Experimental Examination of Intraspecific Density-Dependent Competition during the Breeding Period in Monarch Butterflies (Danaus plexippus)

A central goal of population ecology is to identify the factors that regulate population growth. Monarch butterflies (Danaus plexippus) in eastern North America re-colonize the breeding range over several generations that result in population densities that vary across space and time during the breeding season. We used laboratory experiments to measure the strength of density-dependent intraspecific competition on egg laying rate and larval survival and then applied our results to density estimates of wild monarch populations to model the strength of density dependence during the breeding season. Egg laying rates did not change with density but larvae at high densities were smaller, had lower survival, and weighed less as adults compared to lower densities. Using mean larval densities from field surveys resulted in conservative estimates of density-dependent population reduction that varied between breeding regions and different phases of the breeding season. Our results suggest the highest levels of population reduction due to density-dependent intraspecific competition occur early in the breeding season in the southern portion of the breeding range. However, we also found that the strength of density dependence could be almost five times higher depending on how many life-stages were used as part of field estimates. Our study is the first to link experimental results of a density-dependent reduction in vital rates to observed monarch densities in the wild and show that the effects of density dependent competition in monarchs varies across space and time, providing valuable information for developing robust, year-round population models in this migratory organism.     

Accounting for interspecific competition and age structure in demographic analyses of density dependence improves predictions of fluctuations in population size

Understanding species coexistence has long been a major goal of ecology. Coexistence theory for two competing species posits that intraspecific density dependence should be stronger than interspecific density dependence. Great tits and blue tits are two bird species that compete for food resources and nesting cavities. On the basis of long‐term monitoring of these two competing species at sites across Europe, combining observational and manipulative approaches, we show that the strength of density regulation is similar for both species, and that individuals have contrasting abilities to compete depending on their age. For great tits, density regulation is driven mainly by intraspecific competition. In contrast, for blue tits, interspecific competition contributes as much as intraspecific competition, consistent with asymmetric competition between the two species. In addition, including age‐specific effects of intra‐ and interspecific competition in density‐dependence models improves predictions of fluctuations in population size by up to three times.     

Signs of stabilisation and stable coexistence

Many empirical studies motivated by an interest in stable coexistence have quantified negative density dependence, negative frequency dependence, or negative plant–soil feedback, but the links between these empirical results and ecological theory are not straightforward. Here, we relate these analyses to theoretical conditions for stabilisation and stable coexistence in classical competition models. By stabilisation, we mean an excess of intraspecific competition relative to interspecific competition that inherently slows or even prevents competitive exclusion. We show that most, though not all, tests demonstrating negative density dependence, negative frequency dependence, and negative plant–soil feedback constitute sufficient conditions for stabilisation of two‐species interactions if applied to data for per capita population growth rates of pairs of species, but none are necessary or sufficient conditions for stable coexistence of two species. Potential inferences are even more limited when communities involve more than two species, and when performance is measured at a single life stage or vital rate. We then discuss two approaches that enable stronger tests for stable coexistence‐invasibility experiments and model parameterisation. The model parameterisation approach can be applied to typical density‐dependence, frequency‐dependence, and plant–soil feedback data sets, and generally enables better links with mechanisms and greater insights, as demonstrated by recent studies.     

