Contest competition inDrosophila subobscura

The role of larval intraspecific competition in laboratory populations ofDrosophila subobscura was investigated. Mortality is density-independent during the first 3 days after hatching but becomes density dependent as development proceeds to pupation. Although total biomass per patch was independent of initial egg density, competition betweenDrosophila larvae leads to the formation of smaller pupae. This resulted in a population that was dominated by suppressed individuals. Development rate ofD. subobscura larvae was not affected by high larval densities. Smaller pupae give rise to females with fewer eggs in their ovarioles. A simple simulation model, predicting the effects of intraspecific competition on the fecundity of the nextDrosophila generation is described.     

Partitioning the sources of demographic variation reveals density‐dependent nest predation in an island bird population

Ecological factors often shape demography through multiple mechanisms, making it difficult to identify the sources of demographic variation. In particular, conspecific density can influence both the strength of competition and the predation rate, but density‐dependent competition has received more attention, particularly among terrestrial vertebrates and in island populations. A better understanding of how both competition and predation contribute to density‐dependent variation in fecundity can be gained by partitioning the effects of density on offspring number from its effects on reproductive failure, while also evaluating how biotic and abiotic factors jointly shape demography. We examined the effects of population density and precipitation on fecundity, nest survival, and adult survival in an insular population of orange‐crowned warblers (Oreothlypis celata) that breeds at high densities and exhibits a suite of traits suggesting strong intraspecific competition. Breeding density had a negative influence on fecundity, but it acted by increasing the probability of reproductive failure through nest predation, rather than through competition, which was predicted to reduce the number of offspring produced by successful individuals. Our results demonstrate that density‐dependent nest predation can underlie the relationship between population density and fecundity even in a high‐density, insular population where intraspecific competition should be strong.     

Frequency-dependent community dynamics driven by sexual interactions

Research in community ecology has tended to focus on trophic interactions (e.g., predation, resource competition) as driving forces of community dynamics, and sexual interactions have often been overlooked. Here we discuss how sexual interactions can affect community dynamics, especially focusing on frequency-dependent dynamics of horizontal communities (i.e., communities of competing species in a single ecological guild). By combining mechanistic and phenomenological models of competition, we place sexual reproduction into the framework of modern coexistence theory. First, we review how population dynamics of two species competing for two resources can be represented by the Lotka–Volterra competition model as well as frequency dynamics, and how niche differentiation and overlap produce negative and positive frequency-dependence (i.e., stable coexistence and priority effect), respectively. Then, we explore two situations where sexual interactions change the frequency-dependence in community dynamics: (1) reproductive interference, that is, negative interspecific interactions due to incomplete species recognition in mating trials, can promote positive frequency-dependence and (2) density-dependent intraspecific adaptation load, that is, reduced population growth rates due to adaptation to intraspecific sexual (or social) interactions, produces negative frequency-dependence. We show how reproductive interference and density-dependent intraspecific adaptation load can decrease and increase niche differences in the framework of modern coexistence theory, respectively. Finally, we discuss future empirical and theoretical approaches for studying how sexual interactions and related phenomena (e.g., reproductive interference, intraspecific adaptation load, and sexual dimorphism) driven by sexual selection and conflict can affect community dynamics.     

Year-class coexistence in biennial plants

I extend the well known and biologically well motivated Skellam model of plant population dynamics to biennial plants. The model has two attractors: either one year class competitively excludes the other, resulting in 2-cycles with only vegetative vs only flowering plants in alternating years, or the two year classes coexist at an interior equilibrium. Contrary to earlier models, these two attractors can exist also simultaneously. I investigate the robustness of the model by including delayed flowering, a common phenomenon in plants, and provide a full numerical bifurcation analysis of the generalized model. High fecundity implies strong competition within year classes and promotes coexistence, whereas high survival results in strong competition between year classes and promotes competitive exclusion. Delayed flowering tends to stabilize the interior equilibrium, but (unlike in density-independent matrix models) the population cycles are robust with respect to some delay in flowering.     

