Intraspecific Density Dependence Research Articles

Successful forest restoration using plantation at high deer density: How neighboring vegetation drives browsing pressure and tree growth

Browsing can be an environmental stress to forest ecosystems, where the composition and structure of plant communities depend on the balance between ungulate browsing and the ability of plants to tolerate and acclimate to this stressor. Tree planting is a management tool for restoring forest whose integrity has been compromised by the intensity and repetition of browsing. This tool may be insufficient when the stress leading to forest degradation is ongoing. Balsam fir (Abies balsamea (L.) Mill.) plantations are used on Anticosti Island (Quebec, Canada) to restore winter habitat for white-tailed deer (Odocoileus virginianus Zimmermann, 1780). To promote natural regeneration and growth of planted firs that are the staple winter forage of deer, enclosures were temporarily erected and deer densities reduced to decrease browsing pressure. The stress induced by heavy browsing at the time of fence removal and the density of competitors in regenerating cutblocks, however, could compromise fir growth and forest recruitment. Our objective was to identify factors influencing browsing intensity and balsam fir’s growth following fence removal. We aimed to identify factors influencing lateral browsing intensity and number of trunk reiterations on fir at the landscape scale, including distance and size of the surrounding forests, and at the local scale through intraspecific density-dependence and interspecific associative effects. We then sought to identify the stressors most influencing balsam fir growth among browsing variables and a neighborhood crowding index (NCI). We measured browsing intensity and growth of 114 balsam firs 8 years following planting and again 2 and 4 years after dismantling two management enclosures. At the landscape scale, our results showed that when balsam fir was more than 150 m from a forest edge, the probability of browsing was close to zero and consistently a low percentage of shelter area surrounding a focal fir led to a decrease in browsing pressure. At the local scale, intensity of lateral browsing increased with the NCI of white spruce (Picea glauca (Moench) Voss). Fir growth was negatively influenced by the additive effects of browsing and NCI. Finally, our results showed that 19.7% (95 %CI: 4.5%, 42.6%) of the effect of the surrounding vegetation on growth was mediated by browsing. The success of forest ecosystem restoration through planting depends on density of neighboring tree species that can generate environmental stress by influencing growth indirectly through browsing as well as directly through competition for resources.

The existence and strength of higher order interactions is sensitive to environmental context.

One strategy for understanding the dynamics of any complex system, such as a community of competing species, is to study the dynamics of parts of the system in isolation. Ecological communities can be decomposed into single species, and pairs of interacting species. This reductionist strategy assumes that whole-community dynamics are predictable and explainable from knowledge of the dynamics of single species and pairs of species. This assumption will be violated if higher order interactions (HOIs) are strong. Theory predicts that HOIs should be common. But it is difficult to detect HOIs, and to infer their long-term consequences for species coexistence, solely from short-term data. I conducted a protist microcosm experiment to test for HOIs among competing bacterivorous ciliates, and test the sensitivity of HOIs to environmental context. I grew three competing ciliate species in all possible combinations at each of two resource enrichment levels, and used the population dynamic data from the one- and two-species treatments to parameterize a competition model at each enrichment level. I then compared the predictions of the parameterized model to the dynamics of the whole community (three-species treatment). I found that the existence, and thus strength, of HOIs was environment dependent. I found a strong HOI at low enrichment, which enabled the persistence of a species that would otherwise have been competitively excluded. At high enrichment, three-species dynamics could be predicted from a parameterized model of one- and two-species dynamics, provided that the model accounted for nonlinear intraspecific density dependence. The results provide one of the first rigorous demonstrations of the long-term consequences of HOIs for species coexistence, and demonstrate the context dependence of HOIs. HOIs create difficult challenges for predicting and explaining species coexistence in nature.

