Exploring density-dependent relationships in demographic parameters in populations of birds at a large spatial scale

The importance of density-dependent processes in natural populations is widely accepted, but the issue of the shape of density-dependent relationships (such as influenced by vagueness, or time-delay) remains unresolved. We explored the density-dependent relationships in demographic parameters for 12 species of birds in Britain using large-scale, long-term data sets. We predicted that a negative relation between density and demographic parameters should be observed for the stable species, whereas the decreasing or increasing species should display a positive relation if the environment changes progressively through time bringing about a continuous change in density dependence. Our prediction was verified for nine species out of 12; however, we observed, for the three remaining species, a significant decrease of survival rates through time that seems to be involved in a long-term population decline. In all cases where a density-dependent relation was found, we observed an important variance around the relation. In one case, we showed that this variance increased significantly with density. We found evidence for time-delayed effects of density dependence both for survival and breeding performance. In two species, our results suggest the existence of complex interactions (compensatory mechanisms) between survival and breeding performance or between the different components of breeding performance.     

Responses to environmental stressors in an estuarine fish: Interacting stressors and the impacts of local adaptation

Common killifish, Fundulus heteroclitus, are found in marshes and estuaries along the Atlantic coast of North America from Newfoundland to Florida. Although these habitats are highly productive, they are also characterized by variation in a number of abiotic stressors, including temperature, salinity, oxygen, and anthropogenic toxicants, which vary substantially in both space and time. In order to survive in these habitats, killifish must be able to cope with these stressors, both individually and in combination. There is substantial evidence to suggest that populations of F. heteroclitus have undergone local adaptation to multiple abiotic stressors, including temperature, salinity, and toxicants, but most studies have examined the effects of single stressors in isolation. Here I review some of the studies on local adaptation in F. heteroclitus, focusing on the molecular basis of local adaptation to abiotic stressors, and the acute responses to these stressors both singly and in combination. This work demonstrates that there are substantial interactions between the responses to both natural and anthropogenic stressors at the cellular level.     

Population-level ecological effect assessment: estimating the effect of toxic chemicals on density-dependent populations

We examined the relationship between individual-level and population-level effects of toxic chemicals, employing the equilibrium population size as an index of population-level effects. We first analyzed two-stage matrix models considering four life-history types and four density-dependent models, and then we analyzed ecotoxicological and life-history data of the fathead minnow (Pimephales promelas) and brook trout (Salvelinus fontinalis) as real examples. Our elasticity analysis showed that toxic impacts on density-dependent populations depended largely on the differences in density-dependence and in life histories of the organisms. In particular, the importance of adult survivability was considerably increased in iteroparous organisms with density-dependent juvenile survivability or fertility. Our results also suggested that population-level effects, as indicated by the percentage reduction in equilibrium population size, were often greater than the percentage reductions in vital rates of individuals. Our analysis indicates that assessing population-level risk and developing a risk-reduction strategy without considering density-dependence can be risky.     

Density-dependent demographic variation determines extinction rate of experimental populations

Understanding population extinctions is a chief goal of ecological theory. While stochastic theories of population growth are commonly used to forecast extinction, models used for prediction have not been adequately tested with experimental data. In a previously published experiment, variation in available food was experimentally manipulated in 281 laboratory populations of Daphnia magna to test hypothesized effects of environmental variation on population persistence. Here, half of those data were used to select and fit a stochastic model of population growth to predict extinctions of populations in the other half. When density-dependent demographic stochasticity was detected and incorporated in simple stochastic models, rates of population extinction were accurately predicted or only slightly biased. However, when density-dependent demographic stochasticity was not accounted for, as is usual when forecasting extinction of threatened and endangered species, predicted extinction rates were severely biased. Thus, an experimental demonstration shows that reliable estimates of extinction risk may be obtained for populations in variable environments if high-quality data are available for model selection and if density-dependent demographic stochasticity is accounted for. These results suggest that further consideration of density-dependent demographic stochasticity is required if predicted extinction rates are to be relied upon for conservation planning.     

