Evidence for an even sex allocation in haplodiploid cyclical parthenogens

Recent theoretical work has shown that haplodiploid cyclical parthenogens, such as rotifers, are expected to have an equal frequency of male‐producing and resting‐egg producing females during their sexual phase. We tested this prediction by following sexual reproduction dynamics in two laboratory populations and one field population of the rotifer Brachionus plicatilis through two growing seasons. We recorded population density, proportion of sexual females, and sex allocation (the proportion of male‐producing sexual females as a fraction of total sexual females). We found this sex allocation ratio to vary from 0.3 to 1.0 in single sampling events. However, when we computed sex allocation by using the integrated densities of both male‐producing sexual females and resting‐egg producing sexual females over time, the two laboratory populations and one of the two field growing seasons showed sex allocation ratios that did not significantly differ from the expected value of 0.5.     

Density-dependent regulation of natural and laboratory rotifer populations

Density-dependent regulation of abundance is fundamentally important in the dynamics of most animal populations. Density effects, however, have rarely been quantified in natural populations, so population models typically have a large uncertainty in their predictions. We used models generated from time series analysis to explore the form and strength of density-dependence in several natural rotifer populations. Population growth rate (r) decreased linearly or non-linearly with increased population density, depending on the rotifer species. Density effects in natural populations reduced r to 0 at densities of 1–10 l^–1 for 8 of the 9 rotifer species investigated. The sensitivities of these species to density effects appeared normally distributed, with a mean r=0 density of 2.3 l^–1 and a standard deviation of 1.9. Brachionus rotundiformis was the outlier with 10–100× higher density tolerance. Density effects in laboratory rotifer populations reduced r to 0 at population densities of 10–100 ml^–1, which is 10⁴ higher than densities in natural populations. Density effects in laboratory populations are due to food limitation, autotoxicity or to their combined effects. Experiments with B. rotundiformis demonstrated the absence of autotoxicity at densities as high as 865 ml^–1, a much higher density than observed in natural populations. It is, therefore, likely that food limitation rather than autotoxicity plays a major role in regulating natural rotifer populations.     

Sex,population dynamics and resting egg production in rotifers

The interaction between sexual reproduction and population growth in the rotifer Brachionus plicatilis was examined using exponential and logistic growth models. A computer simulation was used to explore the effects of the frequency of sex and the proportion of a female's daughters reproducing sexually on population growth rate and resting egg production. Within the parameters of the simulation, the proportion of sexual daughters required for maximum resting egg production increased as the frequency of sex decreased. Adding density-dependence to the model also shifted this optimum towards a higher proportion of sexual daughters. When population growth was unconstrained by density, the optimum proportion of sexual daughters varied between 10–50%, depending on the frequency of sex. This compares to 25–64% when the greatest density restraints were applied at K = 500 females. As the population growth rate (r) increases, the optimum proportion of sexual daughters increases nonlinearly and these effects are exaggerated as sex becomes less frequent. A compilation of published data from laboratory populations of four Brachionus plicatilis strains reveals that the average proportion of sexual daughters in 21% and in close agreement with that predicted by simulation. The limitations of the simulation and its generalizability to other rotifers are discussed.     

镜湖常见臂尾轮虫(Brachionus)种群周年动态及生活史对策

对镜湖臂尾轮虫种群的周年动态进行了研究,测量和分析了它们的个体大小、卵大小和相对卵大小及其与水温和叶绿素a浓度等生态因子间的关系。结果表明,镜湖常见的7种臂尾轮虫中,仅萼花臂尾轮虫(Brachionus calyciflorus)、角突臂尾轮虫(B.angularis)、裂足臂尾轮虫(B.diversicornis)和剪形臂尾轮虫(B.forficula)的种群密度和出现频率较高;前两者种群密度高峰期均出现在2~3月份,4月份后均较低;后两者种群密度在6~9月份较高,而在10月份后很快下降。除剪形臂尾轮虫外,其它3种臂尾轮虫的个体体积均与水温间具有显著的相关性,平均卵体积均与各自的平均被甲长度呈显著的正相关关系。角突臂尾轮虫和萼花臂尾轮虫个体大小与水体叶绿素a浓度间呈显著的负相关关系,它们的卵体积分别与水温和叶绿素a浓度之间具有显著的负相关性,且水温是影响它们卵体积的主要因子。4种臂尾轮虫中,仅角突臂尾轮虫的相对卵体积与水温间表现出显著的正相关关系。     

Sex, parthenogenesis and genetic structure of rotifers: microsatellite analysis of contemporary and resting egg bank populations