Effects of density, species interactions, and environmental stochasticity on the dynamics of British bird communities

Our knowledge of the factors affecting species abundances is mainly based on time‐series analyses of a few well‐studied species at single or few localities, but we know little about whether results from such analyses can be extrapolated to the community level. We apply a joint species distribution model to long‐term time‐series data on British bird communities to examine the relative contribution of intra‐ and interspecific density dependence at different spatial scales, as well as the influence of environmental stochasticity, to spatiotemporal interspecific variation in abundance. Intraspecific density dependence has the major structuring effect on these bird communities. In addition, environmental fluctuations affect spatiotemporal differences in abundance. In contrast, species interactions had a minor impact on variation in abundance. Thus, important drivers of single‐species dynamics are also strongly affecting dynamics of communities in time and space.

Non‐stationary and interactive effects of climate and competition on pink salmon productivity

Pacific salmon (Oncorhynchus spp.) are exposed to increased environmental change and multiple human stressors. To anticipate future impacts of global change and to improve sustainable resource management, it is critical to understand how wild salmon populations respond to stressors associated with human‐caused changes such as climate warming and ocean acidification, as well as competition in the ocean, which is intensified by the large‐scale production and release of hatchery reared salmon. Pink salmon (O. gorbuscha) are a keystone species in the North Pacific Ocean and support highly valuable commercial fisheries. We investigated the joint effects of changes in ocean conditions and salmon abundances on the productivity of wild pink salmon. Our analysis focused on Prince William Sound in Alaska, because the region accounts for ~50% of the global production of hatchery pink salmon with local hatcheries releasing 600–700 million pink salmon fry annually. Using 60 years of data on wild pink salmon abundances, hatchery releases, and ecological conditions in the ocean, we find evidence that hatchery pink salmon releases negatively affect wild pink salmon productivity, likely through competition between wild and hatchery juveniles in nearshore marine habitats. We find no evidence for effects of ocean acidification on pink salmon productivity. However, a change in the leading mode of North Pacific climate in 1988–1989 weakened the temperature–productivity relationship and altered the strength of intraspecific density dependence. Therefore, our results suggest non‐stationary (i.e., time varying) and interactive effects of ocean climate and competition on pink salmon productivity. Our findings further highlight the need for salmon management to consider potential adverse effects of large‐scale hatchery production within the context of ocean change.

Quantifying the nature and strength of intraspecific density dependence in Arctic mosquitoes.

Processes that change with density are inherent in all populations, yet quantifying density dependence with empirical data remains a challenge. This is especially true for animals recruiting in patchy landscapes because heterogeneity in habitat quality in combination with habitat choice can obscure patterns expected from density dependence. Mosquitoes (Diptera: Culicidae) typically experience strong density dependence when larvae compete for food, however, effects vary across species and contexts. If populations experience intense intraspecific density-dependent mortality then overcompensation can occur, where the number of survivors declines at high densities producing complex endogenous dynamics. To seek generalizations about density dependence in a widespread species of Arctic mosquito, Aedes nigripes, we combined a laboratory experiment, field observations, and modeling approaches. We evaluated alternative formulations of discrete population models and compared best-performing models from our lab study to larval densities from ponds in western Greenland. Survivorship curves from the lab were the best fit by a Hassell model with compensating density dependence (equivalent to a Beverton-Holt model) where peak recruitment ranged from 8 to 80 mosquitoes per liter depending on resource supply. In contrast, our field data did not show a signal of strong density dependence, suggesting that other processes such as predation may lower realized densities in nature, and that expected patterns may be obscured because larval abundance covaries with resources (cryptic density dependence). Our study emphasizes the importance of covariation between the environment, habitat choice, and density dependence in understanding population dynamics across landscapes, and demonstrates the value of pairing lab and field studies.

Multispecies integrated population model reveals bottom‐up dynamics in a seabird predator–prey system