Multiple stressor effects on coral reef ecosystems

Global climate change has profound implications on species distributions and ecosystem functioning. In the coastal zone, ecological responses may be driven by various biogeochemical and physical environmental factors. Synergistic interactions can occur when the combined effects of stressors exceed their individual effects. The Red Sea, characterized by strong gradients in temperature, salinity, and nutrients along the latitudinal axis provides a unique opportunity to study ecological responses over a range of these environmental variables. Using multiple linear regression models integrating in situ, satellite and oceanographic data, we investigated the response of coral reef taxa to local stressors and recent climate variability. Taxa and functional groups responded to a combination of climate (temperature, salinity, air‐sea heat fluxes, irradiance, wind speed), fishing pressure and biogeochemical (chlorophyll a and nutrients ‐ phosphate, nitrate, nitrite) factors. The regression model for each species showed interactive effects of climate, fishing pressure and nutrient variables. The nature of the effects (antagonistic or synergistic) was dependent on the species and stressor pair. Variables consistently associated with the highest number of synergistic interactions included heat flux terms, temperature, and wind speed followed by fishing pressure. Hard corals and coralline algae abundance were sensitive to changing environmental conditions where synergistic interactions decreased their percentage cover. These synergistic interactions suggest that the negative effects of fishing pressure and eutrophication may exacerbate the impact of climate change on corals. A high number of interactions were also recorded for algae, however for this group, synergistic interactions increased algal abundance. This study is unique in applying regression analysis to multiple environmental variables simultaneously to understand stressor interactions in the field. The observed responses have important implications for understanding climate change impacts on marine ecosystems and whether managing local stressors, such as nutrient enrichment and fishing activities, may help mitigate global drivers of change.     

Behavioral interactions under multiple stressors: temperature and salinity mediate aggression between an invasive and a native fish

Interspecific aggression is a critical determinant of the success and competitive superiority of many invasive over native species. While single abiotic stressors can alter aggression levels, the manner in which multiple stressors may alter the strength and outcome of interspecific interactions and hence the invasion potential of a species is still poorly understood, even though multiple stressors are prevalent in many ecosystems. Furthermore, multiple stressors may interact to produce synergistic or antagonistic effects on individual level behaviors, thereby modulating invasive-native species interactions in unexpected ways. Here we examined the effect of two key abiotic stressors in freshwater ecosystems-temperature and salinity-on interspecific aggression between the invasive eastern mosquito fish (Gambusia holbrooki) and juveniles of the native Australian bass (Macquaria novemaculeata). Under controlled laboratory conditions, individuals were exposed to low or high salinity levels (15 and 35‰), and low or high temperatures (21 and 28 °C), and the frequency of interspecific aggressive behaviors was scored. The effect of temperature and salinity on interspecific aggression was antagonistic for both M. novemaculeata and G. holbrooki. While elevated temperature promoted aggression, elevated salinity partially or entirely negated this effect. Moreover, regardless of temperature, M. novemaculeata was more aggressive than G. holbrooki under elevated salinity. In addition to this, the native displayed more aggression to smaller than larger heterospecifics when exposed to elevated salinity alone, while G. holbrooki showed no such preference. These results highlight the importance of considering the interplay between multiple abiotic stressors and behavioral interactions between invasive and native species, combined with the modulating effect of species-specific and size based responses to those stressors.     

The devil is in the detail: Nonadditive and context‐dependent plant population responses to increasing temperature and precipitation

In climate change ecology, simplistic research approaches may yield unrealistically simplistic answers to often more complicated problems. In particular, the complexity of vegetation responses to global climate change begs a better understanding of the impacts of concomitant changes in several climatic drivers, how these impacts vary across different climatic contexts, and of the demographic processes underlying population changes. Using a replicated, factorial, whole‐community transplant experiment, we investigated regional variation in demographic responses of plant populations to increased temperature and/or precipitation. Across four perennial forb species and 12 sites, we found strong responses to both temperature and precipitation change. Changes in population growth rates were mainly due to changes in survival and clonality. In three of the four study species, the combined increase in temperature and precipitation reflected nonadditive, antagonistic interactions of the single climatic changes for population growth rate and survival, while the interactions were additive and synergistic for clonality. This disparity affects the persistence of genotypes, but also suggests that the mechanisms behind the responses of the vital rates differ. In addition, survival effects varied systematically with climatic context, with wetter and warmer + wetter transplants showing less positive or more negative responses at warmer sites. The detailed demographic approach yields important mechanistic insights into how concomitant changes in temperature and precipitation affect plants, which makes our results generalizable beyond the four study species. Our comprehensive study design illustrates the power of replicated field experiments in disentangling the complex relationships and patterns that govern climate change impacts across real‐world species and landscapes.     