Cyclically parthenogenetic rotifers are a valuable model for investigating the relationship between reproductive mode and population structure, although advances in this field have been hindered by low allozyme variability in these organisms. A high genotypic diversity is predicted after population establishment, which would be eroded by clonal selection during the parthenogenetic phase. The resting egg bank, produced sexually, is presumed to store high levels of genetic diversity, with subsequent effects on planktonic population structure. Here, we provide the first application of microsatellite markers to a rotifer planktonic population and its associated resting egg bank. Seven polymorphic microsatellite loci were screened in populations of the rotifer Brachionus plicatilis in a temporary pond to analyse: (i) the genetic structure of the resting egg bank; (ii) the changes in the genetic structure of rotifer populations during the parthenogenetic phase; and (iii) the population structure after its initiation from resting eggs. Microsatellites proved to be a useful tool for clone identification, revealing a surprisingly high clonal diversity in rotifer populations. The last sample in the parthenogenetic phase showed evidence of clonal selection, as indicated by a low observed clonal diversity and the appearance of linkage disequilibria. The resting egg bank, analysed comprehensively for the first time in any zooplankter, is in Hardy-Weinberg and linkage equilibrium, and contains a high genotypic diversity. Unexpectedly, the resting egg bank differed from the planktonic population in its allelic composition, suggesting that resting egg hatching is biased.     

Variability for mixis initiation in Brachionus plicatilis

Deductions from both evolutionary models and inductive argumentation from empirical data support the notion of intraspecific variability for the initiation of sexual reproduction (mixis) within rotifer populations. In this study, we focus on the time and density at which mixis is initiated in a growing population. Cyclical parthenogenetic clones of Brachionus plicatilis established by hatching of resting eggs, isolated from a natural habitat, have been tested at the start of their sexual phase. Clones exhibited great variation for this trait, their time of switching to sexual reproduction being correlated with population density. Most of the variation for mixis initiation has either low or no heritability and is caused by individual environmental factors.     

Population dynamics of three songbird species in a nestbox population in Central Europe show effects of density,climate and competitive interactions

Unravelling the contributions of density‐dependent and density‐independent factors in determining species population dynamics is a challenge, especially if the two factors interact. One approach is to apply stochastic population models to long‐term data, yet few studies have included interactions between density‐dependent and density‐independent factors, or explored more than one type of stochastic population model. However, both are important because model choice critically affects inference on population dynamics and stability. Here, we used a multiple models approach and applied log‐linear and non‐linear stochastic population models to time series (spanning 29 years) on the population growth rates of Blue Tits Cyanistes caeruleus, Great Tits Parus major and Pied Flycatchers Ficedula hypoleuca breeding in two nestbox populations in southern Germany. We focused on the roles of climate conditions and intra‐ and interspecific competition in determining population growth rates. Density dependence was evident in all populations. For Blue Tits in one population and for Great Tits in both populations, addition of a density‐independent factor improved model fit. At one location, Blue Tit population growth rate increased following warmer winters, whereas Great Tit population growth rates decreased following warmer springs. Importantly, Great Tit population growth rate also decreased following years of high Blue Tit abundance, but not vice versa. This finding is consistent with asymmetric interspecific competition and implies that competition could carry over to influence population dynamics. At the other location, Great Tit population growth rate decreased following years of high Pied Flycatcher abundance but only when Great Tit population numbers were low, illustrating that the roles of density‐dependent and density‐independent factors are not necessarily mutually exclusive. The dynamics of this Great Tit population, in contrast to the other populations, were unstable and chaotic, raising the question of whether interactions between density‐dependent and density‐independent factors play a role in determining the (in) stability of the dynamics of species populations.     

Co‐adaptation impacts the robustness of predator–prey dynamics against perturbations

Global change threatens the maintenance of ecosystem functions that are shaped by the persistence and dynamics of populations. It has been shown that the persistence of species increases if they possess larger trait adaptability. Here, we investigate whether trait adaptability also affects the robustness of population dynamics of interacting species and thereby shapes the reliability of ecosystem functions that are driven by these dynamics. We model co‐adaptation in a predator–prey system as changes to predator offense and prey defense due to evolution or phenotypic plasticity. We investigate how trait adaptation affects the robustness of population dynamics against press perturbations to environmental parameters and against pulse perturbations targeting species abundances and their trait values. Robustness of population dynamics is characterized by resilience, elasticity, and resistance. In addition to employing established measures for resilience and elasticity against pulse perturbations (extinction probability and return time), we propose the warping distance as a new measure for resistance against press perturbations, which compares the shapes and amplitudes of pre‐ and post‐perturbation population dynamics. As expected, we find that the robustness of population dynamics depends on the speed of adaptation, but in nontrivial ways. Elasticity increases with speed of adaptation as the system returns more rapidly to the pre‐perturbation state. Resilience, in turn, is enhanced by intermediate speeds of adaptation, as here trait adaptation dampens biomass oscillations. The resistance of population dynamics strongly depends on the target of the press perturbation, preventing a simple relationship with the adaptation speed. In general, we find that low robustness often coincides with high amplitudes of population dynamics. Hence, amplitudes may indicate the robustness against perturbations also in other natural systems with similar dynamics. Our findings show that besides counteracting extinctions, trait adaptation indeed strongly affects the robustness of population dynamics against press and pulse perturbations.     

