic pollutants to the local environment.

This study investigated the role of a permanently manned Australian Antarctic research station (Casey Station) as a source of contemporary persistent organic pollutants (POPs) to the local environment. Polybrominated diphenyl ethers (PBDEs) and poly- and perfluoroalkylated substances (PFASs) were found in indoor dust and treated wastewater effluent of the station. PBDE (e.g., BDE-209 26-820 ng g(-1) dry weight (dw)) and PFAS levels (e.g., PFOS 3.8-2400 ng g(-1) (dw)) in dust were consistent with those previously reported in homes and offices from Australia, reflecting consumer products and materials of the host nation. The levels of PBDEs and PFASs in wastewater (e.g., BDE-209 71-400 ng L(-1)) were in the upper range of concentrations reported for secondary treatment plants in other parts of the world. The chemical profiles of some PFAS samples were, however, different from domestic profiles. Dispersal of chemicals into the immediate marine and terrestrial environments was investigated by analysis of abiotic and biotic matrices. Analytes showed decreasing concentrations with increasing distance from the station. This study provides the first evidence of PFAS input to Polar regions via local research stations and demonstrates the introduction of POPs recently listed under the Stockholm Convention into the Antarctic environment through local human activities.

A model to resolve organochlorine pharmacokinetics in migrating humpback whales.

Humpback whales are iconic mammals at the top of the Antarctic food chain. Their large reserves of lipid-rich tissues such as blubber predispose them to accumulation of lipophilic contaminants throughout their lifetime. Changes in the volume and distribution of lipids in humpback whales, particularly during migration, could play an important role in the pharmacokinetics of lipophilic contaminants such as the organochlorine pesticide hexachlorobenzene (HCB). Previous models have examined constant feeding and nonmigratory scenarios. In the present study, the authors develop a novel heuristic model to investigate HCB dynamics in a humpback whale and its environment by coupling an ecosystem nutrient-phytoplankton-zooplankton-detritus (NPZD) model, a dynamic energy budget (DEB) model, and a physiologically based pharmacokinetic (PBPK) model. The model takes into account the seasonal feeding pattern of whales, their energy requirements, and fluctuating contaminant burdens in the supporting plankton food chain. It is applied to a male whale from weaning to maturity, spanning 20 migration and feeding cycles. The model is initialized with environmental HCB burdens similar to those measured in the Southern Ocean and predicts blubber HCB concentrations consistent with empirical concentrations observed in a southern hemisphere population of male, migrating humpback whales. Results show for the first time some important details of the relationship between energy budgets and organochlorine pharmacokinetics.

Uptake of zwitterionic antibiotics by rice (Oryza sativa L.) in contaminated soil.

Antibiotics, including members of the tetracycline and fluoroquinolone families, are emerging organic environmental contaminants. Uptake from soil by plants is a means for antibiotics to enter terrestrial food chains. Chemical exchange between plant and the soil/water matrix occurs simultaneously with degradation in the soil/water matrix. In this study, the comparative temporal behaviour of rice (Oryza sativa L.) towards the zwitterionic antibiotics oxytetracycline, chlortetracycline and norfloxacin at initial soil/water concentrations of 10, 20 and 30 µg g(-1) (dry weight) is investigated. This is accomplished within the framework of an activity-based mass-conserving dynamic model. Plant antibiotic concentrations are observed to increase to a maximum then decline. Maximum concentrations in rice are compound-dependent linear functions of initial soil/water concentrations, but the relationships are not related to the compound octan-1-ol/water distribution ratio (DOW). The times required to attain maximal concentrations are independent of initial soil/water levels for a given antibiotic, but again vary between antibiotics and are not related to DOW values. Translocation from root to other tissues is not observed. The magnitudes of Root Concentration Factors (RCFs), the ratio of root and soil/water concentrations, are consistent with significant sorption to soil and consequent relatively low concentrations in interstitial water.

Constructing ecologies.

We synthesize the generic properties of ecologically realistic multi-trophic level models and define criteria for ecological realism. We define an "ecospace" in which all ecologically realistic dynamics are confined, and construct "resource rays" that define the resources available to each species at every point in the ecospace. Resource rays for a species are lines from a vertex of maximum resource to the opposite boundary where no resources are available. The growth functions of all biota normally decrease along their resource rays, and change sign from positive to negative. This property prescribes that each species must have a zero isosurface within the ecospace. We illustrate our conditions on a highly cited three trophic level model from population dynamics, showing how to extend this system biologically consistently to a closed ecological system. Our synthesis extends the concept of carrying capacity of population models to explicitly include exhaustion of limiting resources, and so allows for population biology models to be considered as ecologically closed systems with respect to a key limiting nutrient. This approach unifies many theoretical and applied models in a common biogeochemical framework, facilitates better understanding of the key structures of complex ecologies, and suggests strategies for efficient design of experiments.

Predicting the accumulation of organic contaminants from soil by plants.

Analytic expressions for maximum chemical concentration attained in plants, and time this takes for uptake from surrounding soil were derived from a simple two-compartment soil/water-plant model. To illustrate, for the antibiotic norflxacin undergoing first order loss in the soil/water phase with a rate constant of 0.544 days⁻¹, maximum concentration in soybean P(MAX) is predicted to occur after 2.79 days exposure and be independent of initial soil/water concentration SW0 of 52.5 mg kg⁻¹ dry weight. For soybean, the relationship between P(MAX) and SW0 is P(MAX) = 0.047SW0, resulting in predicted maximum levels of 2.20 mg kg⁻¹ dry weight. Modelled plant concentrations agreed well with experimental data (R² = 0.91).