AbstractAssessing the effects of climate and interspecific relationships on communities is challenging because of the complex interplay between species population dynamics, their interactions, and the need to integrate information across several biological levels (individuals, populations, communities). Usually used to quantify single‐species demography, integrated population models (IPMs) have recently been extended to communities. These models allow fitting multispecies matrix models to data from multiple sources while simultaneously accounting for uncertainty in each data source. We used multispecies IPMs accommodating climatic variables to quantify the relative contribution of climate vs. interspecific interactions on demographic parameters, such as survival and breeding success, in the dynamics of a predator–prey system. We considered a stage‐structured predator–prey system combining 22 yr of capture–recapture data and population counts of two seabirds, the Brown Skua (Catharacta lönnbergi) and its main prey the Blue Petrel (Halobaena caerulea), both breeding on the Kerguelen Islands in the Southern Ocean. Our results showed that climate and predator–prey interactions drive the demography of skuas and petrels in different ways. The breeding success of skuas appeared to be largely driven by the number of petrels and to a lesser extent by intraspecific density dependence. In contrast, there was no evidence of predation effects on the demographic parameters of petrels, which were affected by oceanographic factors. We conclude that bottom‐up mechanisms are the main drivers of this skua–petrel system.

Neural ordinary differential equations for ecological and evolutionary time‐series analysis

Abstract Inferring the functional shape of ecological and evolutionary processes from time‐series data can be challenging because processes are often not describable with simple equations. The dynamical coupling between variables in time series further complicates the identification of equations through model selection as the inference of a given process is contingent on the accurate depiction of all other processes. We present a novel method, neural ordinary differential equations (NODEs), for learning ecological and evolutionary processes from time‐series data by modelling dynamical systems as ordinary differential equations and dynamical functions with artificial neural networks (ANNs). Upon successful training, the ANNs converge to functional shapes that best describe the biological processes underlying the dynamics observed, in a way that is robust to mathematical misspecifications of the dynamical model. We demonstrate NODEs in a population dynamic context and show how they can be used to infer ecological interactions, dynamical causation and equilibrium points. We tested NODEs by analysing well‐understood hare and lynx time‐series data, which revealed that prey–predator oscillations were mainly driven by the interspecific interaction, as well as intraspecific densitydependence, and characterised by a single equilibrium point at the centre of the oscillation. Our approach is applicable to any system that can be modelled with differential equations, and particularly suitable for linking ecological, evolutionary and environmental dynamics where parametric approaches are too challenging to implement, opening new avenues for theoretical and empirical investigations.

Temporal patterns and ecosystem correlates of chum salmon (Oncorhynchus keta) migration phenology in the Pacific Northwest

Understanding and quantifying migration phenology of commercially harvested Pacific salmon (Oncorhynchus spp.) is a cornerstone for managing sustainable populations. Here, we use a multidecadal data time series together with a hypothesis-driven framework to evaluate migration phenology in adult fall and winter ecotype chum salmon (O. keta) in a poorly studied but highly managed system — the South Puget Sound (SPS) of Washington State, USA. Using generalized additive mixed models that accounted for temporal autoregressive dynamics, we examined the effect of commercial harvest, climate variation, intraspecific density dependence, and predator buffering on migration timing and run duration. SPS chum salmon are migrating earlier over time, especially the winter ecotype that showed the strongest temporal shift from historical timing. Migration timing shifts were closely associated with regional marine climate regimes, local scale freshwater availability, and increasing pinniped abundance. We conclude that there is potential for the winter ecotype migration converging with that of the fall ecotype and that directional change in migration phenology may be driven by a unique combination of ecosystem factors.

Intra‐ and interspecific density dependence mediates weather effects on the population dynamics of a plant–insect herbivore system

Temperature and precipitation are two major factors determining arthropod population densities, but the effects from these climate variables are seldom evaluated in the same study system and in combination with inter‐ and intraspecific density dependence. In this study, I used a 19 year time series on plant variables (shoot height and flowering incidence) and insect density in order to understand direct and indirect effects of climatic fluctuations on insect population densities. The study system includes two closely related leaf beetle species (Galerucella spp.) and a flower feeding weevil Nanophyes marmoratus attacking the plant purple loosestrife Lythrum salicaria. Results suggest that both intraspecific density dependence and weather variables affected Galerucella population densities, with interactive effects of rain and temperature on insect densities that depended on the timing relative to insect life cycles. In spring, high temperatures increased Galerucella densities only when combined with high rain, as low rain implies a high drought risk. Low temperatures are only beneficial if combined with little rain, as high rain cause chilly and wet conditions that are bad for insects. In summer, interactive effects of rain and temperature are different because high temperatures and little rain cause drought that induce wilting in plants, thus reducing food availability for the leaf feeding larvae. In contrast, the density of the flower feeding weevil was less affected by temperature and precipitation directly, and more indirectly interspecific density dependent effects through reduced resource availability caused by previous Galerucella damage.

