Evolutionarily stable strategies of migration in heterogeneous environments

By means of a simulation model we are showing that the rates of migration can be related to avoidance of competition between relatives, especially in clonal organisms. This could result in a strong selective pressure for migration, even at a high cost. In addition, if the habitat is fragmented, migration can strongly affect local dynamics and result in a dramatic decrease of the densities in some places. In parthenogenetically reproducing organisms like aphids, the level of relatedness in local populations is expected to be very high and therefore they can serve as a good model group for testing these hypotheses. This revised version was published online in July 2006 with corrections to the Cover Date.     

Delayed female reproduction in equilibrium and chaotic populations

Behavioural and life history polymorphisms are often observed in animal populations. We analyse the timing of maturation and reproduction in risky and resource-limited environments. Field and laboratory evidence suggests that female voles and mice, for example, can adjust their breeding according to the level of risk to their own survival and to survival probabilities and recruitment of young produced under different environmental conditions. Under risky or harsh conditions breeding can be postponed until later in the current breeding season or even to the next breeding season. We develop a population dynamics and life history model for polymorphism in reproduction (co-existence of breeding and non-breeding behaviours) of females in an age-structured population, with two temporally distinct mating events within the breeding season. We assume that, after overwintering, the females can breed in spring and again in summer or they can delay breeding in spring and breed in summer only. Young females born in spring can either mature and breed in summer or stay immature and postpone breeding over the winter to the next breeding season. We show that an evolutionarily stable breeding strategy is either an age-structured combination of pure breeding behaviours (old females breed and young delay maturity) or a mixed breeding behaviour within age-classes (a fraction of females breed and the rest of the age class postpones breeding). Co-occurrence of mixed reproductive behaviour in spring and summer within a single breeding season is observed in fluctuating populations only. The reproductive patterns depend on intraspecific, possibly interspecific, and ecological factors. The density dependence (e.g. social suppression) and predation risk are shown to be possible evolutionary mechanisms in adjusting the relative proportions of the different but co-existing reproductive behaviours.     

Pigment variability in larval Sebastes jordani off central California

Atypical pigment was observed in pre‐flexion, flexion and post‐flexion larval Sebastes collected off central California in 1991 and 1992 that otherwise resembled Sebastes jordani . Atypical pigment occurred on the axillary region at the base of the pectoral fin (most prominent on the inner edge of the pectoral fin base adjacent to the gut), on the median and distal regions of the pectoral fin and on the median and distal regions of the pelvic fin. In addition, lower and upper jaw pigment was observed at a much smaller size in these specimens than previously described in the literature. Identifications of these atypical specimens as S. jordani were confirmed using meristic and otolith characters as well as mitochondrial DNA sequence data. The ontogenetic variability of S. jordani is described. Specimens collected north of Point Conception were more pigmented than larval S. jordani described in the literature (collected predominately south of Point Conception), suggesting geographic variation in pigment development during the larval stage.     

The life cycles and growth of Carabus glabratus and C. violaceus in Budalen, central Norway

Abstract. 1. The seasonal activity patterns of adults and larvae, and the seasonal development of female gonads showed that Curubus glabrafus Payk. and Curubus violaceus L. living in a subalpine habitat are spring (summer) breeders with biennial life cycles.
2. As a rule, the adults and third stage larvae hibernate, but C.glubrutus at least can hibernate also at the second larval stage, particularly in years in which unfavourable climatic conditions in the spring delay breeding.
3. The adaptive strategies underlying these life history patterns and their possible significance for species diversification and coexistence are discussed.     

Trade-offs in larval performance on normal and novel hosts

The evolution of host specialization in phytophagous insects is generally thought to involve genetic trade-offs that prevent individuals from maximizing fitness simultaneously on two or more hosts. Several hypotheses, however, have suggested that trade-offs may not be evident in experiments comparing larval performance on normal and novel hosts. Tests on survivorship, growth rate, and pupal mass among families of the swallowtail butterfly Papilio oregonius on its normal host and on a novel host provide support for these hypotheses, although they do not discriminate among them. Families differed in their relative performance on the hosts, but there was no evidence of a negative genetic correlation between hosts for any of the measures of performance. In addition, there were no correlations among the different measures, corroborating an earlier result suggesting that these different components of performance in the P. machaon species group are under at least partially separate genetic control. These results and similar results published for other insects have now produced a body of studies indicating that genetic trade-offs in individual components of larval performance may not be a major factor preventing shifts onto novel host plants. Trade-offs leading to the evolution of host specialization are more likely to involve coordination among the various components of performance together with ecological factors that allow higher fitness on one host than on others.     