Reproductive allocation patterns in different density populations of spring wheat

The effects of increased intraspecific competition on size hierarchies （size inequality） and reproductive allocation were investigated in populations of the annual plant, spring wheat （Triticurn aestivurn）. A series of densities （100, 300, 1 000, 3 000 and 10 000 plants/m^2） along a gradient of competition intensity were designed in this experiment. The results showed that average shoot biomass decreased with increased density. Reproductive allocation was negatively correlated to Gini coefficient （R^2 = 0.927）, which suggested that reproductive allocation is inclined to decrease as size inequality increases. These results suggest that both vegetative and reproductive structures were significantly affected by intensive competition. However, results also indicated that there were different relationships between plant size and reproductive allocation pattern in different densities. In the lowest density population, lacking competition （100 plants/m^2）, individual reproductive allocation was size independent but, in high density populations （300, 1 000, 3 000 and 10 000 plants/m^2）, where competition occurred, individual reproductive allocation was size dependent： the small proportion of larger individuals were winners in competition and got higher reproductive allocation （lower marginal reproductive allocation; MRA）, and the larger proportion of smaller individuals were suppressed and got lower reproductive allocation （higher MRA）. In conclusion, our results support the prediction that elevated intraspecific competition would result in higher levels of size inequality and decreased reproductive allocation （with a negative relationship between them）. However, deeper analysis indicated that these frequency- and size-dependent reproductive strategies were not evolutionarily stable strategies.     

Change of dynamic regimes in the population of species with short life cycles: Results of an analytical and numerical study

There is a phenomenon of multiregimism found in the elementary mathematical model of population dynamics, meaning the possibility for different dynamic regimes to exist under the same conditions, with transition to these regimes dependent on the initial numerical values. The effect in question comes into existence in the model which has several different limiting regimes (attractors): equilibrium, regular fluctuations, and chaotic attractor. The revealed phenomenon of multiregimism lets us explain the initiation of fluctuations as well as disappearance of fluctuations. Adequacy of the model's dynamic regimes is depicted by their correlation with the actual dynamics of population size of bank vole (Myodes glareolus). It is shown that the impact of climatic factors on a reproductive process of a population noticeably extends the range of possible dynamic regimes and, in fact, leads to random migration over attraction basins of these regimes.     

Maternal provisioning is structured by species’ competitive neighborhoods

Differential maternal provisioning of offspring in response to environmental conditions has been argued as ‘the missing link’ in plant life histories. Although empirical evidence suggests that maternal provisioning responses to abiotic conditions are common, there is little understanding of how differences in maternal provisioning manifest in response to competition. Frequency manipulations are commonly employed in ecological studies to assess the strength of interspecific competition, relative to intraspecific competition, and we used frequency manipulations to test how competition in two soil moisture environments affects maternal provisioning of seed mass. Specifically, for 15 pairs comprised from 25 annual plant species that occur in California, we varied the relative frequencies of conspecific to heterospecific competitors from 90% (intraspecific competition) to 10% (interspecific competition). We found that conspecific frequency affected maternal provisioning (seed mass) in 12 of the 25 species (eight significantly (p < 0.05), four marginally significantly (p < 0.07)), and that these responses included both increased (five species) and decreased (six species) seed mass, as well as one species with opposing directions of response to conspecific frequency that depended on the soil moisture environment. Conspecific frequency also affected per capita fecundity (seed number) for 17 of the 25 species (15 significantly (p < 0.05), two marginally significantly (p < 0.09)), which generally decreased seed number as conspecific frequency increased. The direction and magnitude of frequency‐dependent seed mass depended on the identity of the competitor, even among species whose fecundity was not affected by competitor identity; the latter finding reveals competitive differences among species that would otherwise appear to be competitively equivalent. Our research demonstrates how species responses to different competitive environments manifest through maternal provisioning, and that these responses alter previous estimates of environmentally‐determined maternal provisioning and reproductive output; future study is needed to understand their combined effects on population and community dynamics.     

