The relative importance of sexual and clonal reproduction for population growth in the perennial herb Fragaria vesca

The relative importance of sexual and clonal reproduction for population growth in clonal plants is highly variable. Clonal reproduction is often more important than sexual reproduction but there is considerable interspecific variation and the importance of the two reproductive modes can change with environmental conditions. We carried out a demographic study on the woodland strawberry (Fragaria vesca), a widespread clonal herb, at 12 sites in Switzerland during 2 years. Study sites were selected in two different habitats, i.e., forest and forest edge. We used periodic matrix models to estimate annual population growth rates and carried out prospective analyses to identify life cycle components that influence population growth rates most. Retrospective analyses were applied to study how the two different habitats affected population dynamics. Furthermore, we tested whether trade-offs between sexual and clonal reproduction occurred. There were large differences in annual population growth rates between sites and large within-site differences between years. Results of the prospective analyses clearly indicate that clonal reproduction is the dominant reproductive pathway whereas sexual reproduction is rather insignificant for population growth. Compared to forest habitats, forest edge habitats had higher population growth rates in the first year but smaller growth rates in the second year. We attribute these differing habitat effects to different water availabilities during consecutive years. No trade-offs between sexual and clonal reproduction were found. In conclusion, population growth of F. vesca relies heavily on clonal reproduction. Furthermore, reproduction and survival rates of F. vesca depend highly on spatio-temporal variation of environmental conditions.     

Bridging the gap between theory and data in ecological models

In September 2011, the University of Essex, UK, hosted an interdisciplinary conference, Mathematical and Theoretical Ecology 2011 (MATE 2011), with the theme of ‘Linking models with ecological processes’. The aim of the meeting was to create discussion and debate between modellers and empiricists working in ecology. A wide range of topics were discussed at the meeting including evolutionary and community models of ecosystem structure, epidemiological models, non-linear models of population dynamics, spatiotemporal models, individual and collective movement behaviour, and applications of ecological models to engineering problems. In this introductory article, we provide a report of the MATE 2011 meeting, and briefly review the most recent relevant research in the fields of mathematical and theoretical ecology. We introduce and summarise the eight contributed articles that were selected for this special issue. The diverse range of topics and the wide range of mathematical, statistical and computational tools used illustrate the broad appeal and depth of research in the rich field of mathematical and theoretical ecology.     

Stochastic density dependence in population size of a benthic clonal invertebrate: the regulating role of fission

The influence of environmental variation on the demography of clonal organisms has been poorly studied. I utilise a matrix model of the population dynamics of the intertidal zoanthid Palythoa caesia to examine how density dependence and temporal variation in demographic rates interact in regulating population size. The model produces realistic simulations of population size, with erratic fluctuations between soft lower and upper boundaries of approximately 55 and 90% cover. Cover never exceeds the maximum possible of 100%, and the population never goes to extinction. A sensitivity analysis indicates that the model’s behaviour is driven by density dependence in the fission of large colonies to produce intermediate sized colonies. Importantly, there is no density-dependent mortality in the model, and density dependence in recruitment, while present, is unimportant. Thus it appears that the main demographic processes which are considered to regulate population size in aclonal organisms may not be important for clonal species. Received: 18 August 1999 / Accepted: 29 October 1999     

大沙鼠种群年龄结构的季节变化和繁殖特征

2003~2004年在内蒙古包头市达茂联合旗腾格淖尔地区对大沙鼠(Rhombomys opimus)年龄组成的季节变化及其繁殖特征进行了研究。结果表明,大沙鼠的年龄组成存在着明显的季节变化和年度之间的差异,在2月至次年的4月没有幼体,6月无亚成体;大沙鼠的性比无论从总体还是各个年龄组看,均存在着差异,特别到老体,这种差异更加突出。在内蒙古包头腾格淖尔地区的大沙鼠一年中部分个体参与第三胎繁殖,而且其繁殖状况存在着年际之间的差异。     

A stochastic model for the mutation-selection balance in an infinite asexual population with a genome of fixed size

A stochastic model is presented which describes the evolution of a genome of a haploid species in an infinite population. The genome is a finite set of elements. The elements are divided into different classes according to their effect on the fitness of the organism. Repeated mutations of the genome elements are permitted, in particular positive mutations are introduced. The distribution of the deleterious elements in the genome with respect to the impact on the fitness is given after the replication step. The steady state is fully described including the distribution and the fitness.     