Pathogens, density dependence and the coexistence of tropical trees

There is increasing interest in the role played by density-dependent mortality from natural enemies, particularly plant pathogens, in promoting the coexistence and diversity of tropical trees. Here, we review four issues in the analysis of pathogen-induced density dependence that have been overlooked or inadequately addressed. First, the methodology for detecting density dependence must be robust to potential biases. Observational studies, in particular, require a careful analysis to avoid biases generated by measurement error, and existing studies could be criticized on these grounds. Experimental studies manipulating plant density and pathogen incidence will often be preferable, or should be run in parallel. Second, the form of density dependence is not well understood and, in particular, there are no data indicating whether pathogens cause compensating or overcompensating density responses. Owing to this, we argue that the potential for pathogen-induced density dependence to generate diversity-enhancing outcomes, such as the Janzen-Connell effect, remains uncertain, as coexistence is far more probable if density dependence is overcompensating. Third, there have been few studies examining the relative importance of intra- or interspecific density dependence resulting from pathogens (or, more widely, natural enemies). This is essentially equivalent to asking to what extent pathogens are host-specific. If pathogens are generalists, then mortality rates will respond to overall plant density, irrespective of plant species identity. This will weaken the intraspecific density dependence and reduce the diversity-promoting effects of pathogens. Finally, we highlight the need for studies that integrate observations and experiments on pathogens and density dependence into the whole life cycle of trees, because as yet it is not possible to be certain of the degree to which pathogens contribute to observed dynamics.     

Bias in the detection of negative density dependence in plant communities

Regression dilution is a statistical inference bias that causes underestimation of the strength of dependency between two variables when the predictors are error‐prone proxies (EPPs). EPPs are widely used in plant community studies focused on negative density‐dependence (NDD) to quantify competitive interactions. Because of the nature of the bias, conspecific NDD is often overestimated in recruitment analyses, and in some cases, can be erroneously detected when absent. In contrast, for survival analyses, EPPs typically cause NDD to be underestimated, but underestimation is more severe for abundant species and for heterospecific effects, thereby generating spurious negative relationships between the strength of NDD and the abundances of con‐ and heterospecifics. This can explain why many studies observed rare species to suffer more severely from conspecific NDD, and heterospecific effects to be disproportionally smaller than conspecific effects. In general, such species‐dependent bias is often related to traits associated with likely mechanisms of NDD, which creates false patterns and complicates the ecological interpretation of the analyses. Classic examples taken from literature and simulations demonstrate that this bias has been pervasive, which calls into question the emerging paradigm that intraspecific competition has been demonstrated by direct field measurements to be generally stronger than interspecific competition.     

Effects of intraspecific and interspecific density on the demography of a perennial herb, Sanicula europaea

We examined the effects of intraspecific and interspecific competition on demographic processes in the perennial herb Sanicula europaea by manipulating the density of neighbouring plants. We followed the response in terms of survival, growth and reproduction and in terms of seedling recruitment. The demographic data from all phases of the life cycle enabled us to assess also the overall effects of treatments on population growth rate (λ) by transition matrix models. We also decomposed the differences in λ between control and treatments, using life table response experiments (LTRE). To study the effects of competition on recruitment in more detail and to evaluate the role of seed availability, we sowed seeds at different densities with or without vegetation removal.
Vegetative growth and flowering frequency of established individuals was not significantly affected by removal treatments, which suggest no, or a delayed response to released competition. Neighbour removal had no effect on seedling emergence but enhanced recruitment through a higher seedling survival. Conspecific and simultaneous conspecific and heterospecific removal of plants led to an increase in population growth rate (λ), whereas heterospecific removal alone led to a decrease. Emergence of seedlings and fate of vegetative established individuals contributed most to differences in λ between the control and the different treatments. Seed addition enhanced seedling emergence but, as seedling and juvenile survival were density dependent, densities of established individuals appear not to be seed limited.
In S. europaea removal treatments had different effects on established individuals and recruitment. This suggests that studies quantifying the effects of competition over the entire life cycle and performed in a natural environment are necessary to assess the importance of competition in perennial plant populations.     

Pollination outcomes reveal negative density‐dependence coupled with interspecific facilitation among plants

Pollination is thought to be under positive density‐dependence, destabilising plant coexistence by conferring fitness disadvantages to rare species. Such disadvantage is exacerbated by interspecific competition but can be mitigated by facilitation and intraspecific competition. However, pollinator scarcity should enhance intraspecific plant competition and impose disadvantage on common over rare species (negative density‐dependence, NDD). We assessed pollination proxies (visitation rate, pollen receipt, pollen tubes) in a generalised plant community and related them to conspecific and heterospecific density, expecting NDD and interspecific facilitation due to the natural pollinator scarcity. Contrary to usual expectations, all proxies indicated strong intraspecific competition for common plants. Moreover interspecific facilitation prevailed and was stronger for rare than for common plants. Both NDD and interspecific facilitation were modulated by specialisation, floral display and pollinator group. The combination of intraspecific competition and interspecific facilitation fosters plant coexistence, suggesting that pollination can be a niche axis maintaining plant diversity.     