濒危植物峨眉含笑的种内、种间竞争

研究植物种内、种间竞争关系是探究植物濒危原因的重要方式之一, 根据竞争来源和竞争预测模型可以制定具有针对性的保护策略。本文以雅安周公山峨眉含笑(Michelia wilsonii)野生种群为研究对象, 使用逐步扩大范围法确定峨眉含笑的竞争范围半径, 运用Hegyi单木竞争模型计算竞争指数(competition index, CI), 分析其种内、种间竞争关系。结果表明: 峨眉含笑的最适竞争范围半径是10 m, 能较好地反映其种内竞争强度; 峨眉含笑的竞争压力主要来自种内, 种内竞争指数(348.72, 62.52%)远大于种间竞争指数(209.03, 37.48%); 小树、中树阶段个体的竞争强度较大, 平均竞争指数(3.97、3.14)远高于总体平均竞争指数(2.68); 内有21种竞争木, 其中杉木(Cunninghamia lanceolata)、华中樱桃(Prunus conradinae)、细刺枸骨(Ilex hylonoma)是峨眉含笑的主要竞争树种; 胸径与竞争指数间服从指数函数关系(CI = 3.8907e^-0.048x, R² = 0.1087, P < 0.01), 随着对象木胸径的增大, 竞争指数不断降低, 当胸径达到30 cm后, 竞争强度基本稳定。综上, 小树、中树阶段的峨眉含笑个体受到极强的种内竞争, 初入老树阶段的个体受到较强的种间竞争。为更好地保护峨眉含笑的天然种群, 降低竞争对种群更替的影响, 在林分管理中需要促进小树、中树的个体更新, 减轻植株间的竞争消耗, 加速峨眉含笑的生长和维持种群稳定。     

Interspecific competition counteracts negative effects of dispersal on adaptation of an arthropod herbivore to a new host

Dispersal and competition have both been suggested to drive variation in adaptability to a new environment, either positively or negatively. A simultaneous experimental test of both mechanisms is however lacking. Here, we experimentally investigate how population dynamics and local adaptation to a new host plant in a model species, the two‐spotted spider mite (Tetranychus urticae), are affected by dispersal from a stock population (no‐adapted) and competition with an already adapted spider mite species (Tetranychus evansi). For the population dynamics, we find that competition generally reduces population size and increases the risk of population extinction. However, these negative effects are counteracted by dispersal. For local adaptation, the roles of competition and dispersal are reversed. Without competition, dispersal exerts a negative effect on adaptation (measured as fecundity) to a novel host and females receiving the highest number of immigrants performed similarly to the stock population females. By contrast, with competition, adding more immigrants did not result in a lower fecundity. Females from populations with competition receiving the highest number of immigrants had a significantly higher fecundity than females from populations without competition (same dispersal treatment) and than the stock population females. We suggest that by exerting a stronger selection on the adapting populations, competition can counteract the migration load effect of dispersal. Interestingly, adaptation to the new host does not significantly reduce performance on the ancestral host, regardless of dispersal rate or competition. Our results highlight that assessments of how species can adapt to changing conditions need to jointly consider connectivity and the community context.     

Linking the continental migratory cycle of the monarch butterfly to understand its population decline

