The Effects of Seed Dispersal on the Simulation of Long-Term Forest Landscape Change

The study of forest landscape change requires an understanding of the complex interactions of both spatial and temporal factors. Traditionally, forest gap models have been used to simulate change on small and independent plots. While gap models are useful in examining forest ecological dynamics across temporal scales, large, spatial processes, such as seed dispersal, cannot be realistically simulated across large landscapes. To simulate seed dispersal, spatially explicit landscape models that track individual species distribution are needed. We used such a model, LANDIS, to illustrate the implications of seed dispersal for simulating forest landscape change. On an artificial open landscape with a uniform environment, circular-shaped tree species establishment patterns resulted from the simulations, with areas near seed sources more densely covered than areas further from seed sources. Because LANDIS simulates at 10-y time steps, this pattern reflects an integration of various possible dispersal shapes and establishment that are caused by the annual variations in climate and other environmental variables. On real landscapes, these patterns driven only by species dispersal radii are obscured by other factors, such as species competition, disturbance, and landscape structure. To further demonstrate the effects of seed dispersal, we chose a fairly disturbed and fragmented forest landscape (approximately 500,000 ha) in northern Wisconsin. We compared the simulation results of a map with tree species (seed source locations) realistically parameterized (the real scenario) against a randomly parameterized species map (the random scenario). Differences in the initial seed source distribution lead to different simulation results of species abundance with species abundance starting at identical levels under the two scenarios. This is particularly true for the first half of the model run (0–250 y). Under the random scenario, infrequently occurring and shade tolerant species tend to be overestimated, while midabundant and midshade tolerant species tend to be underestimated. The over- and underestimation of species abundance diminish when examining long-term (500 y) landscape dynamics, because stochastic factors, such as fire, tend to make the landscapes under both scenarios converge. However, differences in spatial patterns, and especially species age-cohort distributions, can persist under the two scenarios for several hundred years. Received 24 November 1998; accepted 17 March 1999.     

Testing the mechanisms by which source-sink dynamics alter competitive outcomes in a model system

Dispersal among sites can affect within-site competitive outcomes via source-sink dynamics. Source-sink dynamics are thought to affect competitive outcomes primarily via spatial subsidies: by redistributing individuals from sources to sinks, source-sink dynamics can alter competitive outcomes in both sources and sinks. However, dispersal also can affect competitive outcomes via demography modification, which occurs when dispersal alters the parameters governing species' per capita demographic rates. For instance, dispersal of exploitative competitors might cause extinction of some of the resources for which competition occurs, thereby altering the competition coefficients. I used protist microcosms as a model system to test whether spatial subsidies alone could explain the effects of source-sink dynamics on competitive outcomes. I examined the long-term outcome of exploitative competition among three bacterivorous ciliate protists in microcosms of high enrichment (sources) and low enrichment (sinks) in both the presence and the absence of dispersal. Dispersal altered competitive outcomes. Fitting mathematical models to the population dynamics revealed that spatial subsidies were insufficient to account for the effects of dispersal. Fitting alternative models strongly suggested that demography modification was an important determinant of competitive outcomes. These results provide the first evidence that dispersal does not simply redistribute competitors but can alter their per capita demographic rates.     

Strong microsite control of seedling recruitment in tundra

The inclusion of environmental variation in studies of recruitment is a prerequisite for realistic predictions of the responses of vegetation to a changing environment. We investigated how seedling recruitment is affected by seed availability and microsite quality along a steep environmental gradient in dry tundra. A survey of natural seed rain and seedling density in vegetation was combined with observations of the establishment of 14 species after sowing into intact or disturbed vegetation. Although seed rain density was closely correlated with natural seedling establishment, the experimental seed addition showed that the microsite environment was even more important. For all species, seedling emergence peaked at the productive end of the gradient, irrespective of the adult niches realized. Disturbance promoted recruitment at all positions along the environmental gradient, not just at high productivity. Early seedling emergence constituted the main temporal bottleneck in recruitment for all species. Surprisingly, winter mortality was highest at what appeared to be the most benign end of the gradient. The results highlight that seedling recruitment patterns are largely determined by the earliest stages in seedling emergence, which again are closely linked to microsite quality. A fuller understanding of microsite effects on recruitment with implications for plant community assembly and vegetation change is provided.     

