Simulating density-dependent relationships in south Texas northern bobwhite populations

Density dependence influences northern bobwhite (Colinus virginianus) reproduction and overwinter mortality. However, the functional forms of these density-dependent relationships or the factors that influence them during the annual life cycle events of this bird are not clear. We used a systems analysis approach with a compartment model based on difference equations (Δt = 3 months) for bobwhites in South Texas to simulate population behavior using 16 different functional forms of density-dependent production and overwinter mortality. During the reproductive season, a weak linear density-dependent relationship resulted in the longest population persistence (up to 100.0 yr), whereas a reverse-sigmoid density-dependent relationship had the worst population persistence (2.5–3.5 yr). Regarding overwinter mortality, a sigmoid or weak linear density-dependent relationship and a weak linear or no density-dependent reproduction relationship had the longest population persistence (87.5–100.0 yr). Weak linear density-dependent reproduction with either sigmoid or weak linear overwinter mortality produced stable fall population trends. Our results indicated that density dependence may have a greater influence on overwinter survival of bobwhites than previously thought. Inclusion of density-dependent functional relationships that represent both density-dependent reproduction and overwinter mortality, were critical for our simulation model to function properly. Therefore, integrating density-dependent relationships for both reproductive and overwinter periods of the annual cycle of bobwhite life history events is essential for conducting realistic bobwhite population simulation analyses that can be used to test different management scenarios in an integrated and interdisciplinary manner. © 2012 The Wildlife Society.     

Reproductive asynchrony increases with environmental disturbance

While it is widely recognized that the manner in which organisms adjust their timing of reproduction reflects evolutionary strategies aimed at minimizing offspring mortality or maximizing reproductive output, the conditions under which the evolutionarily stable strategy involves synchronous or asynchronous reproduction is a matter of considerable discord. A recent theoretical model predicts that whether a population displays reproductive synchrony or asynchrony will depend on the relative scales of intrinsic regulation and environmental disturbance experienced by reproducing individuals. This model predicts that, under conditions of negligible competition and large-scale environmental perturbation, evolution of a single mixed strategy will result in asynchronous reproduction. We tested this prediction using empirical data on large-scale climatic fluctuation and the annual timing of reproduction by three species of flowering plants covering 1300-population-years and four degrees of latitude in Norway. In agreement with model predictions, within populations of all three species reproductive asynchrony increased with the magnitude of large-scale climatic perturbation, but bore no relation to the strength of local density dependence. These results suggest that mixed evolutionarily stable strategies can arise from the interplay of combinations of agents of selection and the scale at which they operate; hence it is fruitless to associate synchronous versus asynchronous timing with particular single factors like climate, competition, or predation.     

Delayed density-dependent season length alone can lead to rodent population cycles

Studies of cyclic microtine populations (voles and lemmings) have suggested a relationship between the previous year's population density and the subsequent timing of the onset of reproduction by overwintered breeding females. No studies have explored the importance of this relationship in the generation of population cycles. Here we mathematically examine the implications of variation in reproductive season length caused by delayed density-dependent changes in its start date. We demonstrate that when reproductive season length is a function of past population densities, it is possible to get realistic population cycles without invoking any changes in birth rates or survival. When parameterized for field voles (Microtus agrestis) in Kielder Forest (northern England), our most realistic model predicts population cycles of similar periodicity to the Kielder populations. Our study highlights the potential importance of density-dependent reproductive timing in microtine population cycles and calls for investigations into the mechanism(s) underlying this phenomenon.     

Coadaptation of prenatal and postnatal maternal effects

In a wide variety of species, a female's age of first reproduction influences offspring size and survival, suggesting that there exists an optimal timing of reproduction. Mothers in many species also influence offspring size and survival after birth through variation in parental care. We experimentally separated these effects in the burying beetle Nicrophorus vespilloides to test for coadaptation between prenatal and postnatal maternal effects associated with age at first reproduction. Females that reproduced early produced offspring with lower birth weight. The amount of parental care depended on the age of first reproduction of the caretaker, as did the extent of offspring begging. As predicted for a coadaptation of maternal effects, prenatal and postnatal effects were opposite for different-aged mothers, and larval weight gain and survival was greatest when the age of the caretaker and birth mother matched. Thus, prenatal effects intrinsically associated with age of first reproduction can be ameliorated by innate plasticity in postnatal care. A coadaptation of prenatal and postnatal maternal effects may evolve to allow variable timing of the first reproductive attempt. Such a coadaptation might be particularly valuable when females are constrained from reproducing at an optimal age, as, for example, in species that breed on scarce and unpredictable resources.     