Synergism of Metabolite Action in Plant Responses to Stresses

A diversity of unfavorable environmental factors determines complex interactions between phytohormones and signal elements and also the formation and combined action of metabolites in plants. This permits the plant to overcome stresses and to realize its potential capacities. Synergism between plant chemical constituents during ontogeny was revealed in the case of their responses to various abiotic stressors, such as pathogens or pests, and also during competition between plants. Jasmonic, salicylic, and abscisic acids, ethylene, hydrogen peroxide, NO, antioxidants, defensive proteins, and enzymes can manifest synergistic plant responses to unfavorable factors.__________Translated from Fiziologiya Rastenii, Vol. 52, No. 4, 2005, pp. 614–621.Original Russian Text Copyright © 2005 by Ryabushkina.     

Biotic interactions modify multiple‐stressor effects on juvenile brown trout in an experimental stream food web

Agricultural land use results in multiple stressors affecting stream ecosystems. Flow reduction due to water abstraction, elevated levels of nutrients and chemical contaminants are common agricultural stressors worldwide. Concurrently, stream ecosystems are also increasingly affected by climate change. Interactions among multiple co‐occurring stressors result in biological responses that cannot be predicted from single‐stressor effects (i.e. synergisms and antagonisms). At the ecosystem level, multiple‐stressor effects can be further modified by biotic interactions (e.g. trophic interactions). We conducted a field experiment using 128 flow‐through stream mesocosms to examine the individual and combined effects of water abstraction, nutrient enrichment and elevated levels of the nitrification inhibitor dicyandiamide (DCD) on survival, condition and gut content of juvenile brown trout and on benthic abundance of their invertebrate prey. Flow velocity reduction decreased fish survival (?12% compared to controls) and condition (?8% compared to initial condition), whereas effects of nutrient and DCD additions and interactions among these stressors were not significant. Negative effects of flow velocity reduction on fish survival and condition were consistent with effects on fish gut content (?25% compared to controls) and abundance of dominant invertebrate prey (?30% compared to controls), suggesting a negative metabolic balance driving fish mortality and condition decline, which was confirmed by structural equation modelling. Fish mortality under reduced flow velocity increased as maximal daily water temperatures approached the upper limit of their tolerance range, reflecting synergistic interactions between these stressors. Our study highlights the importance of indirect stressor effects such as those transferred through trophic interactions, which need to be considered when assessing and managing fish populations and stream food webs in multiple‐stressor situations. However, in real streams, compensatory mechanisms and behavioural responses, as well as seasonal and spatial variation, may alter the intensity of stressor effects and the sensitivity of trout populations.     

Emergent interactions influence functional traits and success of dune building ecosystem engineers

Aims Dune building processes are affected by interactions between the growth of ecosystem engineering dune grasses and environmental factors associated with disturbance such as sand burial and sea spray. Research investigating how species interactions influence dune community structure and functional trait responses in high abiotic stress environments is minimal. We investigated how species interactions influence the functional trait responses of three dominant dune grasses to common abiotic stressors.Methods We performed a multi-factorial greenhouse experiment by planting three common dune grasses (Ammophila breviligulata Fern., Uniola paniculata L. and Spartina patens Muhl.) in different interspecific combinations, using sand burial and sea spray as abiotic stressors. Sand burial was applied once at the beginning of the study. Sea spray was applied three times per week using a calibrated spray bottle. Morphological functional trait measurements (leaf elongation, maximum root length, aboveground biomass and belowground biomass) were collected at the end of the study. The experiment continued from May 2015 to August 2015.Important findings Species interactions between A. breviligulata and U. paniculata negatively affected dune building function traits of A. breviligulata, indicating that interactions with U. paniculata could alter dune community structure. Furthermore, A. breviligulata had a negative interaction with S. patens, which decreased S. patens functional trait responses to abiotic stress. When all species occurred together, the interactions among species brought about coexistence of all three species. Our data suggest that species interactions can change traditional functional trait responses of dominant species to abiotic stress.     