The temporal structure of the environment may influence range expansions during climate warming

Understanding the processes that influence range expansions during climate warming is paramount for predicting population extirpations and preparing for the arrival of non‐native species. While climate warming occurs over a background of variation due to cyclical processes and irregular events, the temporal structure of the thermal environment is largely ignored when forecasting the dynamics of non‐native species. Ecological theory predicts that high levels of temporal autocorrelation in the environment – relatedness between conditions occurring in close temporal proximity – will favor populations that would otherwise have an average negative growth rate by increasing the duration of favorable environmental periods. Here, we invoke such theory to explain the success of biological invasions and evaluate the hypothesis that sustained periods of high environmental temperature can act synergistically with increases in mean temperature to favor the establishment of non‐native species. We conduct a 60‐day field mesocosm experiment to measure the population dynamics of the non‐native cladoceran zooplankter Daphnia lumholtzi and a native congener Daphnia pulex in ambient temperature environments (control), warmed with recurrent periods of high environmental temperatures (uncorrelated‐warmed), or warmed with sustained periods of high environmental temperatures (autocorrelated‐warmed), such that both warmed treatments exhibited the same mean temperature but exhibited different temporal structures of their thermal environments. Maximum D. lumholtzi densities in the warmed‐autocorrelated treatment were threefold and eightfold higher relative to warmed‐uncorrelated and control treatments, respectively. Yet, D. lumholtzi performed poorly across all experimental treatments relative to D. pulex and were undetectable by the end of the experiment. Using mathematical models, we show that this increase in performance can occur alongside increasing temporal autocorrelation and should occur over a broad range of warming scenarios. These results provide both empirical and theoretical evidence that the temporal structure of the environment can influence the performance of species undergoing range expansions due to climate warming.     

The Flask model: emergence of nutrient-recycling microbial ecosystems and their disruption by environment-altering 'rebel' organisms

Here we introduce a new model of life–environment interaction, which simulates an evolving microbial community in a 'Fask' of liquid with prescribed inputs of nutrients. The flask is seeded with a clonal population of 'microbes' that are subject to mutation on genetic loci that determine their nutrient uptake patterns, release patterns, and their effects on, and response to, other environmental variables. In contrast to existing models of life-environment interaction, notably Daisyworld, what benefits the individual organisms is decoupled from their 'global' (system-level) effects. A robust property of the model is the emergence of ecosystems that tend toward a state where nutrients are efficiently utilised and differentially recycled, with a correlated increase in total population. Organisms alter the environment as a free 'by-product' of their growth, and their growth is constrained by adverse environmental effects. This introduces environmental feedback, which can disrupt the model ecosystems, even though there are no constraints on the conditions to which the organisms can theoretically adapt. 'Rebel' organisms can appear that grow rapidly by exploiting an under-utilised resource, but in doing so shift the environment away from the state to which the majority of the community are adapted. The result can be a population crash with lossof recycling, followed by later recovery, or in extreme cases, a total extinction of the system. Numerous runs of these 'flask' ecosystems show that tighter environmental constraints on growth make the system more vulnerable to internally generated ecosystem extinction.     

Temporal coincidence of environmental stress events modulates predation rates

Climate warming experiments generally test the ecological effects of constant treatments while neglecting the influence of more realistic patterns of environmental fluctuations. Thus, little is known regarding how the temporal interaction between multiple episodes of thermal stress influences biotic interactions. We measured the sensitivity of predation rate in an intertidal sea star to changing levels of temporal coincidence of underwater and aerial thermal stress events. In laboratory trials, we controlled for intensity, variance and temporal patterning of both underwater and aerial body temperature. Predation rate decreased as underwater and aerial thermal stress episodes became temporally non-coincident, despite a similar intensity and variance among treatments. Experiments under constant conditions were a poor predictor of more complex environmental scenarios because of these strong temporal interactions. Such temporal interactions may be widespread in various ecosystems, suggesting a strong need for empirical studies and models that link environmental complexity, physiology, behaviour and species interactions.     

Anthropogenic disturbance in a changing environment: modelling lifetime reproductive success to predict the consequences of multiple stressors on a migratory population

Animals make behavioural and reproductive decisions that maximise their lifetime reproductive success, and thus their fitness, in light of periodic and stochastic variability of the environment. Modelling the variation of an individual's energy levels formalises this tradeoff and helps to quantify the population‐level consequences of stressors (e.g. disturbance from human activities and environmental change) that can affect behaviour or physiology. In this study, we develop a dynamic state variable model for the spatially explicit behaviour, physiology and reproduction of a female, long‐lived, migratory marine vertebrate. The model can be used to investigate the spatio‐temporal patterns of behaviour and reproduction that allow an individual to maximise its overall reproductive output. We parametrised the model for eastern North Pacific blue whales Balaenoptera musculus, and used it to predict the effects of changing environmental conditions and increasing human disturbance on the population's vital rates. In baseline conditions, the model output had high fidelity to observed energy dynamics, movement patterns and reproductive strategies. Simulated scenarios suggested that environmental changes could have severe consequences on the population's vital rates, but that individuals could tolerate high levels of anthropogenic disturbance. However, this ability depended on where, when and how often disturbance occurred. In scenarios with both environmental change and anthropogenic disturbance, synergistic interactions caused stronger effects than in isolation. In general, larger body size offered a buffer against stochasticity and disturbance, and, consequently, we predicted juveniles to be more susceptible to disturbance. We also predicted that females prioritise their own survival at the expense of the current reproductive attempt, presumably the result of their long lifespan. Our approach provides a general framework to make predictions of the cumulative and synergistic effects of human disturbance and climate change on migratory populations, which can inform effective management and conservation efforts.     