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.     

European butterfly populations vary in sensitivity to weather across their geographical ranges

Aim

The aim was to assess the sensitivity of butterfly population dynamics to variation in weather conditions across their geographical ranges, relative to sensitivity to density dependence, and determine whether sensitivity is greater towards latitudinal range margins.

Location

Europe.

Time period

1980–2014.

Major taxa studied

Butterflies.

Methods

We use long‐term (35 years) butterfly monitoring data from > 900 sites, ranging from Finland to Spain, grouping sites into 2° latitudinal bands. For 12 univoltine butterfly species with sufficient data from at least four bands, we construct population growth rate models that include density dependence, temperature and precipitation during distinct life‐cycle periods, defined to accommodate regional variation in phenology. We use partial R² values as indicators of butterfly population dynamics' sensitivity to weather and density dependence, and assess how these vary with latitudinal position within a species' distribution.

Results

Population growth rates appear uniformly sensitive to density dependence across species' geographical distributions, and sensitivity to density dependence is typically greater than sensitivity to weather. Sensitivity to weather is greatest towards range edges, with symmetry in northern and southern parts of the range. This pattern is not driven by variation in the magnitude of weather variability across the range, topographic heterogeneity, latitudinal range extent or phylogeny. Significant weather variables in population growth rate models appear evenly distributed across the life cycle and across temperature and precipitation, with substantial intraspecific variation across the geographical ranges in the associations between population dynamics and specific weather variables.

Main conclusions

Range‐edge populations appear more sensitive to changes in weather than those nearer the centre of species' distributions, but density dependence does not exhibit this pattern. Precipitation is as important as temperature in driving butterfly population dynamics. Intraspecific variation in the form and strength of sensitivity to weather suggests that there may be important geographical variation in populations' responses to climate change.     

Assessing the spatial variability of density dependence in waterfowl populations

Population models commonly assume that the demographic parameters are spatially invariant, but there is considerable evidence that population growth rate (r) and the strength of density dependence (β) can vary over a species' range. To address this issue we developed a spatially explicit Gompertz population model based on the spatially varying coefficients approach to assess the spatial variation in population drivers. The model was fit to spatially stratified time series population estimates of the mallard Anas platyrhynchos in western North America. We included precipitation during the previous year and spring maximum temperature in the current year as environmental factors in the density dependent population model. Because density dependent models can give biased estimates for time series of abundance data, we fit a naïve model without informative priors and a model where we constrained the mean and variance of r to biologically realistic values that were derived via a comparative demography approach. In the naïve model, r and β were not separately identifiable and their values were overestimated, leading to unrealistic population growth. The naïve model also implied spatial variation in population r and the return time to equilibrium [1?(– β)] across the survey area. In contrast, in the informative model, r and the return time to equilibrium did not vary markedly among populations and were generally equal across populations. The effects of the climatic factors were similar across models. Population growth rates in the Prairie‐pothole region were positively correlated with precipitation, while in Alaska rates were positively correlated with spring temperature. Although it has been argued in the past that adding ecological realism could help avoid the pitfalls associated with density dependent models, our results demonstrate that imposing constraints on the population parameters is still the best course of action.     