Positive interactions can increase size inequality in plant populations

1.  Large variation in the size of individuals is a ubiquitous feature of natural plant populations. While the role of competition in generating this variation has been studied extensively, the potential effects of positive interactions among plants, which are common in high-stress environments, have not been investigated.
2.  Using an individual-based 'zone-of-influence' model, we investigate the effects of competition, abiotic stress and facilitation on size inequality in plant monocultures. In the model, stress reduces the growth rate of plants, and facilitation ameliorates the effects of stress. Both facilitation and competition occur in overlapping zones of influence. We tested some of the model's predictions with a field experiment using the clonal grass Elymus nutans in an alpine meadow.
3.  Facilitation increased the size inequality of model populations when there was no density-dependent mortality. This effect decreased with density as competition overwhelmed facilitation. The lowest size inequality was found at intermediate densities both with the model and in the field.
4.  When density-dependent mortality was included in the model, stress delayed its onset and reduced its rate by reducing growth rates, so the number of survivors at any point in time was higher under harsh than under more benign conditions. Facilitation increased size inequality during self-thinning.
5.   Synthesis . Our results demonstrate that facilitation interacts with abiotic stress and competition to influence the degree of size inequality in plant populations. Facilitation increased size inequality at low to intermediate densities and during self-thinning.     

An improved null model for assessing the net effects of multiple stressors on communities

Ecological stressors (i.e., environmental factors outside their normal range of variation) can mediate each other through their interactions, leading to unexpected combined effects on communities. Determining whether the net effect of stressors is ecologically surprising requires comparing their cumulative impact to a null model that represents the linear combination of their individual effects (i.e., an additive expectation). However, we show that standard additive and multiplicative null models that base their predictions on the effects of single stressors on community properties (e.g., species richness or biomass) do not provide this linear expectation, leading to incorrect interpretations of antagonistic and synergistic responses by communities. We present an alternative, the compositional null model, which instead bases its predictions on the effects of stressors on individual species, and then aggregates them to the community level. Simulations demonstrate the improved ability of the compositional null model to accurately provide a linear expectation of the net effect of stressors. We simulate the response of communities to paired stressors that affect species in a purely additive fashion and compare the relative abilities of the compositional null model and two standard community property null models (additive and multiplicative) to predict these linear changes in species richness and community biomass across different combinations (both positive, negative, or opposite) and intensities of stressors. The compositional model predicts the linear effects of multiple stressors under almost all scenarios, allowing for proper classification of net effects, whereas the standard null models do not. Our findings suggest that current estimates of the prevalence of ecological surprises on communities based on community property null models are unreliable, and should be improved by integrating the responses of individual species to the community level as does our compositional null model.     

The relative roles of density and climatic variation on population dynamics and fecundity rates in three contrasting ungulate species

The relative influences of density-dependent and -independent processes on vital rates and population dynamics have been debated in ecology for over half a century, yet it is only recently that both processes have been shown to operate within the same population. However, generalizations on the role of each process across species are rare. Using a process-orientated generalized linear modelling approach we show that variations in fecundity rates in populations of three species of ungulates with contrasting life histories are associated with density and winter weather in a remarkably similar manner. However, there are differences and we speculate that they are a result of differences in size between the species. Much previous research exploring the association between vital rates, population dynamics and density-dependent and -independent processes has used pattern-orientated approaches to decompose time-series into contributions from density-dependent and -independent processes. Results from these analyses are sometimes used to infer associations between vital rates, density and climatic variables. We compare results from pattern-orientated analyses of time-series with process-orientated analyses and report that the two approaches give different results. The approach of analysing relationships between vital rates, density and climatic variables may detect important processes influencing population dynamics that time-series methodologies may overlook.     