Spatiotemporal population distributions and their implications for species coexistence in a variable environment

A population experiences environmental variation both directly, through effects on life history parameters such as fecundity, and indirectly, through effects on the population distributions of competitors and thus on the distribution of competition. Which spatial and temporal scales of environmental variation most influence the coexistence of two species thus depends in part on the degree to which the resident population responds to different scales of variation. In this paper, I calculate an approximation for a spatiotemporal population distribution as the result of a filter function convolved with the environmental variation. I find that there is no straightforward connection between spatial or temporal scales inherent to an organism's life history, such as mean lifetime or dispersal distance, and the population's sensitivity to variation at different scales. Rather, life history traits interact sensitively with the way environmental variation affects the organism. I comment on the implications for variation-mediated coexistence.     

Movement patterns, social dynamics, and the evolution of cooperation

The structure of social interactions influences many aspects of social life, including the spread of information and behavior, and the evolution of social phenotypes. After dispersal, organisms move around throughout their lives, and the patterns of their movement influence their social encounters over the course of their lifespan. Though both space and mobility are known to influence social evolution, there is little analysis of the influence of specific movement patterns on evolutionary dynamics. We explored the effects of random movement strategies on the evolution of cooperation using an agent-based prisoner’s dilemma model with mobile agents. This is the first systematic analysis of a model in which cooperators and defectors can use different random movement strategies, which we chose to fall on a spectrum between highly exploratory and highly restricted in their search tendencies. Because limited dispersal and restrictions to local neighborhood size are known to influence the ability of cooperators to effectively assort, we also assessed the robustness of our findings with respect to dispersal and local capacity constraints. We show that differences in patterns of movement can dramatically influence the likelihood of cooperator success, and that the effects of different movement patterns are sensitive to environmental assumptions about offspring dispersal and local space constraints. Since local interactions implicitly generate dynamic social interaction networks, we also measured the average number of unique and total interactions over a lifetime and considered how these emergent network dynamics helped explain the results. This work extends what is known about mobility and the evolution of cooperation, and also has general implications for social models with randomly moving agents.     

Eco-Evolutionary Feedback and the Tuning of Proto-Developmental Life Cycles

Multicellular organisms depend on developmental programs to coordinate growth and differentiation from single cells, but the origins of development are unclear. A possible starting point is stochastic phenotypic variation generated by molecular noise. Given appropriate environmental conditions, noise-driven differentiation could conceivably evolve so as to come under regulatory control; however, abiotic conditions are likely to be restrictive. Drawing from an experimental system, we present a model in which environmental fluctuations are coupled to population growth. We show that this coupling generates stable selection for a single optimal strategy that is largely insensitive to environmental conditions, including the number of competitors, carrying capacity of the environment, difference in growth rates among phenotypic variants, and population density. We argue that this optimal strategy establishes stabilizing conditions likely to improve the quality and reliability of information experienced by evolving organisms, thus increasing opportunity for the evolutionary emergence of developmental programs.     

Impacts of environmental variability in open populations and communities: "inflation" in sink environments

Ecological communities are typically open to the immigration and emigration of individuals, and also variable through time. In this paper we argue that interesting and potentially important effects arise when one splices together spatial fluxes and temporal variability. The particular system we examine is a sink habitat, where a species faces deterministic extinction but is rescued by recurrent immigration. We have shown, using a simple extension of the canonical exponential growth model in a time-varying environment, that variation "inflates" the average abundance of sink populations. We can analytically quantify the magnitude of this effect in several special cases (square-wave temporal variation and Gaussian stochastic variation). The inflationary effect can be large in "intermittent" sinks (where there are periods with positive growth), and when temporal variation is strongly autocorrelated. The effect appears to be robust to incorporation of demographic stochasticity (due to discrete birth-death-immigration processes), and to direct density dependence. With discrete generations, however, one can observe a wide range of effects of temporal variation, including depression as well as inflation. We argue that the inflationary effect of temporal variation in sink habitats can have important implications for community structure, because it can increase the average abundance (and hence local impacts) of species that on average are being excluded from a local community. We illustrate the latter effect using a familiar model of exploitative competition for a single limiting resource. We demonstrate that temporal variation can reverse local competitive dominance, even to the extent of allowing an inferior competitor maintained by immigration to exclude a competing species that would be locally superior in a constant environment.     