Density dependence in blowfly populations: experimental evaluation of non‐parametric time‐series modelling

Two approaches for describing density dependence in demographic rates of stage‐structured populations are compared in this study. Time‐series data from laboratory blowfly populations (Lucilia sericata) have been analysed in a separate study, with a statistical modelling approach that incorporated density dependences as unspecified (non‐parametric) functions. In this study, we assessed density‐dependent structures by manipulating densities of larvae and adults in cohorts of blowflies and measuring the demographic rates. We here compare the density‐dependent structures revealed by the cohort experiments with those estimated by the non‐parametric model. This model estimates the demographic rates to have the following density‐dependent structures: (i) larval survival was non‐linearly density‐dependent (a ‘humped’ function), (ii) adult survival was density‐independent, and (iii) reproductive rate decreased with adult density. In the cohort experiments reported here, (i) juvenile survival exhibited a positive density dependence in low densities (facilitation), which became negative at higher densities (competition). Pupal and adult size decreased with initial larval density. (ii) Adult survival was reduced by high initial larval density, but it was independent of adult density. (iii) Reproductive rate was reduced by high initial larval density, and by high adult density in populations of large individuals (from low larval density). Hence, the results from these experiments support the non‐parametric model estimates regarding density‐dependent structures of demographic rates in the blowfly populations. The mean demographic rates, however, were apparently underestimated by the model. We conclude that non‐parametric modelling is a useful first approach for exploratory analysis of ecological time‐series data.     

Experimental evidence that competition and habitat use shape the individual fitness surface

A key prediction made by theories of density‐dependent competition is that resource overlap should increase the intensity of competition. By extension, we can predict that competition should lead to density‐dependent natural selection. I studied natural selection on limb length and body size in a total of seven populations of Anolis sagrei over 3 years in the Bahamas. Experimental manipulations of population density on small off‐shore cays revealed that the strength of natural selection on body size increased with density, suggesting that density‐dependent intraspecific competition drives natural selection. At low density, reduced competition revealed significant selection on limb length driven by changes in perch diameter, indicating that selection favoured a match between morphology and habitat. The role habitat played in shaping selection was further illuminated by inter‐annual changes in vegetation structure stemming from variation in precipitation among years. Thus, changes in both the intensity of competition across spatial replicates, and in resource availability through time, revealed changes in the targets of natural selection. Results provide empirical support for the long‐standing hypothesis that density‐dependent natural selection shapes the fitness surface of Greater Antilles anoles.     

Patterns and processes in shorebird survival rates: a global review

Changes in demographic rates underpin changes in population size, and understanding demographic rates can greatly aid the design and development of strategies to maintain populations in the face of environmental changes. However, acquiring estimates of demographic parameters at relevant spatial scales is difficult. Measures of annual survival rates can be particularly challenging to obtain because large‐scale, long‐term tracking of individuals is difficult and the resulting data contain many inherent biases. In recent years, advances in both tracking and analytical techniques have meant that, for some taxonomic groups, sufficient numbers of survival estimates are available to allow variation within and among species to be explored. Here we review published estimates of annual adult survival rates in shorebird species across the globe, and construct models to explore the phylogenetic, geographical, seasonal and sex‐based variation in survival rates. Models of 295 survival estimates from 56 species show that survival rates calculated from recoveries of dead individuals or from return rates of marked individuals are significantly lower than estimates from mark–recapture models. Survival rates also vary across flyways, largely as a consequence of differences in the genera that have been studied and the analytical methods used, with published estimates from the Americas and from smaller shorebirds (Actitis, Calidris and Charadrius spp.) tending to be underestimated. By incorporating the analytical method used to generate each estimate within a mixed model framework, we provide method‐corrected species‐specific and genus‐specific adult annual survival estimates for 52 species of 15 genera.     

