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     

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.     

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.     

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.     

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.

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     

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.     

Temporal and spatial variation in the nitrite-oxidizing bacterial community of a grassland soil

Abstract The temporal and spatial distribution of the nitrite-oxidizing community of a non-fertilized, semi-natural grassland soil was studied to obtain more insight into the possible variation in nitrate production in this soil throughout the year. Data describing the size, potential nitrite-oxidizing activity and serotype composition of the nitrite-oxidizing community are reported. In addition, several abiotic soil parameters potentially related to the activity of this community were measured. Whereas numbers and potential activities largely varied with time and place, the specific affinity for nitrite oxidation, defined as the ratio V _max/ K _m, was relatively constant. The serotypes Nitrobacter agilis, N. winogradskyi and N. hamburgensis were all present in the top 5-cm soil in every 500-g sample examined, showing that these species co-exist in this soil.     

Density-dependent resistance of the gypsy moth <Emphasis Type="Italic">Lymantria dispar</Emphasis> to its nucleopolyhedrovirus,and the consequences for population dynamics

The processes controlling disease resistance can strongly influence the population dynamics of insect outbreaks. Evidence that disease resistance is density-dependent is accumulating, but the exact form of this relationship is highly variable from species to species. It has been hypothesized that insects experiencing high population densities might allocate more energy to disease resistance than those at lower densities, because they are more likely to encounter density-dependent pathogens. In contrast, the increased stress of high-density conditions might leave insects more vulnerable to disease. Both scenarios have been reported for various outbreak Lepidoptera in the literature. We tested the relationship between larval density and disease resistance with the gypsy moth (Lymantria dispar) and one of its most important density-dependent mortality factors, the nucleopolyhedrovirus (NPV) LdMNPV, in a series of bioassays. Larvae were reared in groups at different densities, fed the virus individually, and then reared individually to evaluate response to infection. In this system, resistance to the virus decreased with increasing larval density. Similarly, time to death was faster at high densities than at lower densities. Implications of density–resistance relationships for insect–pathogen population dynamics were explored in a mathematical model. In general, an inverse relationship between rearing density and disease resistance has a stabilizing effect on population dynamics.     

Mutation and control of the human immunodeficiency virus

We examine the dynamics of infection by the human immunodeficiency virus (HIV), as well as therapies that minimize viral load, restore adaptive immunity, and use minimal dosage of anti-HIV drugs. Virtual therapies for wild-type infections are demonstrated; however, the HIV infection is never cured, requiring continued treatment to keep the condition in remission. With high viral turnover and mutation rates, drug-resistant strains of HIV evolve quickly. The ability of optimal therapy to contain drug-resistant strains is shown to depend upon the relative fitness of mutant strains.     

Sex-specific facilitation and reproduction of the gynodioecious cushion plant Arenaria polytrichoides on the Himalaya-Hengduan mountains,SW ChinaOA

When benefiting other beneficiaries, cushion plants may reciprocally receive feedback effects. The feedback effects on different sex morphs, however, remains unclear. In this study, taking the gynodioecious Arenaria polytrichiodes as a model species, we aimed to assess the sex-specific facilitation intensity of cushion plant by measuring the beneficiary cover ratio, and to assess the potential costs in cushion reproductive functions by measuring the flower and fruit cover ratios. The total benef...     

Density-dependent growth and reproduction of the apple snail, Pomacea canaliculata: a density manipulation experiment in a paddy field

To examine density dependence in the survival, growth, and reproduction of Pomacea canaliculata, we conducted an experiment in which snail densities were manipulated in a paddy field. We released paint-marked snails of 15–20 mm shell height into 12 enclosures (pens) of 16 m² at one of five densities – 8, 16, 32, 64, or 128 snails per pen. The survival rate of released snails was 95% and was independent of snail density. The snail density had a significant effect on the growth and egg production of individual snails. This density dependence may have been caused by reduced food availability. The females at high density deposited fewer and smaller egg masses than those at low density, and consequently produced fewer eggs. The females at densities 8 and 16 deposited more than 3000 eggs per female, while the females at density 128 oviposited only 414 eggs. The total egg production per pen was, however, higher at higher snail density. The survival rates of juvenile snails were 21%–37% and were independent of adult density. The juvenile density was positively correlated with the total egg production per pen and hence was higher at higher adult density. However, the density of juveniles larger than 5 mm in shell height, i.e., juveniles that can survive an overwintering period, was not significantly different among density treatments. These results suggest that snail density after the overwintering period is independent of the density in the previous year. Thus, density dependence in growth and reproduction might regulate the population of P. canaliculata in paddies. Received: October 23, 1998 / Accepted: July 16, 1999     

A stochastic model for extinction and recurrence of epidemics: estimation and inference for measles outbreaks

Epidemic dynamics pose a great challenge to stochastic modelling because chance events are major determinants of the size and the timing of the outbreak. Reintroduction of the disease through contact with infected individuals from other areas is an important latent stochastic variable. In this study we model these stochastic processes to explain extinction and recurrence of epidemics observed in measles. We develop estimating functions for such a model and apply the methodology to temporal case counts of measles in 60 cities in England and Wales. In order to estimate the unobserved spatial contact process we suggest a method based on stochastic simulation and marginal densities. The estimation results show that it is possible to consider a unified model for the UK cities where the parameters depend on the city size. Stochastic realizations from the dynamic model realistically capture the transitions from an endemic cyclic pattern in large populations to irregular epidemic outbreaks in small human host populations.