Range position and climate sensitivity: The structure of among‐population demographic responses to climatic variation

Species’ distributions will respond to climate change based on the relationship between local demographic processes and climate and how this relationship varies based on range position. A rarely tested demographic prediction is that populations at the extremes of a species’ climate envelope (e.g., populations in areas with the highest mean annual temperature) will be most sensitive to local shifts in climate (i.e., warming). We tested this prediction using a dynamic species distribution model linking demographic rates to variation in temperature and precipitation for wood frogs (Lithobates sylvaticus) in North America. Using long‐term monitoring data from 746 populations in 27 study areas, we determined how climatic variation affected population growth rates and how these relationships varied with respect to long‐term climate. Some models supported the predicted pattern, with negative effects of extreme summer temperatures in hotter areas and positive effects on recruitment for summer water availability in drier areas. We also found evidence of interacting temperature and precipitation influencing population size, such as extreme heat having less of a negative effect in wetter areas. Other results were contrary to predictions, such as positive effects of summer water availability in wetter parts of the range and positive responses to winter warming especially in milder areas. In general, we found wood frogs were more sensitive to changes in temperature or temperature interacting with precipitation than to changes in precipitation alone. Our results suggest that sensitivity to changes in climate cannot be predicted simply by knowing locations within the species’ climate envelope. Many climate processes did not affect population growth rates in the predicted direction based on range position. Processes such as species‐interactions, local adaptation, and interactions with the physical landscape likely affect the responses we observed. Our work highlights the need to measure demographic responses to changing climate.     

Effects of positive interactions, size symmetry of competition and abiotic stress on self-thinning in simulated plant populations

Background and Aims

Competition drives self-thinning (density-dependent mortality) in crowded plant populations. Facilitative interactions have been shown to affect many processes in plant populations and communities, but their effects on self-thinning trajectories have not been investigated.

Methods

Using an individual-based ‘zone-of-influence’ model, we studied the potential effects of the size symmetry of competition, abiotic stress and facilitation on self-thinning trajectories in plant monocultures. In the model, abiotic stress reduced the growth of all individuals and facilitation ameliorated the effects of stress on interacting individuals.

Key Results

Abiotic stress made the log biomass – log density relationship during self-thinning steeper, but this effect was reduced by positive interactions among individuals. Size-asymmetric competition also influenced the self-thinning slope.

Conclusions

Although competition drives self-thinning, its course can be affected by abiotic stress, facilitation and competitive symmetry.     

Identifying the interacting roles of stressors in driving the global loss of canopy‐forming to mat‐forming algae in marine ecosystems

Identifying the type and strength of interactions between local anthropogenic and other stressors can help to set achievable management targets for degraded marine ecosystems and support their resilience by identifying local actions. We undertook a meta‐analysis, using data from 118 studies to test the hypothesis that ongoing global declines in the dominant habitat along temperate rocky coastlines, forests of canopy‐forming algae and/or their replacement by mat‐forming algae are driven by the nonadditive interactions between local anthropogenic stressors that can be addressed through management actions (fishing, heavy metal pollution, nutrient enrichment and high sediment loads) and other stressors (presence of competitors or grazers, removal of canopy algae, limiting or excessive light, low or high salinity, increasing temperature, high wave exposure and high UV or CO₂), not as easily amenable to management actions. In general, the cumulative effects of local anthropogenic and other stressors had negative effects on the growth and survival of canopy‐forming algae. Conversely, the growth or survival of mat‐forming algae was either unaffected or significantly enhanced by the same pairs of stressors. Contrary to our predictions, the majority of interactions between stressors were additive. There were however synergistic interactions between nutrient enrichment and heavy metals, the presence of competitors, low light and increasing temperature, leading to amplified negative effects on canopy‐forming algae. There were also synergistic interactions between nutrient enrichment and increasing CO₂ and temperature leading to amplified positive effects on mat‐forming algae. Our review of the current literature shows that management of nutrient levels, rather than fishing, heavy metal pollution or high sediment loads, would provide the greatest opportunity for preventing the shift from canopy to mat‐forming algae, particularly in enclosed bays or estuaries because of the higher prevalence of synergistic interactions between nutrient enrichment with other local and global stressors, and as such it should be prioritized.     