Infanticide in the evolution of reproductive synchrony: effects on reproductive success

Synchronous breeding in animals and plants has stimulated both a theoretical and empirical examination of the possible benefits of active synchronization. The selective pressures of predation and infanticide are the strongest candidates proposed to explain the evolution of reproductive synchrony. Alternatively, breeding asynchronously with conspecifics may ensure a greater availability of resources per breeder. However, the possible fitness benefits resulting from active asynchronization have not yet received attention in evolutionary ecology. Here we present a hypothesis, based on a graphical model, illustrating the costs and benefits of the two modes of reproduction. We tested the hypothesis empirically using a 2 x 2 full factorial study design, where reproductive synchrony and infanticide tactics were manipulated in enclosed populations of the bank vole. The results reveal a relationship between infanticide tactics and breeding synchrony as illustrated by our hypothesis. In general, female reproductive success (number and size of offspring surviving to weaning) was significantly lower in infanticidal populations. Moreover, an asynchronous breeding pattern proved to be advantageous in the noninfanticidal population but this advantage of asynchrony was lost as infanticide became common in the population. Our findings support the idea that synchronous reproduction could have evolved as a counterstrategy against infanticide.     

Timing of births and juvenile mortality in the South American fur seal in Peru

In most species, synchronous, seasonal reproduction is usually associated with higher offspring survival in animals giving birth around the peak, relative to those breeding at the extremes of the reproductive season. In contrast, in the South American fur seal ( Arctocephalus australis ) at Punta San Juan, Peru, females pupping around the peak of births had a greater probability of losing their pups and/or tended to lose them at an earlier age than females breeding early or late in the season. However, the timing of breeding in this population varies little between years and is consistently synchronous. Individual females maintained their relative breeding times in consecutive years, regardless of whether or not they lost their pup the previous year. Thus, pup mortality seems to have no effect on the timing of reproduction in this population.
High breeding densities and the consequent high pup mortality in the S. American fur seal in Peru may have resulted from intense poaching outside protected areas and are of recent origin. The reproductive synchrony in this population could have originally evolved as a response to seasonal variations in food availability and weather conditions, differences in female or pup body condition, predation pressure, sexual selection and/or harassment avoidance, but at the present high density levels has become maladaptive.     

Optimal individual growth and reproduction in perennial species with indeterminate growth

Summary A model predicting optimal timing of growth and reproduction in perennial species with indeterminate growth living in a seasonal environment, is presented. According to the model, the optimal fraction of growing season devoted to growth decreases with increasing individual age and size, which leads to S-shaped growth curves. Winter mortality seems to be a crucial factor affecting the timing of growth and reproduction, under the same function describing the dependence of growth rate and reproductive rate on body size. When winter mortality is heavy, it is often optimal to start reproducing in the first year, and to devote a large proportion of the subsequent years to reproduction, thus leading to small adult body sizes.The model has been applied to two species of mollusc and one species of fish. The model predictions fit well to the field data for these three species.     

Evolutionary responses to harvesting in ungulates

1. We investigate the evolutionary responses to harvesting in ungulates using a state-dependent, stochastic, density-dependent individual-based model of red deer Cervus elaphus (L.) females subject to different harvesting regimes. 2. The population's mean weight at first reproduction shifts towards light weights as harvesting increases, and its distribution changes from a single peak distribution under very low or high harvest rates, to a bimodal distribution under intermediate harvest rates. 3. These results suggest that, consistent with previous studies on aquatic species, harvesting-induced mortality may drive adaptive responses in ungulates by reducing the fitness benefits from adult survival and growth in favour of early and lightweight reproduction. 4. Selective harvesting for heavy animals has no additional effect on the evolutionarily stable strategy, suggesting that harvest rate is more important than the degree of selectivity in driving adaptive responses. However, selective harvesting of light females is positively associated with maturation weights even higher than those of a nonharvested population, probably due to the reduction in the fitness value of the offspring. 5. The average number of weight at maturation strategies in the population declines but the total number of strategies across all simulations increases with harvest rate, suggesting that harvesting-induced selection on weight at maturity overcomes the increase in strategy diversity expected from density-dependent release. 6. Yield initially increases with harvesting due to enhanced productivity of light females experiencing density-dependent release. However, it crashes under intense harvesting resulting in a population skewed to light, young and, therefore, less reproductive animals.     