Threats to several of the world's great animal migrations necessitate a research agenda focused on identifying drivers of their population dynamics. The monarch butterfly is an iconic species whose continental migratory population in eastern North America has been declining precipitously. Recent analyses have linked the monarch decline to reduced abundance of milkweed host plants in the USA caused by increased use of genetically modified herbicide‐resistant crops. To identify the most sensitive stages in the monarch's annual multi‐generational migration, and to test the milkweed limitation hypothesis, we analyzed 22 years of citizen science records from four monitoring programs across North America. We analyzed the relationships between butterfly population indices at successive stages of the annual migratory cycle to assess demographic connections and to address the roles of migrant population size versus temporal trends that reflect changes in habitat or resource quality. We find a sharp annual population decline in the first breeding generation in the southern USA, driven by the progressively smaller numbers of spring migrants from the overwintering grounds in Mexico. Monarch populations then build regionally during the summer generations. Contrary to the milkweed limitation hypothesis, we did not find statistically significant temporal trends in stage‐to‐stage population relationships in the mid‐western or northeastern USA. In contrast, there are statistically significant negative temporal trends at the overwintering grounds in Mexico, suggesting that monarch success during the fall migration and re‐establishment strongly contributes to the butterfly decline. Lack of milkweed, the only host plant for monarch butterfly caterpillars, is unlikely to be driving the monarch's population decline. Conservation efforts therefore require additional focus on the later phases in the monarch's annual migratory cycle. We hypothesize that lack of nectar sources, habitat fragmentation, continued degradation at the overwintering sites, or other threats to successful fall migration are critical limiting factors for declining monarchs.     

The effect of the Holling type II functional response on apparent competition

This article analyzes the classical 2-resource-1-consumer apparent competition community module with the Holling type II functional response. Two types of resource regulation (top-down vs. combined top-down and bottom-up) and two types of consumer behaviors (inflexible consumers with fixed preferences for resources vs. adaptive consumers) are considered. When resources grow exponentially and consumers are inflexible foragers, one resource is always outcompeted due to strong apparent competition. Density dependent resource growth relaxes apparent competition so that resources can coexist. As multiple attractors (either equilibria or limit cycles) coexist, population dynamics and community composition depend on initial population densities. Population dynamics change dramatically when consumers forage adaptively. In this case, the results both for top-down, and combined top-down and bottom-up regulation are similar and they show that species persistence occurs for a much larger set of parameter values when compared with inflexible consumers. Moreover, population dynamics will be chaotic when resource carrying capacities are high enough. This shows that adaptive consumer switching can destabilize population dynamics.     

Multiple attractors in a discrete competition model

The existence of multiple attractors in a competition model implies that the question of coexistence vs. extinction can depend on initial conditions. A discrete stage-structured model of two competing species is derived from a well-tested single-species model of insect populations, and is shown to exhibit multiple attractors for parameter values similar to those used in laboratory experiments which demonstrated chaos in population dynamics. The corresponding basins of attraction are investigated and shown to have very complex structures, and the initial stage structure of the populations is shown to have a significant impact on final outcomes.     

On evolution under asymmetric competition

The evolutionary consequences of asymmetric competition between species are poorly understood in comparison with symmetric competition. A model for evolution of body size under asymmetric competition within and between species is described. The model links processes operating at the scale of the individual to that of macroscopic evolution through a stochastic mutation–selection process. Phase portraits of evolution in a phenotype space characteristically show character convergence and parallel character shifts, with character divergence being relatively uncommon. The asymptotic states of evolution depend very much on the properties of asymmetric competition. Given relatively weak asymmetries between species, a single equilibrium point exists; this is a local attractor, and its position is determined by the intra- and interspecific asymmetries. When the asymmetries are made stronger, several fixed points may come about, creating further equilibrium points which are local attractors. It is also possible for periodic attractors to occur; such attractors comprise Red Queen dynamics with phenotype values that continue to change without ever settling down to constant values. From certain initial conditions, evolution leading to extinction of one of the species is also a likely outcome.     

Evolution of reproductive effort in viscous populations: the importance of population dynamics

I present two ecological models for the evolution of reproductive effort in viscous populations with empty sites. In contrast with previous studies, I show that limited dispersal needs not have a positive effect on the evolutionarily stable allocation of resources to fecundity versus survival. Rather, depending on the feedback between the trait and the population dynamics, population viscosity may have no effect or even lead to a decrease in the evolutionarily stable reproductive effort when individuals can degrade their environment during their lifetime. I show that the different evolutionary outcomes can be explained by the asymmetry in the level of kin competition resulting from investing into juveniles or into adults.     