Density-dependent population regulation detected in short time series of saproxylic beetles

Understanding the regulation of natural populations has been a long-standing problem in ecology. Here we analyze the population dynamics of 17 species of saproxylic beetles in Shizuoka Prefecture, Japan collected over 11–12 years using autoregressive integrated moving average (ARIMA) models. We first examined the dynamics for indications of the order of the ARIMA models and evaluated the time series to determine that it was not simply a random, white noise sequence. All species dynamics were not mere random noise, and ARIMA models up to lag 3 were considered. The best model was selected from the possible ones using several criteria: model convergence, weak residual autocorrelation, the small sample AIC must be among the smallest that were not significantly different, and the lag indicated by the cutoff values in the detrended partial autocorrelation function. We found significant and nearly significant direct density-dependence for 14 of the 17 species, varying from −0.709 and stronger. The characteristic return rates were strong and only one species had a weak return rate (>0.9), implying that these species were strongly regulated by density-dependent factors. We found that populations with higher order ARIMA models (lag 2 and 3) had weaker return rates than populations with ARIMA models with only one lag, suggesting that species with more complex dynamics were more weakly regulated. These results contrast with previous suggestions that 20+ years are needed to detect density dependence from population time series and that most populations are weakly regulated.     

Molecular Ecology of Rotifers: From Population Differentiation to Speciation

The development of cost-effective molecular tools allowing the amplification of minute amounts of DNA effectively opened the field of molecular ecology for rotifers. Here I review these techniques and the advances they have provided in the understanding of sibling species complexes, clonal structure, resting egg banks, population structure, phylogeographic patterns and phylogenetic relationships in rotifers. Most of the research to date has focused on the rotifer species complex Brachionus plicatilis. The use of DNA sequence and microsatellite variation, in the context of the background knowledge of life history, mating behaviour, and temporal population dynamics in these organisms have revolutionised our views into the processes shaping the genetic diversity in aquatic invertebrates. Rotifers have populations with a very high number of clones in genetic equilibrium. In temporary populations clonal selection is effective in eroding the number of clones. Rotifer populations are strongly differentiated genetically for neutral markers, even at small geographical scales, and exhibit deep phylogeographic structure which might reflect the impact of Pleistocene glaciations. Despite the high potential for dispersal afforded by resting eggs, rotifers display persistent historical colonisation effects, with gene flow effective only at a local scale and with marked isolation by distance. Instances of long-distance transcontinental migration resulting in successful colonisation have also been revealed. B. plicatilis is composed of a group of several ancient species and sympatry is common. Despite this, the presence of cosmopolitan species in this species complex cannot be discounted. I discuss future priorities and point out the main areas where our knowledge is still insufficient.     

Population responses to perturbations: the importance of trait-based analysis illustrated through a microcosm experiment

Environmental change continually perturbs populations from a stable state, leading to transient dynamics that can last multiple generations. Several long-term studies have reported changes in trait distributions along with demographic response to environmental change. Here we conducted an experimental study on soil mites and investigated the interaction between demography and an individual trait over a period of nonstationary dynamics. By following individual fates and body sizes at each life-history stage, we investigated how body size and population density influenced demographic rates. By comparing the ability of two alternative approaches, a matrix projection model and an integral projection model, we investigated whether consideration of trait-based demography enhances our ability to predict transient dynamics. By utilizing a prospective perturbation analysis, we addressed which stage-specific demographic or trait-transition rate had the greatest influence on population dynamics. Both body size and population density had important effects on most rates; however, these effects differed substantially among life-history stages. Considering the observed trait-demography relationships resulted in better predictions of a population's response to perturbations, which highlights the role of phenotypic plasticity in transient dynamics. Although the perturbation analyses provided comparable predictions of stage-specific elasticities between the matrix and integral projection models, the order of importance of the life-history stages differed between the two analyses. In conclusion, we demonstrate how a trait-based demographic approach provides further insight into transient population dynamics.     