Lolium multiflorum density responses under ozone and herbicide stress

Adaptations to overcrowding of individual plants result in density dependant control of growth and development. There is little information on how anthropogenic stresses modify these responses. We investigated whether combinations of diclofop‐methyl herbicide and tropospheric ozone alter the pattern of expected growth compensation with density changes resulting from intraspecific competition in Lolium multiforum Lam (Poacea) plants. Individual plant vegetative parameters and total seed production were assessed for plants growing under various densities and different herbicide rates and ozone treatments. The stressors differently changed the frequency distribution for average individual plant weight resulting from increasing densities. Only herbicide affected seedling mortality. Plants were able to compensate during grain filling maintaining similar seed production – density relationships in all treatments. Our findings contribute to the understanding of the impact of stress factors on the demographic changes in plant populations. Important ecological implications arise: (i) contrasting responses to ozone and herbicide, alone and in combination of individual plants resulted in different biomass – density relationships; (ii) stress effects on plant populations could not be predicted from individual responses; and (iii) changes in competitive outcome by single or combined stress factors may alter the expected genotype frequency in a crowded population with few dominant individuals.     

Apparent survival of tropical birds in a wet,premontane forest in Costa Rica

Despite the importance of tropical birds in the development of life history theory, we lack information about demographic rates and drivers of population dynamics for most species. We used a 7‐year (2007–2013) capture‐mark‐recapture dataset from an exceptionally wet premontane forest at mid‐elevation in Costa Rica to estimate apparent survival for seven species of tropical passerines. For four of these species, we provide the first published demographic parameters. Recapture probabilities ranged from 0.21 to 0.53, and annual estimates of apparent survival varied from 0.23 to 1.00. We also assessed the consequences of inter‐annual variation in rainfall on demographic rates. Our results are consistent with inter‐annual rainfall increasing estimates of apparent survival for two species and decreasing estimates for three species. For the three species where we could compare our estimates of apparent survival to estimates from drier regions, our estimates were not consistently higher or lower than those published previously. The temporal and spatial variability in demographic rates we document within and among species highlights the difficulties of generalizing life history characteristics across broad biogeographic gradients. Most importantly, this work emphasizes the context‐specific role of precipitation in shaping tropical avian demographic rates and underscores the need for mechanistic studies of environmental drivers of tropical life histories.     

Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra‐ and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five‐fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra‐ and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.     

The interaction effect between intraspecific competition and seed quality on the life‐history traits of the seed‐feeding beetle Acanthoscelides macrophthalmus

It has been shown that intraspecific competition and resource quality may affect life‐history traits of insects, such as body size, fecundity, and survival. However, intraspecific competition and resource quality may interact with each other. The study of such interacting effects is crucial for understanding the influence of these ecological variables on the selection of specific life‐history traits. Here, we investigated whether the interaction between intraspecific larval competition and variation in resource quality affects adult emergence and survival, egg size, fecundity, body size, and sexual size dimorphism (SSD) of the seed‐feeding beetle Acanthoscelides macrophthalmus (Schaeffer) (Coleoptera: Chrysomelidae: Bruchinae) when infesting Leucaena leucocephala (Lam.) De Wit (Fabaceae), its host plant. In the laboratory, beetles were reared on seeds that differed in quality (e.g., different hardness, seed size, water content), in the presence or absence of larval competition. Body size and SSD did not differ between treatments (with and without competition), nor were they affected by varying resource quality. Females subjected to competition during the larval stage and females emerging from seeds of higher quality, displayed the highest fecundity. The proportion of emergent adults was higher in the absence of competition. In addition, larger eggs were laid on the low‐quality resource in the absence of competition, showing a trade‐off between egg size and egg number. Adult survival differed among treatments and resource qualities, suggesting a higher investment in adult survival for individuals emerging from seeds of low quality in the presence of competition. Whether changes in specific traits could be selected for in detriment of others will depend on the strength of intraspecific competition, the variation in resource quality, and the plasticity in the life‐history traits investigated. This needs further clarification.     