Dietary niche and population dynamic feedbacks in a novel habitat

Population dynamics and resource use are often intricately connected via density‐dependent intraspecific competition. However, experimental studies of concurrent change in population and resource use dynamics are scarce. In particular, the impact of factors such as genetic diversity, which can affect both population dynamics and competition, remains unexplored. Using stable isotope analysis and periodic population censuses, we quantified both diet and population dynamics in wheat‐adapted Tribolium castaneum (flour beetle) populations provided with an additional novel resource (corn). Populations were initiated with different levels of genetic variation for traits relevant to population growth and resource use (e.g. fecundity and survival).We found that high population size decreased subsequent corn use, and high corn use in turn lowered population size. Surprisingly, we did not detect a significant effect of founding genetic variation on resource niche expansion, although genetic variation increased overall population size and stability. In contrast, dietary niche expansion decreased both population size and stability. Finally, larval and adult niche dynamics were uncorrelated, suggesting that various life stages perceive or respond differentially to intraspecific competition and resource availability. Our experiments indicate that population performance in a novel habitat depends on stage‐specific interactions between resource use, standing genetic variation, and population size.     

Population Interactions and the Determinants of Population Size

Abstract The flux of individuals within populations is dependent upon six controlling processes: the intrinsic rate of increase of the plant, intraspecific competition for resources, interspecific competition, natural enemies, mutualisms and refuge effects such as the immigration of seeds from other populations. Although population interactions are generally believed to play a major role in determining the flux of individuals within populations, little attempt has been made to quantify the strength of these interactions and their role in the dynamics of populations. In this paper we examine the role of competition, herbivory and mutualistic interactions in determining the dynamics of a range of annual plant species. Firstly, it is shown that the dynamics of three weed species ( Bromus sterilis, Galium aparine, Papaver rhoeas ) in an experimental community in an arable cropping system of winter wheat are determined primarily by the rapid population growth of B. sterilis . Interactions between the species play a minor part in the dynamics of the system. Secondly, it is shown that current levels of grazing by overwintering populations of brent geese have a minor impact on the abundance of Salicornia europaea , but that increased grazing has the potential to reduce abundance and increase the instability of S. europaea populations. This is a consequence of the aggregative response of the geese, which results in an increasing proportion of the seeds of S. europaea being eaten as plant density increases. Thirdly, it is shown that there is a complex interaction between root pathogenic and arbuscular mycorrhizal fungi in natural field populations of Vulpia ciliata and that the benefit of mycorrhizal fungi to the plant is in providing protection against pathogens.     

The frequency of multiple paternity predicts variation in testes size among island populations of house mice

Polyandry generates selection on males through sperm competition, which has broad implications for the evolution of ejaculates and male reproductive anatomy. Comparative analyses across species and competitive mating trials within species have suggested that sperm competition can influence the evolution of testes size, sperm production and sperm form and function. Surprisingly, the intraspecific approach of comparing among population variation for investigating the selective potential of sperm competition has rarely been explored. We sampled seven island populations of house mice and determined the frequency of multiple paternity within each population. Applying the frequency of multiple paternity as an index of the risk of sperm competition, we looked for selective responses in male reproductive traits. We found that the risk of sperm competition predicted testes size across the seven island populations of house mice. However, variation in sperm traits was not explained by sperm competition risk. We discuss these findings in relation to sperm competition theory, and other intrinsic and extrinsic factors that might influence ejaculate quality.     