Stony coral populations are more sensitive to changes in vital rates in disturbed environments

Reef-building corals, like many long-lived organisms, experience environmental change as a combination of separate but concurrent processes, some of which are gradual yet long-lasting, while others are more acute but short-lived. For corals, some chronic environmental stressors, such as rising temperature and ocean acidification, are thought to induce gradual changes in colonies' vital rates. Meanwhile, other environmental changes, such as the intensification of tropical cyclones, change the disturbance regime that corals experience. Here, we use a physiologically structured population model to explore how chronic environmental stressors that impact the vital rates of individual coral colonies interact with the intensity and magnitude of disturbance to affect coral population dynamics and cover. We find that, when disturbances are relatively benign, intraspecific density dependence driven by space competition partially buffers coral populations against gradual changes in vital rates. However, the impact of chronic stressors is amplified in more highly disturbed environments, because disturbance weakens the buffering effect of space competition. We also show that coral cover is more sensitive to changes in colony growth and mortality than to external recruitment, at least in open populations, and that space competition and size structure mediate the extent and pace of coral population recovery following a large-scale mortality event. Understanding the complex interplay among chronic environmental stressors, mass-mortality events, and population size structure sharpens our ability to manage and to restore coral-reef ecosystems in an increasingly disturbed future.

Density‐dependent egg production in chub mackerel in the Kuroshio Current system

AbstractA paradigm of proportionality between spawning stock biomass (SSB) and total egg production (TEP) has been largely untested at multidecadal scales mainly because of difficulty in estimating annual TEP. Recently, this paradigm was directly tested for sardine (Sardinops melanostictus) and anchovy (Engraulis japonicus) at a multidecadal scale to reveal that SSB–TEP proportionality was partially distorted by intraspecific (sardine) and interspecific (anchovy) density dependence in total egg production per spawner individual (TEPPS) or unit weight (TEPPSW). In the present study, we demonstrate intraspecific density dependence in TEPPS/TEPPSW for chub mackerel (Scomber japonicus) in the Kuroshio Current system, using a proxy for TEPPS/TEPPSW, calculated from snapshot abundance data based on fishery‐independent egg surveys in combination with fishery‐dependent stock assessment data, at a multidecadal scale (38 years). TEPPS/TEPPSW exponentially declined with SSB, indicating a strong intraspecific density dependence in TEPPS/TEPPSW in chub mackerel. The observed phenomenon for chub mackerel was similar to that for sardine. Hence, intraspecific density dependence in TEPPS/TEPPSW may be a phenomenon that is generally applicable for species with a high maximum biomass and large population fluctuations. Lastly, we recommend the application of a TEP‐based framework to studies on recruitment mechanisms of fish.

Variation in Reproductive Investment among Ohio Reservoir Largemouth Bass Populations

AbstractReproductive investment, expressed in terms of fecundity and egg quality produced by individual female fish, varies among populations for many species, reflecting plasticity in life history strategies related to intraspecific density dependence or environmental variability. Fisheries scientists and managers have long sought to understand differences in Largemouth Bass Micropterus salmoides population dynamics, yet basic life history traits, such as variation in reproductive investment, remain largely unknown. We determined the fecundity–weight and ovary energy density–weight relationships (where fecundity indexes egg quantity and ovary energy density indexes egg quality) for Largemouth Bass populations from 19 Ohio reservoirs and examined factors that were correlated with population‐specific differences. We found that both relationships varied among reservoir populations, with some interannual variation in reservoirs sampled in multiple years. We then related fecundity and ovary energy density to (1) population characteristics, including population density (as indexed by standard shoreline boat electrofishing CPUE, fish/h) and growth (as indexed by the growth coefficient k from the von Bertalanffy growth function); and (2) reservoir characteristics, including reservoir size (as surface area) and productivity (as Secchi transparency depth). We found that populations with greater fecundity also exhibited greater ovary energy density, indicating that the quantity and quality of reproductive output were greater in some populations than in others. Populations with high reproductive investment were associated with low Largemouth Bass density, fast growth, large reservoir size, and high productivity (i.e., low Secchi transparency). The among‐population variation in reproductive investment provides evidence of density‐dependent control of fecundity and egg quality.