恢复措施对退化草地羊草种群有性生殖数量特征的影响

在封育和翻耙处理的退化羊草草地上,对羊草种群有性生殖的数量特征进行了比较研究．结果表明,翻耙样地羊草种群密度下降显著,生殖蘖密度变化不显著,生殖蘖穗长、节间长、生殖生长比率、穗节数、小穗数、小花数、饱满籽实数和结实率都有显著增长,籽实产量、净籽实重和千粒重分别是封育样地的2．68、3．49和1．3倍．在翻耙样地中,羊草种群发育有复小穗的生殖蘖比封育样地多16％,生殖蘖上复小穗的数量也显著增加,生殖蘖以相对少的小穗,生产了较多的小花和更多的籽实来提高种群的籽实产量．两个样地中羊草种群的生物量生殖分配在种群对生殖蘖RA1、生殖蘖对穗序RA4、穗序对籽实RA6 3个层次上无显著差异,而种群对穗序RA2、生殖蘖对籽实RA5的生殖分配差异显著,种群对籽实RA3的生殖分配差异极显著．     

The invisible niche: Weakly density-dependent mortality and the coexistence of species

Weakly density-dependent effects, characterized by fractional scaling exponents close to one, are rarely studied in the ecological literature. Here, we consider the effect of an additional weakly density-dependent term on a simple competition model. Our investigation reveals that weak density-dependence opens up an “invisible niche”. This niche does not constitute a new mechanism for coexistence, but is a previously unexplored consequence of known mechanisms. In the invisible niche a weaker competitor can survive at very low density. Coexistence thus requires large habitat size. Such niches, if found in nature, would have a direct impact on species-area laws and species-abundance curves and should therefore receive more attention.     

Strong density-dependent survival and recruitment regulate the abundance of a coral reef fish

Debate on the control of population dynamics in reef fishes has centred on whether patterns in abundance are determined by the supply of planktonic recruits, or by post-recruitment processes. Recruitment limitation implies little or no regulation of the reef-associated population, and is supported by several experimental studies that failed to detect density dependence. Previous manipulations of population density have, however, focused on juveniles, and there have been no tests for density-dependent interactions among adult reef fishes. I tested for population regulation in Coryphopterus glaucofraenum, a small, short-lived goby that is common in the Caribbean. Adult density was manipulated on artificial reefs and adults were also monitored on reefs where they varied in density naturally. Survival of adult gobies showed a strong inverse relationship with their initial density across a realistic range of densities. Individually marked gobies, however, grew at similar rates across all densities, suggesting that density-dependent survival was not associated with depressed growth, and so may result from predation or parasitism rather than from food shortage. Like adult survival, the accumulation of new recruits on reefs was also much lower at high adult densities than at low densities. Suppression of recruitment by adults may occur because adults cause either reduced larval settlement or reduced early post-settlement survival. In summary, this study has documented a previously unrecorded regulatory mechanism for reef fish populations (density-dependent adult mortality) and provided a particularly strong example of a well-established mechanism (density-dependent recruitment). In combination, these two compensatory mechanisms have the potential to strongly regulate the abundance of this species, and rule out the control of abundance by the supply of recruits.     

On the effects of spatial heterogeneity on the persistence of interacting species

 The dynamics of two interacting theoretical populations inhabiting a heterogeneous environment are modelled by a system of two weakly coupled reaction–diffusion equations having spatially dependent reaction terms. Longterm persistence of both populations is guaranteed by an invasibility condition, which is itself expressed via the signs of certain eigenvalues of related linear elliptic operators with spatially dependent lowest order coefficients. The effects of change in these coefficients upon the eigenvalues are here exploited to study the effects of spatial heterogeneity on the persistence of interacting species through two particular ecological topics of interest. The first concerns when the location of favorable hunting grounds within the overall environment does or does not affect the success of a predator in predator–prey models, while the second concerns cases of competition models in which the outcome of competition in a spatially varying environment differs from that which would be expected in a spatially homogeneous environment. Received: 9 June 1997     

Regeneration growth of the invasive clonal forb Rorippa austriaca (Brassicaceae) in relation to fertilization and interspecific competition

A special type of clonal growth, spread by lateral roots, ishypothesized to be a favourable trait of invasive, opportunistic plant speciesof disturbed habitats. We tested this hypothesis for the invasive forbRorippa austriaca (Brassicaceae). Regenerationfrom root fragments, subsequent vegetative spread and allocation patterns inrelation to varied nutrient supply and intensity and pattern of interspecificcompetition were analyzed in container experiments. Regeneration success fromroot fragments was 100% and clonal spread was rapid but vegetativeperformance was strongly reduced under unfertilized conditions and,particularly, when interspecific competition was present. While the ratio ofabove- to belowground bio-mass did not differ considerably betweentreatments, R. austriaca allocated a high amount ofresources to belowground growth resulting in low aboveground but highbelowground biomass at harvest time relative to the matrix vegetation.Differences in shoot number or biomass between simulated gaps and denselyvegetated quadrants in the containers were (relatively) weak.Reproductive effort was less reduced under low resource levels, and the clonesdid not set seed at all, irrespective of the treatment. Our results show thatclonal growth by lateral roots and plasticity in clonal growth patterns inR. austriaca promote both exploitation of gaps andnutrient-rich microsites and resistance to competitors. Such plasticity,combined with its ability to regenerate from widely-dispersed rootfragments, contribute to the ability of the species to invade and persistwithindisturbed and spatially heterogeneous habitats.     