Temperature change and complex dynamics

Density-dependent factors, such as population growth rate and migration, influence dynamic behaviour in ecological models. Temperature, an abiotic and density-independent factor, is also an important determinant of insect population growth. We investigated the endogenous dynamics of a density-dependent response-surface model that included temperature, based on time series for two aphid species. We investigated the effects of temperature and random noise on the model dynamics. In most cases, an increase in temperature resulted in a higher predicted equilibrium density; it could induce complex dynamics. Noise at the level of the natural variation in temperature resulted in extinctions in some models. Our results from these models indicate that aphid populations might become more abundant, and less stable in some circumstances, if there is climate warming. Received: 25 November 1996 / Accepted: 30 June 1997     

The generality of management recommendations across populations of an invasive perennial herb

Demographic models are widely used to produce management recommendations for different species. For invasive plants, current management recommendations to control local population growth are often based on data from a limited number of populations per species, and the assumption of stable population structure (asymptotic dynamics). However, spatial variation in population dynamics and deviation from a stable structure may affect these recommendations, calling into question their generality across populations of an invasive species. Here, I focused on intraspecific variation in population dynamics and investigated management recommendations generated by demographic models across 37 populations of a short-lived, invasive perennial herb (Lupinus polyphyllus). Models that relied on the proportional perturbations of vital rates (asymptotic elasticities) indicated an essential role for plant survival in long-term population dynamics. The rank order of elasticities for different vital rates (survival, growth, retrogression, fecundity) varied little among the 37 study populations regardless of population status (increasing or declining asymptotically). Summed elasticities for fecundity increased, while summed elasticities for survival decreased with increasing long-term population growth rate. Transient dynamics differed from asymptotic dynamics, but were qualitatively similar among populations, that is, depending on the initial size structure, populations tended to either increase or decline in density more rapidly than predicted by asymptotic growth rate. These findings indicate that although populations are likely to exhibit transient dynamics, management recommendations based on asymptotic elasticities for vital rates might be to some extent generalised across established populations of a given short-lived invasive plant species.     

Transient Responses to Spatial Perturbations in Advective Systems

We study the transient dynamics, following a spatially-extended perturbation of models describing populations residing in advective media such as streams and rivers. Our analyses emphasize metrics that are independent of initial perturbations—resilience, reactivity, and the amplification envelope—and relate them to component spatial wavelengths of the perturbation using spatial Fourier transforms of the state variables. This approach offers a powerful way of understanding the influence of spatial scale on the initial dynamics of a population following a spatially variable environmental perturbation, an important property in determining the ecological implications of transient dynamics in advective systems. We find that asymptotically stable systems may exhibit transient amplification of perturbations (i.e., have positive reactivity) for some spatial wavelengths and not others. Furthermore, the degree and duration of amplification varies strongly with spatial wavelength. For two single-population models, there is a relationship between transient dynamics and the response length that characterizes the steady state response to spatial perturbations: a long response length implies that peak amplification of perturbations is small and occurs fast. This relationship holds less generally in a specialist consumer-resource model, likely due to the model’s tendency for flow-induced instabilities at an alternative characteristic spatial scale.     

Brachionus plicatilis tolerance to low oxygen concentrations

Tolerance to low oxygen concentrations is expected in Brachionus plicatilis, a rotifer adapted to live in saline warm waters. The population dynamics of a clone of this species, isolated from an endorreic saline lake, was studied under controlled laboratory conditions. Although their growth and metabolism is extremely reduced, B. plicatilis populations are able to maintain relatively high-density populations (a mean of 35 ind ml^–1) in oxygen concentrations below 1 mg 1^–1, for more than one month. Major features of population growth related to oxygen are discussed.     

斜生栅藻密度对新安江萼花臂尾轮虫种群动态的影响

以浓度分别为1.0×106、2.0×106、4.0×106和8.0×106cells·mL-1的斜生栅藻(Scenedesmus obliquus)为轮虫食物,在25℃下,应用群体累积培养和单个体培养法研究了食物浓度对采自新安江水域(屯溪段)的萼花臂尾轮虫的种群动态影响。结果表明:食物浓度对轮虫的种群增长率和休眠卵产量均有显著的影响,均表现为低食物浓度下(≤4.0×106cells·mL-1)萼花臂尾轮虫的种群增长率和休眠卵产量较小且无差异,高食物浓度下(8.0×106cells·mL-1)轮虫的种群增长率和休眠卵产量较大;食物浓度对轮虫的混交雌体百分率和受精率无显著影响;在食物浓度为4.0×106cells·mL-1时萼花臂尾轮虫的净生殖率最大,世代时间最长,而其内禀增长率在食物浓度为1.0×106cells·mL-1时最小;在食物浓度为8.0×106cells·mL-1时平均寿命和生命期望最长。     