Modeling as a tool to manage ecosystems under multiple stresses: an application to Lake Ontario

Present-day ecosystem management involves understanding of the synergistic effect of multiple stressors on multiple and frequently nebulous management end-points. An example is the simultaneous management of nutrient load reductions and salmon stocking in Lake Ontario. In this study, a simple whole-lake annual time scale model was developed to assess the relationship between these two stressors and various ecosystem responses. The model was used to explore the utility of some possible management end-points for ecosystem health. In historical simulations, production per stocked fish and salmon survival appeared to be good indicators, while nutrient recycling rate and average ecosystem-wide food limitation were found to be fairly unresponsive to the two stressors. The model was further used to predict long term averages of salmon biomass and selected health indicators at various sustained loading and stocking rates. Salmon biomass increased with stocking rate at all stocking rates examined, but the rate of increase declined somewhat at high stocking rates. The response of salmon biomass to nutrient loading appeared to be approximately sigmoidal i.e. there was a nutrient threshold below which fish biomass could not be sustained and another nutrient threshold above which salmon biomass either remained constant or even decreased. The response to either stressor was found to be modified by the value of the other stressor, illustrating the importance of ecosystem-level models for aquatic ecosystem management.     

Describing and understanding behavioral responses to multiple stressors and multiple stimuli

Understanding the effects of environmental change on natural ecosystems is a major challenge, particularly when multiple stressors interact to produce unexpected “ecological surprises” in the form of complex, nonadditive effects that can amplify or reduce their individual effects. Animals often respond behaviorally to environmental change, and multiple stressors can have both population‐level and community‐level effects. However, the individual, not combined, effects of stressors on animal behavior are commonly studied. There is a need to understand how animals respond to the more complex combinations of stressors that occur in nature, which requires a systematic and rigorous approach to quantify the various potential behavioral responses to the independent and interactive effects of stressors. We illustrate a robust, systematic approach for understanding behavioral responses to multiple stressors based on integrating schemes used to quantitatively classify interactions in multiple‐stressor research and to qualitatively view interactions between multiple stimuli in behavioral experiments. We introduce and unify the two frameworks, highlighting their conceptual and methodological similarities, and use four case studies to demonstrate how this unification could improve our interpretation of interactions in behavioral experiments and guide efforts to manage the effects of multiple stressors. Our unified approach: (1) provides behavioral ecologists with a more rigorous and systematic way to quantify how animals respond to interactions between multiple stimuli, an important theoretical advance, (2) helps us better understand how animals behave when they encounter multiple, potentially interacting stressors, and (3) contributes more generally to the understanding of “ecological surprises” in multiple stressors research.     

Lotka's Game in Predator–Prey Theory: Linking Populations to Individuals

This paper further examines an individual-based model of a spatially distributed predator–prey population that demonstrates strong spatial structuring in contrast with predictions from its representative analytic formulation. Examination of a small, localized population reveals that extinctions due to demographic stochasticity dominate the dynamics. Local extinction dynamics produce wave pulses and the interactions of these wave pulses constitute global dynamics. The results motivate a population-level cell-based model with each cell representing a local population and parameterized by local extinction probabilities, rather than individual-based interaction rates. A detailed comparison of spatiotemporal plots from the two modelling frameworks shows that the population-level model captures the broad range of dynamics exhibited by the individual-based model. The agreement between these two complementary theoretical frameworks, one formulated at the level of individuals, the other at the level of populations, provides a mechanistic understanding of the dynamics.     

Global switches and fine-tuning-ABA modulates plant pathogen defense

Plants are obliged to defend themselves against a wide range of biotic and abiotic stresses. Complex regulatory signaling networks mount an appropriate defense response depending on the type of stress that is perceived. In response to abiotic stresses such as drought, cold, and salinity, the function of abscisic acid (ABA) is well documented: elevation of plant ABA levels and activation of ABA-responsive signaling result in regulation of stomatal aperture and expression of stress-responsive genes. In response to pathogens, the role of ABA is more obscure and is a research topic that has long been overlooked. This article aims to evaluate and review the reported modes of ABA action on pathogen defense and highlight recent advances in deciphering the complex role of ABA in plant-pathogen interactions. The proposed mechanisms responsible for positive or negative effects of ABA on pathogen defense are discussed, as well as the regulation of ABA signaling and in planta ABA concentrations by beneficial and pathogenic microorganisms. In addition, the fast-growing number of reports that characterize antagonistic and synergistic interactions between abiotic and biotic stress responses point to ABA as an essential component in integrating and fine-tuning abiotic and biotic stress-response signaling networks.