Evolution of new gene functions: simulation and analysis of the amplification model

Creation of new genes and functions is a central feature of evolution. Duplication of existing genes has long been assumed to be the source of new genes, but the precise mechanism has remained unclear. One suggestion is that new genes are created via temporary amplifications, which simultaneously increase both the selective advantage of weak, pre-existing secondary functions and the target for optimizing mutations. This paper examines the amplification model by formalizing it into a mathematical framework. This framework is used to perform stochastic (Monte Carlo) simulations. In addition, experimental data from Salmonella typhimurium LT2 are used to support the modelling, by providing estimates for parameter values. The results show that amplification of tandem repeats is likely to contribute to creation of new genes in nature.     

Importance of endogenous feedback controlling the long-term abundance of tropical mosquito species

Mosquitoes are a major vector for tropical diseases, so understanding aspects that modify their population dynamics is vital for their control and protecting human health. Maximising the efficiency of control strategies for reducing transmission risk requires as a first step the understanding of the intrinsic population dynamics of vectors. We fitted a set of density-dependent and density-independent models to the long-term time series of six tropical mosquito species from northern Australia. The models’ strength of evidence was assessed using Akaike’s Information Criterion (AIC _c ), Bayesian Information Criterion (BIC) and jack-knifed cross-validation (C-V). Density dependence accounted for more than 99% of the model weight in all model-selection methods, with the Gompertz-logistic (Cushing model) being the best-supported model for all mosquito species (negative density feedback expressed even at low densities). The second-most abundant species, Aedes vigilax (a saline breeder), showed no spatial heterogeneity in its density-dependent response, but the remaining five species had different intrinsic growth rates across 11 study sites. Population densities of saline species were high only during the late dry to early wet season following the highest tides of the month or early flood rains when swamps were mostly saline, whereas those of freshwater species were highest during the mid-wet and mid-dry seasons. These findings demonstrate remarkably strong density dependence in mosquito populations, but also suggest that environmental drivers, such as rainfall and tides, are important in modifying seasonal densities. Neglecting to account for strong density feedback in tropical mosquito populations will clearly result in less effective control.     

Individual optimization efforts and population dynamics: a mathematical model for the evolution of resource allocation strategies, with applications to reproductive and mating systems

We develop a formal framework for the optimal allocation of limited resources that includes and clarifies the interplay between individual optimization and the resulting effects at the population level. As an example, in regard to the evolution of sexual recombination, the paradox of the twofold cost of sex is avoided by distinguishing between the evolution of recombination and the subsequent emergence and stability of different mating types as a result of individual optimization within a population that benefits from recombination.

Effect of lethality on the extinction and on the error threshold of quasispecies

In this paper the effect of lethality on error threshold and extinction has been studied in a population of error-prone self-replicating molecules. For given lethality and a simple fitness landscape, three dynamic regimes can be obtained: quasispecies, error catastrophe, and extinction. Using a simple model in which molecules are classified as master, lethal and non-lethal mutants, it is possible to obtain the mutation rates of the transitions between the three regimes analytically. The numerical resolution of the extended model, in which molecules are classified depending on their Hamming distance to the master sequence, confirms the results obtained in the simple model and shows how an error catastrophe regime changes when lethality is taken in account.     

Effect of plants on the searching efficiency of a generalist predator: the importance of predator-prey spatial association

In most studies of tritrophic interactions, the effect of plants on predators is confounded with changes in prey and predator behaviors after an encounter event. Here, we estimate how the effect of plants on prey distribution (in the absence of the predator) and on predator foraging behavior (in the absence of prey) may influence predation rate of Orius insidiosus (Say) (Heteroptera: Anthocoridae) in 11 plant by prey species combinations. The within-leaf distributions of O. insidiosus and its prey overlapped most on bean plants. The predator's foraging behavior (e.g., walking speed, turning rate) also differed among plant species. Simulations, using the prey distribution data and predator's foraging patterns on leaf surfaces of each plant species, show that, overall, the searching efficiency of O. insidiosus was higher on leaves of bean and corn than of tomato. However, the predator's searching efficiency was not consistent within plant species. Thus, the combined effect of plants directly on the predator and indirectly through the prey influenced the predator's searching efficiency.     