Delayed and inverse density dependence in a chamois population of the Italian Alps

Both density dependent and density independent factors have been considered important determinants of the dynamics of ungulate populations Intraspecific competition for food and the amount of snow cover were suggested in the past as factors that influence the demographic parameters of the chamois Rupicapra rupicapra We present a 10 yr study on a closed population of the Italian Alps Animals, divided by sex and age class (kid, yearling, subadult, adult, old), were counted over the period 1981–1990 The number of chamois hunted and the number of carcasses found each year and the daily snow height were also available We investigated possible correlations between several demographic parameters and both snow cover and chamois density, accounting for sex- and age-differential mortality and a possible delay in density dependence Results are as follows a) there is no statistically significant correlation between snow cover and demographic rates, b) there is no detectable compensation between natural and hunting-related mortality, c) birth rate is density independent for any time delay, d) when calculated from carcasses, total mortality, mortality of 1-yr-old and older males and females significantly depend on total density with 2 years' lag, while kid mortality is not related to density for any time delay, e) when calculated from censuses, all mortality rates significantly depend on total density with 2 years' lag, although the variance explained by delayed density is definitely smaller for kids, f) nonlinear regression of kid mortality from censuses against density of kids in the same year yields a much higher coefficient of determination and inverse density dependence is evidenced However, determination between delayed and inverse density dependence in kid mortality is impossible, given our dataset In general, results support the hypothesis that intraspecific competition, epidemics and predation, rather than weather, play a role in the dynamics of this ungulate population     

Intraspecific competition facilitates the evolution of tolerance to insect damage in the perennial plant Solanum carolinense

Tolerance to herbivory (the degree to which plants maintain fitness after damage) is a key component of plant defense, so understanding how natural selection and evolutionary constraints act on tolerance traits is important to general theories of plant–herbivore interactions. These factors may be affected by plant competition, which often interacts with damage to influence trait expression and fitness. However, few studies have manipulated competitor density to examine the evolutionary effects of competition on tolerance. In this study, we tested whether intraspecific competition affects four aspects of the evolution of tolerance to herbivory in the perennial plant Solanum carolinense: phenotypic expression, expression of genetic variation, the adaptive value of tolerance, and costs of tolerance. We manipulated insect damage and intraspecific competition for clonal lines of S. carolinense in a greenhouse experiment, and measured tolerance in terms of sexual and asexual fitness components. Compared to plants growing at low density, plants growing at high density had greater expression of and genetic variation in tolerance, and experienced greater fitness benefits from tolerance when damaged. Tolerance was not costly for plants growing at either density, and only plants growing at low density benefited from tolerance when undamaged, perhaps due to greater intrinsic growth rates of more tolerant genotypes. These results suggest that competition is likely to facilitate the evolution of tolerance in S. carolinense, and perhaps in other plants that regularly experience competition, while spatio-temporal variation in density may maintain genetic variation in tolerance.     

The shifting balance of facilitation and competition affects the outcome of intra- and interspecific interactions over the life history of California grassland annuals

Trait-based resource competition in plants, wherein more similar plants compete more strongly for resources, is a foundation of niche-based explanations for the maintenance of diversity in plant communities. Alternatively, neutral theory predicts that community diversity can be maintained despite equivalent resource requirements among species. We examined interactions at three life history stages (germination, survival, and juvenile-adult growth) for three native and three exotic California annual species in a glasshouse experiment. We varied plant density and species composition in small pots, with pots planted with either intraspecific seeds or in a three species mix of intra- and interspecific neighbors. We saw a range of facilitative, neutral, and competitive interactions that varied significantly by species, rather than by native or exotic status. There were more competitive interactions at the emergence and juvenile-adult growth stages and more facilitative interactions for survival. Consequently, the relative strength of competition in intraspecific versus mixed-species communities depended on whether we considered only the juvenile-adult growth stage or the entire life history of the interacting plants. Using traditional analysis of juvenile-adult growth only, all species showed negative density-dependent interactions for final biomass production. However, when the net effect of plant interactions from seed to adult was considered, which is a prediction of population growth, two native species ceased to show negative density dependence, and the difference between intraspecific and mixed-species competition was only significant for one exotic species. Results were consistent with predictions of neutral, rather than niche, theory for five of six species.