Spatial and temporal patterns in maternal energetic traits of yellow perch (Perca flavescens) in Lake Erie

1. For many fish species, survival during early life stages is linked to the size and energetic condition of females prior to reproduction. For example, females in good energetic condition are often more fecund and produce larger eggs and offspring than those in poor condition. 2. We measured the characteristics of female yellow perch (Perca flavescens) that may influence annual population fluctuations. From 2005 to 2007, we measured spatial variation in female reproductive traits, such as age, length, mass and energy density (J g^?1) of somatic tissues and ovaries among four spawning aggregations of yellow perch in western and central Lake Erie. 3. Maternal traits, such as somatic energy density and spawner age distribution, differed between the western and central basin, whereas reproductive traits, such as fecundity and ovarian energy density, differed across years. 4. To understand the implications of observed differences in demographic rates (growth and mortality rates) between basins, we developed a deterministic model to simulate the total egg production in the western and central basins under different scenarios of fishing mortality. 5. High growth rates and low mortality rates combined to produce higher modelled estimates of total egg production in the central than in the western basin, and a larger proportion of eggs were produced by old age classes in the central basin than in the western basin. 6. Our results demonstrate that changing harvest levels for populations with different demographic rates can influence total reproductive output through complex interactions between age‐specific mortality, growth and size‐specific fecundity, which has implications for the population dynamics of yellow perch and related species across a broad geographic range.     

Optimal resource allocation to seeds and vegetative propagules under density-dependent regulation in Syneilesis palmata (Compositae)

Syneilesis palmata reproduces by both seeds and vegetative propagules (short rhizomes). The latter result in the production of new plants that are larger in size and hence have a higher survival probability and a higher growth rate than seeds. A previous study predicted that the optimal reproductive strategy, in terms of maximizing population growth rate (a fitness measure under no density regulations), was pure vegetative reproduction. However, high resource investment to vegetative propagules can cause local crowding resulting in reduced demographic performances of the plants, because the vegetative propagules of Syneilesis are produced close to one another. We examined, in this situation, the impact of allocating a certain proportion of reproductive resource to seeds with relatively greater capacity for dispersal. We simulated dynamics of hypothetical Syneilesis populations with various reproductive resource allocation balances (from pure seed to pure vegetative reproduction), using a density-dependent matrix model. In the model, it was assumed that plants from vegetative propagules experienced density-dependent reduction in their survival probabilities, but this was not the case for plants originating from seeds. Each allocation strategy was evaluated based on an equilibrium population density, a fitness measure under density-dependent regulations. The optimal reproductive strategy predicted was pure vegetative reproduction. Unrealistic conditions were required for seed reproduction to be favoured, such as the production of seeds one hundred times the normal number per unit resource investment. However, the conditions were fairly relaxed compared with those required in the model where no density effects were incorporated. This indicates that escape from local crowding is likely to be one of the roles of seed production in Syneilesis.     

Warmer springs lead to mistimed reproduction in great tits (Parus major)

In seasonal environments, the main selection pressure on the timing of reproduction (the ultimate factor) is synchrony between offspring requirements and food availability. However, reproduction is initiated much earlier than the time of maximum food requirement of the offspring. Individuals should therefore start reproduction in response to cues (the proximate factors), available in the environment of reproductive decision making, which predict the later environment of selection. With increasing spring temperatures over the past decades, vegetation phenology has advanced, with a concomitant advancement in the reproduction of some species at higher trophic levels. However, a mismatch between food abundance and offspring needs may occur if changes in the environment of decision making do not match those in the environment of selection. Date of egg laying in a great tit (Parus major) population has not advanced over a 23-year period, but selection for early laying has intensified. We believe that this is the first documented case of an adaptive response being hampered because a changing abiotic factor affects the environment in which a reproductive decision is made differently from the environment in which selection occurs.     