Rosa multiflora’s performance under water stress: the role of positive and negative density-dependent intraspecific interactions

It is well known that biotic interactions may significantly alter plant population and community dynamics in natural ecosystems. Multiple studies have reported that density-dependent positive interactions can modify the effect of extreme stress on plant performance (in line with predictions by the stress gradient hypothesis, SGH). However, the performance of invasive species and the role of intraspecific density dependence, either negative or positive, in successful invasions are not well understood. In the present study, we tested if monocultures of Rosa multiflora, a highly invasive shrub, experience a group advantage under extremely low water availability. Using a manipulative greenhouse study, we tested the growth response of R. multiflora seedlings to a 4-level conspecific density treatment and a 4-level water availability gradient. Overall, our results provide preliminary evidence that biotic interactions under stressful conditions between R. multiflora individuals do not experience a group advantage, as predicted. Instead, our study indicates that seedling performance was strongly driven by negative density dependence, in which individuals grown at higher conspecific densities displayed lower plant performance, regardless of water availability. This demonstrates that R. multiflora experiences intense intraspecific competition even under a stressful, low resource environment. Thus, management efforts of R. multiflora need to ensure that re-sprouting of individuals after treatment is eliminated, as release from negative density dependence could lead the invader population to rebound.

Disentangling ecologically equivalent from neutral species: The mechanisms of population regulation matter.

The neutral theory of biodiversity explored the structure of a community of ecologically equivalent species. Such species are expected to display community drift dynamics analogous to neutral alleles undergoing genetic drift. While entire communities of species are not ecologically equivalent, recent field experiments have documented the existence of guilds of such neutral species embedded in real food webs. What demographic outcomes of the interactions within and between species in these guilds are expected to produce ecological drift versus coexistence remains unclear. To address this issue, and guide empirical testing, we consider models of a guild of ecologically equivalent competitors feeding on a single resource to explore when community drift should manifest. We show that community drift dynamics only emerge when the density-dependent effects of each species on itself are identical to its density-dependent effects on every other guild member. In contrast, if each guild member directly limits itself more than it limits the abundance of other guild members, all species in the guild are coexisting, even though they all are ecologically equivalent with respect to their interactions with species outside the guild (i.e. resources, predators, mutualists). Hence, considering only interspecific ecological differences generating density dependence, and not fully accounting for the preponderance of mechanisms causing intraspecific density dependence, will provide an incomplete picture for segregating between neutrality and coexistence. We also identify critical experiments necessary to disentangle guilds of ecologically equivalent species from those experiencing ecological drift, as well as provide an overview of ways of incorporating a mechanistic basis into studies of species coexistence and neutrality. Identifying these characteristics, and the mechanistic basis underlying community structure, is not merely an exercise in clarifying the semantics of coexistence and neutral theories, but rather reflects key differences that must exist among community members in order to determine how and why communities are structured.

Disentangling density‐dependent effects on egg production and survival from egg to recruitment in fish

AbstractUnderstanding of density‐dependent effects is key to achieving sustainable management of self‐regulating biological resources such as fish stocks. Traditionally, density‐dependent effects on population abundance in fish have been considered to occur from hatching to recruitment, based on the paradigm of proportionality between spawning stock biomass and total egg production. Here, we demonstrate how the existence of intraspecific and interspecific density dependence in egg production changes the current understanding of density‐dependent processes in the life history of fish, by disentangling density‐dependent effects on egg production and survival from egg to recruitment, using sardine (Sardinops melanostictus, Clupeidae) and anchovy (Engraulis japonicus, Engraulidae) as model species. For sardine, strong intraspecific density‐dependent effects occurred in egg production, but no density‐dependent effects occurred or if any they were weak enough to be masked by environmental factors from hatching to recruitment. In contrast, for anchovy, interspecific density‐dependent effects occurred in egg production. In the survival after hatching, anchovy experienced stronger intraspecific density‐dependent effects than currently recognized. This analysis could overturn the current understanding of density‐dependent effects in the life history, highlighting contrasts between the effects on individual quality and population abundance and between the model species. We propose to reconsider the basis of fisheries management and recruitment studies based on the revised understanding of density‐dependent effects in the life history of the respective species.