Long-term population dynamics of two Carex curvula species in the Central Alps on native and alien soils

The demography of two closely related alpine sedges, Carex curvula subsp. curvula and Carex curvula subsp. rosae (=C. curvula and C. rosae) has been investigated on their typical sites in the Central Alps. Both species proliferate vegetatively and develop dense tussocks but they show different dominance behaviours in their respective grasslands. It was hypothesized that this may be caused by different growth abilities. The main aim of the study was to compare the vegetative growth of the species under field conditions, under competition-free conditions and under changed soil conditions. An attempt was also made to clarify whether vegetative growth is density dependent. Permanent plots were established in the respective grasslands of the two species and the ramet density was counted over 3 years. Groups of 10 and of 30 ramets of each species were grown in pots with typical and with alien substrate and their growth was observed for 5 years at the field site. The grassland populations of both species were very stable and the overall ramet growth rate (λ) was close to 1.0. Within the pots, both species reached a high ramet number. Only the group of 30 ramets of C. curvula on alien soil could not recover from the transplantation shock. Within the pots, C. rosae showed a greater ramet turnover and a higher increase in ramets than C. curvula. On their native substrate, both species had a significantly higher ramet increase than on the alien substrate. Ramet growth was found to be density dependent for both species, the increases recorded for the groups of 10 being significantly greater than for the groups of 30. Although C. curvula produced fewer ramets than C. rosae, the aboveground dry weight of the former was significantly higher. This may be decisive for its greater competitive success in closed grasslands. Received: 12 April 1997 / Accepted: 12 December 1997     

Simulating the evolution of a clonal trait in plants with sexual and vegetative reproduction

Aims Phenotypic optimality models neglect genetics. However, especially when heterozygous genotypes are fittest, evolving allele, genotype and phenotype frequencies may not correspond to predicted optima. This was not previously addressed for organisms with complex life histories.Methods Therefore, we modelled the evolution of a fitness-relevant trait of clonal plants, stolon internode length. We explored the likely case of an asymmetric unimodal fitness profile with three model types. In constant selection models (CSMs), which are gametic, but not spatially explicit, evolving allele frequencies in the one-locus and five-loci cases did not correspond to optimum stolon internode length predicted by the spatially explicit, but not gametic, phenotypic model. This deviation was due to the asymmetry of the fitness profile. Gametic, spatially explicit individual-based (SEIB) modeling allowed us relaxing the CSM assumptions of constant selection with exclusively sexual reproduction.Important findings For entirely vegetative or sexual reproduction, predictions of the gametic SEIB model were close to the ones of spatially explicit non-gametic phenotypic models, but for mixed modes of reproduction they approximated those of gametic, not spatially explicit CSMs. Thus, in contrast to gametic SEIB models, phenotypic models and, especially for few loci, also CSMs can be very misleading. We conclude that the evolution of traits governed by few quantitative trait loci appears hardly predictable by simple models, that genetic algorithms aiming at technical optimization may actually miss the optimum and that selection may lead to loci with smaller effects in derived compared with ancestral lines.     

Northern Atlantic Oscillation effects on the temporal and spatial dynamics of green spruce aphid populations in the UK

1. The role of climate variability in determining the spatial and temporal patterns of numerical fluctuations is a central problem in ecology. The influence of the North Atlantic Oscillation (NAO) index on the population dynamics and spatial synchrony of the green spruce aphid Elatobium abietinum across the UK was shown. 2. Fifteen overlapping time series within the UK were analysed; we used nonparametric models for determining the feedback nonlinear structure and the climatic effects. The spatial synchrony of these populations and the relationship between synchrony and NAO was estimated. 3. From the 15 time series across the UK, 11 showed positive and significant NAO effects. In most of the cases the NAO effects were nonlinear showing strong negative effects of low values. The NAO variation improve the explained variance of the first-order feedback models in 14.5%; ranging from 0% to 48%. All data showed strong-nonlinear (concave) feedback structure. In most of the localities the explained variance by the first-order feedback was about 50-60%. 4. The spatial synchrony of the per capita growth rates and residuals is high across long distances for those populations affected by NAO. The correlation function predicts a spatial scale of synchrony of about 350-400 km for NAO influenced populations. 5. We think that simple population theoretical models describing the link between NAO fluctuations and green spruce aphid dynamics may be fundamental for predicting and simulating the consequences of different climatic scenarios of the future.