Backward Bifurcation and Optimal Control in Transmission Dynamics of West Nile Virus

The paper considers a deterministic model for the transmission dynamics of West Nile virus (WNV) in the mosquito-bird-human zoonotic cycle. The model, which incorporates density-dependent contact rates between the mosquito population and the hosts (birds and humans), is rigorously analyzed using dynamical systems techniques and theories. These analyses reveal the existence of the phenomenon of backward bifurcation (where the stable disease-free equilibrium of the model co-exists with a stable endemic equilibrium when the reproduction number of the disease is less than unity) in WNV transmission dynamics. The epidemiological consequence of backward bifurcation is that the classical requirement of having the reproduction number less than unity, while necessary, is no longer sufficient for WNV elimination from the population. It is further shown that the model with constant contact rates can also exhibit this phenomenon if the WNV-induced mortality in the avian population is high enough. The model is extended to assess the impact of some anti-WNV control measures, by re-formulating the model as an optimal control problem with density-dependent demographic parameters. This entails the use of two control functions, one for mosquito-reduction strategies and the other for personal (human) protection, and redefining the demographic parameters as density-dependent rates. Appropriate optimal control methods are used to characterize the optimal levels of the two controls. Numerical simulations of the optimal control problem, using a set of reasonable parameter values, suggest that mosquito reduction controls should be emphasized ahead of personal protection measures.     

茉莉酸甲酯诱导大戟三萜类代谢的研究

京大戟是多年生草本药用植物,入药部分是其干燥根,但可入药的京大戟资源由于生长缓慢以及环境污染的加剧而越发匮乏,因此解决大戟资源日益紧张的问题是当今药用植物资源开发与利用方向的重要课题。京大戟含有三萜类、二萜类、黄酮类等丰富的活性成分,一些常见药用植物的有效成分是三萜类化合物,其在抗病毒、抗肿瘤、免疫调节等方面具有很好的活性。对植物萜类物质代谢起重要作用的关键酶,如3-羟基,3-甲基戊二酰辅酶A还原酶(hmgr)、鲨烯合酶(sqs)、法尼基焦磷酸合酶(fps)的基因克隆及活性研究取得了进展和突破,但通过调控萜类物质代谢途径中关键酶基因的表达来诱导终产物合成的研究鲜有报道。通过研究大戟萜类物质代谢途径进而利用基因工程手段提升目的物质的产量来解决京大戟药源短缺问题具有重要意义。该研究以大戟愈伤组织为材料,使用茉莉酸甲酯分别按时间梯度和浓度梯度进行诱导,将诱导后的愈伤组织分为两部分:一部分提取其总RNA,以actin为内参基因进行反转录,实时定量RT-PCR分析大戟三萜类代谢途径中hmgr、sqs与fps基因的相对表达差异;另一部分用于提取其总三萜并使用分光光度法进行含量测定。实时定量RT-PCR分析结果表明,茉莉酸甲酯可诱导3个基因的表达,但其表达模式不一样。相应的京大戟愈伤组织中总三萜的含量明显提高,最高可较未处理样品增加27%。研究结果可为茉莉酸甲酯促进药用植物大戟三萜类物质积累的分子机制研究提供参考。     

Size-dependent resource allocation among vegetative propagules and male and female functions in the forest herb Laportea bulbifera

Reproductive resource investment among vegetative propagules and male and female sexual function and their size-dependence were investigated in a perennial forest herb, Laportea bulbifera . A theoretical model based on fitness gain curves predicts that optimal investments in three reproductive modes will increase with plant size if fitness returns in all three modes increase but become saturated with investment. In a field population, large plants of L. bulbifera produced both male and female inflorescences with propagules, while small plants produced only vegetative propagules. Biomass of propagules, male inflorescences, and infructescences with achenes were all positively correlated with plant size. The increase in investment with plant size was larger for propagule production than for sexual reproduction. The relationship between propagule biomass and plant size was constant irrespective of year, while the relationship between the biomass of sexual reproductive organs and plant size differed between two successive years. Annual change of individual sex expression was investigated for 25 transplanted plants. Although each plant changed its sex expression variously among male, female and bisexual from year to year, 23 out of 25 plants produced both male and female inflorescences in at least one year. The number of viable (germinated and survived) offspring from seeds was not significantly different from the number from propagules. The production cost of a propagule was higher than that of a seed. Resource allocation theory does not seem to be applicable to size-dependent resource allocation, especially the allocation between seeds and propagules in this species.     