Two-patch dispersal-linked compensatory-overcompensatory spatially discrete population models

We study the role of asynchronous and synchronous dispersals on discrete-time two-patch dispersal-linked population models, where the pre-dispersal local patch dynamics are of mixed compensatory and overcompensatory types. Single-species dispersal-linked models behave as single-species single-patch models whenever all pre-dispersal local patch dynamics are compensatory and dispersal is synchronous. However, the dynamics of the corresponding two-patch population model connected by asynchronous dispersal depends on the dispersal rates. The species goes extinct on at least one patch when the asynchronous dispersal rates are high, while it persists when the rates are low. We use numerical simulations to show that in both synchronous and asynchronous mixed compensatory and overcompensatory systems, symmetric and asymmetric dispersals can control and impede the onset of cyclic population oscillations via period-doubling reversal bifurcations. Also, we show that in mixed systems both asynchronous and synchronous dispersals are capable of altering the pre-dispersal local patch dynamics from overcompensatory to compensatory dynamics. Dispersal-linked population models with ‘unstructured’ overcompensatory pre-dispersal local dynamics connected by synchronous dispersal can generate multiple attractors with fractal basin boundaries. However, mixed compensatory and overcompensatory systems appear to exhibit single attractors and not coexisting (multiple) attractors.     

Influence of density‐dependent competition on foraging and migratory behavior of a subtropical colonial seabird

Density‐dependent competition for food resources influences both foraging ecology and reproduction in a variety of animals. The relationship between colony size, local prey depletion, and reproductive output in colonial central‐place foragers has been extensively studied in seabirds; however, most studies have focused on effects of intraspecific competition during the breeding season, while little is known about whether density‐dependent resource depletion influences individual migratory behavior outside the breeding season. Using breeding colony size as a surrogate for intraspecific resource competition, we tested for effects of colony size on breeding home range, nestling health, and migratory patterns of a nearshore colonial seabird, the brown pelican (Pelecanus occidentalis), originating from seven breeding colonies of varying sizes in the subtropical northern Gulf of Mexico. We found evidence for density‐dependent effects on foraging behavior during the breeding season, as individual foraging areas increased linearly with the number of breeding pairs per colony. Contrary to our predictions, however, nestlings from more numerous colonies with larger foraging ranges did not experience either decreased condition or increased stress. During nonbreeding, individuals from larger colonies were more likely to migrate, and traveled longer distances, than individuals from smaller colonies, indicating that the influence of density‐dependent effects on distribution persists into the nonbreeding period. We also found significant effects of individual physical condition, particularly body size, on migratory behavior, which in combination with colony size suggesting that dominant individuals remain closer to breeding sites during winter. We conclude that density‐dependent competition may be an important driver of both the extent of foraging ranges and the degree of migration exhibited by brown pelicans. However, the effects of density‐dependent competition on breeding success and population regulation remain uncertain in this system.     

The 'ghost of settlement past' determines mortality and fecundity in the barnacle, Semibalanus balanoides

Intraspecific competition is common in many organisms. For many sessile marine invertebrates high settlement densities can lead to competition with conspecifics that can affect the probability of mortality, morphology, and reproductive output. This work aims to determine the affect of recruit density on shell morphology and fecundity and the temporal nature of intraspecific competition for the acorn barnacle, Semibalanus balanoides . Two tile arrays were used; experiment 1 was deployed at the start of the settlement season (43,264 recruits, 4.1% survivorship to adult), experiment 2 was deployed in the middle of the season (5,454 recruits, 42.8% survivorship). Experiment 2 displayed density dependent mortality between recruits and juveniles and adults, whereas no relationship was found for exp. 1. Experiment 2 had higher egg production per individual, but also higher egg production per area of tile compared to exp. 1. A negative relationship was found between recruit density and egg production, however during the summer period, this relationship switched to a positive relationship between juvenile density and egg production, due to high mortality on tiles with high recruit density. Cumulative population density (CPD) was calculated as the cumulative mean density of barnacles per cm^-2 of tile and was used as an index of intraspecific competition. There was a linear negative relationship between mass of eggs per individual and CPD. However, CPD over the summer period more fully explained egg production per individual. A time window was identified when cumulative intraspecific competitive effects were shown to have a strong negative effect on egg production and an increase in mortality, causing a flip from density independent fecundity to density dependent fecundity. These results suggest that recruit density is an important driver of subsequent population processes.