Threshold assessment,categorical perception,and the evolution of reliable signaling

Animals often use assessment signals to communicate information about their quality to a variety of receivers, including potential mates, competitors, and predators. But what maintains reliable signaling and prevents signalers from signaling a better quality than they actually have? Previous work has shown that reliable signaling can be maintained if signalers pay fitness costs for signaling at different intensities and these costs are greater for lower quality individuals than higher quality ones. Models supporting this idea typically assume that continuous variation in signal intensity is perceived as such by receivers. In many organisms, however, receivers have threshold responses to signals, in which they respond to a signal if it is above a threshold value and do not respond if the signal is below the threshold value. Here, we use both analytical and individual-based models to investigate how such threshold responses affect the reliability of assessment signals. We show that reliable signaling systems can break down when receivers have an invariant threshold response, but reliable signaling can be rescued if there is variation among receivers in the location of their threshold boundary. Our models provide an important step toward understanding signal evolution when receivers have threshold responses to continuous signal variation.     

Pelagic food webs: Responses to environmental processes and effects on the environment

Changes in both the environment and environmental research have led to the development of new protocols and approaches. These new approaches consider both the effects of changes in the global environment on living organisms (i.e. the responses of ecosystems to environmental processes) and the feedback responses of these organisms and ecosystems (i.e. the effects of living organisms on the environment). The present paper focuses on pelagic food webs in aquatic ecosystems. We examine three major effects of global environmental changes on aquatic organisms: (i) the release of pollutants and biological agents in lakes and nearshore marine waters; (ii) the loss of biodiversity and the collapse of commercially exploited resources that were heretofore renewable. We develop detailed examples of the effects of human activities on marine organisms (i.e. the effects of nutrient supply on the structure of pelagic food webs in marine systems. Finally, we examine (iii) the food-web-controlled exchanges of CO₂ between the atmosphere and the ocean, as a feedback effect of pelagic ecosystems on the global environment with respect to the ongoing climate change.     

Limits to elevational distributions in two species of emberizine finches: disentangling the role of interspecific competition, autoecology, and geographic variation in the environment

When species' elevational ranges are wider where putative competitors are absent, researchers have concluded that interspecific competition influences elevational distributions. This overlooks the distinction between factors that limit distributions directly and factors that only influence organisms indirectly through covarying regulators or resources. Because elevation affects organisms indirectly, testing whether competition influences elevational ranges relies on the heretofore untested assumption that the relationship between elevation and factors influencing organisms directly is similar across geography. Focusing on Buarremon brush-finches (Aves: Emberizidae), a group in which distributions represent one of the best examples of the potential role of competition limiting elevational ranges, we show that when distributions are compared along axes of climatic variation, some patterns of elevational range variation do appear to be consistent with predictions of the hypothesis that release from competition underlies expanded elevational ranges in allopatry. However, other patterns of expanded elevational ranges in the absence of putative competitors are better explained by hypothesis related to species' autoecology and geographic variation in the environment. This latter finding cautions against using elevation uncritically as a dimension of ecological niches, and suggests that classical examples of interspecific competition may need re-evaluation.     

Memory shapes microbial populations

Correct decision making is fundamental for all living organisms to thrive under environmental changes. The patterns of environmental variation and the quality of available information define the most favourable strategy among multiple options, from randomly adopting a phenotypic state to sensing and reacting to environmental cues. Cellular memory—the ability to track and condition the time to switch to a different phenotypic state—can help withstand environmental fluctuations. How does memory manifest itself in unicellular organisms? We describe the population-wide consequences of phenotypic memory in microbes through a combination of deterministic modelling and stochastic simulations. Moving beyond binary switching models, our work highlights the need to consider a broader range of switching behaviours when describing microbial adaptive strategies. We show that memory in individual cells generates patterns at the population level coherent with overshoots and non-exponential lag times distributions experimentally observed in phenotypically heterogeneous populations. We emphasise the implications of our work in understanding antibiotic tolerance and, in general, bacterial survival under fluctuating environments.     