Optimal rates of bisexual reproduction in cyclical parthenogens with density-dependent growth

This work explores theoretical patterns of reproduction that maximize the production of resting eggs and the long-term fitness of genotypes in cyclical parthenogens. Our focus is on density-dependent reproduction as it influences the consequences of a trade-off between producing amictic daughters – which reproduce parthenogenetically and subitaneously – and producing mictic daughters – which undergo meiosis and bisexual reproduction. Amictic females increase competitive ability and allow the population to achieve a larger size; mictic females directly contribute to population survival through harsh periods by producing resting eggs. Although morphologically indistinguishable, the two types of females differ greatly in their ecological and reproductive roles. What factors underlie the differential allocation of resources to produce amictic and mictic females? Using a demographic model based on readily accessible parameters we demonstrate the existence of a frequency of mictic females that will maximize the population's long-term fitness. This frequency, termed the optimal mictic ratio, m_o, is 1 ? (q/b)^1/2, where q is the mortality rate and b is the maximum birth rate. Using computer simulation we compared the fitness of a population with this constant mictic ratio with populations having multiple switches from complete parthenogenetic growth to complete allocation in mixis (mictic ratio either 0 or 1). Two important conclusions for optimal mixis in density-dependent growth conditions are: (1) intermediate mictic ratios are optimal, and (2) optimal mictic ratios are higher when habitat conditions are better. Physiological cues responding to differences in birth and death rates are common so that it is possible that populations may adjust their relative rates of mictic and amictic female production in response to environmentally induced changes to the optimum mictic ratio. Our analysis demonstrates that different patterns of mixis are expected in different type of habitats. Since the optimal mictic ratio is sensitive to the effects of a variety of environmental challenges, our model makes possible a new means to evaluate life history evolution in cyclical parthenogens.     

Determinants of reproductive success in female adders,Vipera berus

Summary Female lifetime reproductive success in a small population of individually-marked adders in southern Sweden was studied over a period of seven years. Reproductive characteristics varied little from year to year and were consistent through time in individual females. Most females mature at four years of age and reproduce every two years. The total number of offspring produced by a female depends on her adult body size (and thus, litter size) and longevity (and thus, number of litters per lifetime). Adult body size in females is influenced mainly by subadult growth rates. Offspring size depends on maternal body size and a tradeoff between offspring size and offspring number. Maternal age does not affect litter sizes and offspring sizes except through ontogenetic changes in maternal body size.Survival of females after parturition is low because of the high energy costs of reproduction, compounded by low feeding rates of gravid females because of their sedentary behaviour at this time. About one-half of females produce only a single litter during their lifetimes, although some females live to produce four or five litters. On a proximate basis, rates of energy accumulation for growth (in subadults) and reproduction (in adults) may be the most important determinants of fitness in female adders.     

Intergenomic epistasis causes asynchronous hatch times in whitefish hybrids,but only when parental ecotypes differ

Support for the theory of ecological speciation requires evidence for ecological divergence between species which directly or indirectly causes reproductive isolation. This study investigates effects of ecological vs. genetic disparity of parental species on the presence of endogenous selection (deformation and mortality rates) and potential sources of exogenous selection (growth rates and hatch timing) on hybrids. Hybrid embryonic development is analysed in a common‐garden full‐sib cross of three species belonging to two different ecotypes within the Coregonus lavaretus species flock in the central Alpine region of Europe. Although hatch timing was similar across the three species, embryonic growth rates and egg sizes differed between ecotypes. This led to a mismatch between embryonic growth rate and egg size in hybrid crosses that reveals epistasis between the maternal and embryonic genomes and transgressive hatch times that were asynchronous with control crosses. A strong constraint of egg size to embryo size at late development was also evident. We argue that this demonstrates potential for coadaptation of a maternal trait (egg size) with offspring growth rate to be an important source of selection against hybridization between ecotypes with different egg sizes. Implications for the measurement and quantification of early life‐history traits affected by this additive relationship, such as hatch day and larval size, are also discussed.     