Two dimensions of demographic differentiation of species in a mountain grassland community: An experimental test

Abstract There is remarkable variation in life histories of coexisting plant species. These “alternative designs” for the given set of environmental conditions are likely to play a role in species niche differences and thus may underlie species coexistence, although there is no clear demonstration of it. Currently available data on within‐community differentiation concern primarily easy and measurable traits that are not fully informative of life history variation. Their relevance for functional differentiation of species and species coexistence is far from clear. Here, we examined differentiation of coexisting species in demographic parameters and how it determines species’ response to neighbours. We determined these parameters for a set of 21 co‐occurring species by fitting a process‐based model to a long‐term (30 years) data series of shoot counts. We examined the functional relevance of these parameters using species responses in a field removal experiment. We further asked with which functional traits they are correlated. Species were differentiated along two largely independent axes: (a) slow vs. fast, separating species according to instantaneous growth rate, competitive response and intraspecific density dependence; and (b) dispersal vs. local dynamics, which separated species with strong dispersal (by seeds or vegetative) from species that tended to stay in the occupied spot. While the slow‐fast axis was associated with commonly used leaf and seed traits, the dispersal axis was best predicted by lateral spreading distance. Each of these two axes predicted different components of species’ responses to neighbour competition: the slow‐fast axis was a good predictor of the short‐term response, whereas dispersal axis was a good predictor of the long‐term response. Synthesis. Demographic differentiation of coexisting species resembles to an important degree demographic differentiation known from large‐scale comparisons. This differentiation is functionally meaningful also at the fine scale; its role in species’ responses to competition implies it is involved in species niche differentiation and coexistence. While seed and leaf traits are important correlates of demographic differentiation, a hitherto underappreciated trait, viz. lateral spreading distance, is an important predictor of the dispersal axis at the fine scale and should be more widely used. A plain language summary is available for this article.

Coastal phytoplankton community dynamics and coexistence driven by intragroup density‐dependence, light and hydrodynamics

Phytoplanktonic communities maintain a high diversity in a seemingly homogeneous environment, competing for the same set of resources. Many theories have been proposed to explain this coexistence despite likely competition, such as contrasted responses to temporal environmental variation. However, theory has developed at a faster pace than empirical evaluation using field data, which requires inferring drivers of community dynamics from observational time series. Here, we combine autoregressive models with a data set spanning more than 20 years of phytoplankton counts every two weeks, together with nutrients and physical variables. By comparing models dominated by nutrients or physical variables (hydrodynamics and climate), we first explore which abiotic factors contribute more to phytoplankton growth and decline. We find that physical drivers – such as irradiance, wind, and salinity – explain some of the variability in abundances unexplained by biotic interactions. In contrast, responses to nutrients explain less of the phytoplankton variability. Concerning biotic drivers of community dynamics, multivariate autoregressive models reveal that competition between different groups (genera) has a much weaker effect on population growth rates than competition within a group. In fact, the few biotic interactions between genera that are detected are frequently positive. Hence, our system is unlikely to be best represented as a set of competitors whose differing responses to fluctuating environments allow coexistence, as in ‘paradox of the plankton’ models with a storage effect or a relative nonlinearity of competition. Coexistence is more likely to result from high intragroup density‐dependence. Competition between phytoplanktonic groups and nutrient limitation are often invoked as drivers of phytoplankton dynamics; our findings suggest instead that more attention should be given to the physical structure of the environment and natural enemies such as grazers and pathogens, that could explain the high intraspecific density dependence found here.