Temporal genetic stability in natural populations of the waterflea Daphnia magna in response to strong selection pressure

Studies monitoring changes in genetic diversity and composition through time allow a unique understanding of evolutionary dynamics and persistence of natural populations. However, such studies are often limited to species with short generation times that can be propagated in the laboratory or few exceptional cases in the wild. Species that produce dormant stages provide powerful models for the reconstruction of evolutionary dynamics in the natural environment. A remaining open question is to what extent dormant egg banks are an unbiased representation of populations and hence of the species’ evolutionary potential, especially in the presence of strong environmental selection. We address this key question using the water flea Daphnia magna, which produces dormant stages that accumulate in biological archives over time. We assess temporal genetic stability in three biological archives, previously used in resurrection ecology studies showing adaptive evolutionary responses to rapid environmental change. We show that neutral genetic diversity does not decline with the age of the population and it is maintained in the presence of strong selection. In addition, by comparing temporal genetic stability in hatched and unhatched populations from the same biological archive, we show that dormant egg banks can be consulted to obtain a reliable measure of genetic diversity over time, at least in the multidecadal time frame studied here. The stability of neutral genetic diversity through time is likely mediated by the buffering effect of the resting egg bank.     

Estimation of growth regulation in natural populations by extended family of growth curve models with fractional order derivative: Case studies from the global population dynamics database

Estimating the trend in population time series data using growth curve models is a central idea in population ecology. Several models, mainly governed by differential or difference equations, have been applied to real data sets to identify general growth pattern and make predictions. In this article, we analyze ecological time series data by fitting mathematical models governed by fractional differential equations (FDE). The order of the FDE (α) is used to quantify the evidence of memory in the population processes. The application of FDE is exemplified by analyzing time series data on two bird species Phalacrocorax carbo (Great cormorant) and Parus bicolor (Tufted titmouse) and two mammal species Castor canadensis (Beaver) and Ursus americanus (American black bear) extracted from the global population dynamics database. Five different population growth models were fitted to these data; density-independent exponential, negative density-dependent logistic and θ-logistic model, positive density-dependent exponential Allee and strong Allee model. Both ordinary and fractional derivative representations of these models were fitted to the time series data. Markov chain Monte Carlo (MCMC) method was used to estimate the model parameters and Akaike information criterion was used to select the best model. By estimating the return rate for each of the time series, we have shown that populations governed by FDE with a small value of α (high level of memory) return to the stable equilibrium faster. This demonstrates a synergistic interplay between memory and stability in natural populations.     

Divergent density feedback control of migratory predator recovery following sex‐biased perturbations

Uncertainty in risks posed by emerging stressors such as synthetic hormones impedes conservation efforts for threatened vertebrate populations. Synthetic hormones often induce sex‐biased perturbations in exposed animals by disrupting gonad development and early life‐history stage transitions, potentially diminishing per capita reproductive output of depleted populations and, in turn, being manifest as Allee effects. We use a spatially explicit biophysical model to evaluate how sex‐biased perturbation in life‐history traits of individuals (maternal investment in egg production and male‐skewed sex allocation in offspring) modulates density feedback control of year‐class strength and recovery trajectories of a long‐lived, migratory fish—shovelnose sturgeon (Scaphirhynchus platorynchus)—under spatially and temporally dynamic synthetic androgen exposure and habitat conditions. Simulations show that reduced efficiency of maternal investment in gonad development prolonged maturation time, increased the probability of skipped spawning, and, in turn, shrunk spawner abundance, weakening year‐class strength. However, positive density feedback disappeared (no Allee effect) once the exposure ceased. By contrast, responses to the demographic perturbation manifested as strong positive density feedback; an abrupt shift in year‐class strength and spawner abundance followed after more than two decades owing to persistent negative population growth (a strong Allee effect), reaching an alternative state without any sign of recovery. When combined with the energetic perturbation, positive density feedback of the demographic perturbation was dampened as extended maturation time reduced the frequency of producing male‐biased offspring, allowing the population to maintain positive growth rate (a weak Allee effect) and gradually recover. The emergent patterns in long‐term population projections illustrate that sex‐biased perturbation in life‐history traits can interactively regulate the strength of density feedback in depleted populations such as Scaphirhynchus sturgeon to further diminish reproductive capacity and abundance, posing increasingly greater conservation challenges in chemically altered riverscapes.