An individual-based population model for the prediction of rotifer population dynamics and resting egg production

Most species of rotifers have a combination of sexual and asexual reproduction, with sexual reproduction resulting in resting eggs, which can lay dormant for long periods. The occurrence of sexual reproduction affects population dynamics through the temporary presence of male rotifers, and a reduction in the growth of the number of female rotifers. A previously published, individual-based model used dynamic energy budget theory to describe rotifer food intake, growth, egg production, and mortality, but assumed asexual reproduction only. In the current study, we have expanded the model to describe the entire reproductive cycle of the rotifers, making it usable for investigating relationships, such as those between the signal triggering mictic egg production, and the timing and number of resting eggs produced. The model is intended for use in predicting the specific future development of cultures, for instance, as a process model in rotifer or resting egg production for aquaculture. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer Research     

Evolution as a critical component of plankton dynamics

Microevolution is typically ignored as a factor directly affecting ongoing population dynamics. We show here that density-dependent natural selection has a direct and measurable effect on a planktonic predator-prey interaction. We kept populations of Brachionus calyciflorus, a monogonont rotifer that exhibits cyclical parthenogenesis, in continuous flow-through cultures (chemostats) for more than 900 days. Initially, females frequently produced male offspring, especially at high population densities. We observed rapid evolution, however, towards low propensity to reproduce sexually, and by 750 days, reproduction had become entirely asexual. There was strong selection favouring asexual reproduction because, under the turbulent chemostat regime, males were unable to mate with females, produced no offspring, and so had zero fitness. In replicated chemostat experiments we found that this evolutionary process directly influenced the population dynamics. We observed very specific but reproducible plankton dynamics which are explained well by a mathematical model that explicitly includes evolution. This model accounts for both asexual and sexual reproduction and treats the propensity to reproduce sexually as a quantitative trait under selection. We suggest that a similar amalgam of ecological and evolutionary mechanisms may drive the dynamics of rapidly reproducing organisms in the wild.     

Asynchrony between solitary bee emergence and flower availability reduces flower visitation rate and may affect offspring size

Climate change can disrupt plant-pollinator interactions when shifts in the timing of pollinator activity and flowering occur unequally (i.e., phenological asynchrony). Phenological asynchrony between spring-emerging solitary bees and spring-flowering plants may cause bees to experience food deprivation that can affect their reproductive success. However, the mechanisms underlying the effects of food deprivation on solitary bee reproduction remain unknown. We investigated 1) whether food deprivation caused by phenological asynchrony affects solitary bee reproduction by influencing female lifespan and/or visitation to flowers, and 2) the relationship between the magnitude of asynchrony and bee responses. We simulated phenological asynchrony by depriving emerged female Osmia cornifrons (a spring-active solitary bee species) of nectar and pollen for 0 to 16 days. Following asynchrony treatments, we used flight cages to monitor 1) post-treatment female lifespan, 2) flower visitation, and 3) reproduction (i.e., total offspring, offspring weight, sex ratio). We found that post-treatment female lifespan was not affected by phenological asynchrony treatments, but that flower visitation rate and offspring weight decreased as the magnitude of asynchrony increased. Due to low offspring production and a lack of female offspring across treatments, we were unable to assess the effects of phenological asynchrony on total offspring produced or sex ratio. Findings suggest that post-emergence food deprivation caused by phenological asynchrony may affect offspring size by influencing nest-provisioning rates. In solitary bees, body size influences wintering survival, fecundity, and mating success. Thus, phenological asynchrony may have consequences for solitary bee populations that stem from reduced flower visitation rates, and these consequences may increase as the magnitude of asynchrony increases. Because many wild flowering plants and crops rely on pollination services provided by bees for reproductive success, bee responses to phenological asynchrony may also affect wild plant biodiversity and crop yields.