Maternal effects and the outcome of interspecific competition

1. Maternal environmental effects create lagged population responses to past environments. Although they are ubiquitous and vary in expression across taxa, it remains unclear if and how their presence alters competitive interactions in ecological communities.
2. Here, we use a discrete‐time competition model to simulate how maternal effects alter competitive dynamics in fluctuating and constant environments. Further, we explore how omitting maternal effects alter estimates of known model parameters from observational time series data.
3. Our simulations demonstrate that (i) maternal effects change competitive outcomes, regardless of whether competitors otherwise interact neutrally or exhibit non‐neutral competitive differences, (ii) the consequences of maternal effects for competitive outcomes are mediated by the temporal structure of environmental variation, (iii) even in constant conditions, competitive outcomes are influenced by species'' maternal effects strategies, and (iv) in observational time series data, omitting maternal effects reduces variation explained by models and biases parameter estimates, including competition coefficients.
4. Our findings demonstrate that the ecological consequences of maternal effects hinge on the competitive environment. Evolutionary biologists have long recognized that maternal effects can be an important but often overlooked strategy buffering populations from environmental change. We suggest that maternal effects are similarly critical to ecology and call for research into maternal effects as drivers of dynamics in populations and communities.

     

Effects of environmental variables on fish feeding ecology: implications for the performance of baited fishing gear and stock assessment

The effectiveness of baited fishing gear ultimately depends upon behaviour of the target species – activity rhythms, feeding motivation, and sensory and locomotory abilities. While any environmental parameter that mediates feeding or locomotion can have an important influence on the active space presented by the bait and fish catchability, few biologists have considered how such variation in behaviour might affect catch per unit effort (CPUE) and the resultant stock abundance estimates or population parameters. This review reveals that environment‐related variation in feeding behaviour can act through four different mechanisms: metabolic processes, sensory limitations, social interactions and direct impacts. Water temperature, light level, current velocity and ambient prey density are likely to have largest effects on fish catchability, potentially affecting variation in CPUE by a factor of ten. Feeding behaviour is also density‐dependent, with both positive and negative effects. Over time and geographic space a target species can occupy wide ranges of environmental conditions, and in certain cases, spatial and temporal variation in feeding biology could have a larger impact on CPUE than patterns of abundance. Temperature, light and current can be measured with relative facility and corrections to stock assessment models are feasible. Making corrections for biological variables such as prey density and bait competitors will be more difficult because the measurements are often not practical and relationships to feeding catchability are more complex and poorly understood. There is a critical need for greater understanding of how environmental variables affect feeding‐related performance of baited fishing gear. A combination of field observations and laboratory experiments will be necessary to parameterize stock assessment models that are improved to accommodate variation in fish behaviour. Otherwise, survey data could reveal more about variation in behaviour than abundance trends.     

Disentangling the role of connectivity,environmental filtering,and spatial structure on metacommunity dynamics

Dispersal is a key process in metacommunity dynamics, allowing the maintenance of diversity in complex community networks. Geographic distance is usually used as a surrogate for connectivity implying that communities that are closely located are considered more prone to exchange individuals than distant communities. However, in some natural systems, organisms may be subjected to directional dispersal (air or water flows, particular landscape configuration), possibly leading close communities to be isolated from each other and distant communities to be connected. Using geographic distance as a proxy for realised connectivity may then yield misleading results regarding the role of dispersal in structuring communities in such systems. Here, we quantified the relative importance of flow connectivity, geographic distance, and environmental gradients to explain polychaete metacommunity structure along the coasts of the Gulf of Lions (northwest Mediterranean Sea). Flow connectivity was estimated by Lagrangian particle dispersal simulations. Our results revealed that this metacommunity is strongly structured by the environment at large spatial scales, and that both flow connectivity and geographic distance play an important role within homogeneous environments at smaller spatial scales. We thus strongly advocate for a wider use of connectivity measures, in addition to geographic distance, to study spatial patterns of biological diversity (e.g. distance decay) and to infer the processes behind these patterns at different spatial scales. Synthesis Everything is connected, but connections are seldom accurately quantified. Biological communities are often studied separately, using observations, experiments and models to unravel local dynamics of organisms interacting with each other. However, regional processes such as dispersal through ocean and air circulation, likely to connect distant communities and influence their local dynamics, are not always accounted for, or, at best, used as an homogeneous and distance‐related factor. Ocean models have being extensively developed and validated during the past decades with the increasing availability of accurate meteorological data. Using such model outputs, precise quantifi cation of exchange rates of organisms between communities was performed in a marine Mediterranean coastal area. Jointly with local environmental and biological data, these results were used to quantify the effects of realistic connectivity on local and regional polychaete community structure, and revealed that the environmental gradient, geographic distance, and connectivity were responsible for community structure at different spatial scales.