Density dependence in total egg production per spawner for marine fish

AbstractA paradigm of fisheries science holds that spawning stock biomass (SSB) is directly proportional to total egg production (TEP) of fish stocks. This “SSB–TEP proportionality” paradigm has been a basic premise underlying the spawner–recruitment models for fisheries management and numerous studies on recruitment mechanisms of fish. Studies on maternal effects on reproductive potential of a stock have progressed during the last few decades, leading to doubt concerning the paradigm. Nonetheless, a direct test of the paradigm at multidecadal scales has been difficult because of data limitations in the stock assessment systems worldwide. Here, we tested the paradigm for marine fish based on a novel combination of two independent 38‐year time series: fishery‐dependent stock assessment data and fishery‐independent egg survey data. Through this approach, we show that the SSB–TEP proportionality is distorted by density dependence in total egg production per spawner individual (TEPPS) or spawner unit weight (TEPPSW) at a multidecadal scale. The TEPPS/TEPPSW exponentially declined with biomass and thus was density‐dependent for Japanese sardine, a small pelagic species exhibiting a high level of population fluctuation, in the western North Pacific. By contrast, the TEPPS/TEPPSW was sardine‐density‐dependent for Japanese anchovy, another small pelagic species exhibiting a moderate level of population fluctuation well‐known for being out of phase with sardine. Our analysis revealed intraspecific (sardine) and interspecific (anchovy) density dependence in TEPPS/TEPPSW, which was previously unaccounted for in spawner–recruitment relationships. Such density‐dependent effects at the time of spawning should be considered in fisheries management and studies on recruitment mechanisms.

Mechanisms influencing the coexistence of multiple consumers and multiple resources: resource and apparent competition

AbstractInteractions among multiple resources and consumers involve two indirect interactions: resource competition among consumers and apparent competition among resources. However, competition among multiple consumers is typically viewed through the lens of direct interactions embodied in the Lotka‐Volterra competition model, which fails to capture the mechanisms of these indirect interactions. In this paper, I analyze various elaborations of MacArthur's and Tilman's original consumer–resource models including more than two species per trophic level, saturating functional responses, and direct intraspecific density dependence within the consumers. First, the simplest model with two resources and two consumers with linear functional responses is analyzed via the structure of the resulting isoclines, and this is reconciled with Tilman's graphical ZNGI/consumption vector approach. With three species at each trophic level even in this simple model, each consumer is not required to have the largest impact on the resource that most limits its growth for multiple consumers to coexist. In fact, a consumer that is an inferior competitor on each resource in isolation may still coexist with superior competitors, and conversely a consumer that is an inferior competitor on each available resource may still be able to drive all other consumers extinct. However, the maximum number of coexisting consumers is set by the number of available resources. Saturating functional responses do not qualitatively alter the conditions for multiple consumers and resources to coexist at a stable point equilibrium but do increase the range of apparent competitive abilities for resources that can invade and coexist. Saturating functional responses also increase the range of dynamics that the community may display (i.e., limit cycles and chaos), which previous analyses have shown can permit more consumer species than resources to coexist. Adding direct intraspecific density dependence in the consumers, either in the form of feeding interference or density‐dependent demographic rates, permit more consumers to coexist than available resources, even at stable point equilibria. Understanding the indirect effects that cascade through a community is essential to predicting community changes and understanding how species at multiple trophic levels coexist, and these indirect effects should not be shrouded behind the curtain of Lotka‐Volterra competition.

Intra-specific relatedness, spatial clustering and reduced demographic performance in tropical rainforest trees.

Intra-specific negative density dependence promotes species coexistence by regulating population sizes. Patterns consistent with such density dependence are frequently reported in diverse tropical tree communities. Empirical evidence demonstrating whether intra-specific variation is related to these patterns, however, is lacking. The present study addresses this important knowledge gap by genotyping all individuals of a tropical tree in a long-term forest dynamics plot in tropical China. We show that related individuals are often spatially clustered, but having closely related neighbours reduces the growth performance of focal trees. We infer from the evidence that dispersal limitation and negative density dependence are operating simultaneously to impact the spatial distributions of genotypes in a natural population. Furthermore, dispersal limitation decreases local intra-specific genetic diversity and increases negative density dependence thereby promoting niche differences and species coexistence as predicted by theory.