Evolutionarily singular strategies and the adaptive growth and branching of the evolutionary tree

We present a general framework for modelling adaptive trait dynamics in which we integrate various concepts and techniques from modern ESS-theory. The concept of evolutionarily singular strategies is introduced as a generalization of the ESS-concept. We give a full classification of the singular strategies in terms of ESS-stability, convergence stability, the ability of the singular strategy to invade other populations if initially rare itself, and the possibility of protected dimorphisms occurring within the singular strategy's neighbourhood. Of particular interest is a type of singular strategy that is an evolutionary attractor from a great distance, but once in its neighbourhood a population becomes dimorphic and undergoes disruptive selection leading to evolutionary branching. Modelling the adaptive growth and branching of the evolutionary tree can thus be considered as a major application of the framework. A haploid version of Levene's soft selection model is developed as a specific example to demonstrate evolutionary dynamics and branching in monomorphic and polymorphic populations.     

The evolution of parasites in response to tolerance in their hosts: the good, the bad, and apparent commensalism

Tolerance to parasites reduces the harm that infection causes the host (virulence). Here we investigate the evolution of parasites in response to host tolerance. We show that parasites may evolve either higher or lower within-host growth rates depending on the nature of the tolerance mechanism. If tolerance reduces virulence by a constant factor, the parasite is always selected to increase its growth rate. Alternatively, if tolerance reduces virulence in a nonlinear manner such that it is less effective at reducing the damage caused by higher growth rates, this may select for faster or slower replicating parasites. If the host is able to completely tolerate pathogen damage up to a certain replication rate, this may result in apparent commensalism, whereby infection causes no apparent virulence but the original evolution of tolerance has been costly. Tolerance tends to increase disease prevalence and may therefore lead to more, rather than less, disease-induced mortality. If the parasite is selected, even a highly efficient tolerance mechanism may result in more individuals in total dying from disease. However, the evolution of tolerance often, although not always, reduces the individual risk of dying from infection.     

Predicting trait values and measuring selection in complex life histories: reproductive allocation decisions in Soay sheep

Accurate prediction of life history phenomena and characterisation of selection in free-living animal populations are fundamental goals in evolutionary ecology. In density regulated, structured populations, where individual state influences fate, simple and widely used approaches based on individual lifetime measures of fitness are difficult to justify. We combine recently developed structured population modelling tools with ideas from modern evolutionary game theory (adaptive dynamics) to understand selection on allocation of female reproductive effort to singletons or twins in a size-structured population of feral sheep. In marked contrast to the classical selection analyses, our model-based approach predicts that the female allocation strategy is under negligible directional selection. These differences arise because classical selection analysis ignores components of offspring fitness and fails to consider selection over the complete life cycle.     

Evolution of airborne infectious diseases according to changes in characteristics of the host population

The authors predicted evolutionary changes in airborne infectious diseases according to changes in the characteristics of the host population. The predictions were based upon a mathematical model of infectious diseases and the validity of the predictions was verified against the history of man and pathogens. The feature of this model is that it involves a density of pathogens in the environment as an additional variable which can be regarded as more suitable to airborne infectious diseases. In spite of this modification, this study reached a similar conclusion to the threshold density theory: that is, susceptible host density in the absence of the pathogen must be larger than that in the presence of the pathogen, for the pathogen to be persistent. Moreover the authors concluded that one type of pathogen cannot be replaced by another type of pathogen as long as the susceptible host density of the former type is the mininum one. The predictions were considered to be valid for a wide range of infectous diseases. Making use of these principles, the authors predicted that the variety of infectious diseases should increase as host density increases and that pathogens should evolve to be less virulent as the host life-span increases. The finalidea discussed is whether or nor the history of man and pathogen can be verified by the predictions.     

Classic predictions about sex change do not hold under all types of size advantage

Theory predicts that the ‘size advantage’ (rate of increase in male and female fitness with age or size) determines the direction and the timing of sex change in sequential hermaphrodites. Whereas the size advantage is generated by the mating system and would be expected to vary within and between species, the shape or form of the size advantage has rarely been estimated directly. Here, we ask whether theoretical predictions about the timing of sex change hold under different types of size advantage. We model two biological scenarios representing different processes generating the size advantage and find that different types of size advantage can produce patterns that qualitatively differ from classic predictions. Our results demonstrate that a good understanding of sequentially hermaphroditic mating systems, and specifically, a direct assessment of the processes underlying the size advantage is crucial to reliably predict and explain within‐species patterns of the timing of sex change.     

Evolution of complex life cycles in trophically transmitted helminths. I. Host incorporation and trophic ascent

Links between parasites and food webs are evolutionarily ancient but dynamic: life history theory provides insights into helminth complex life cycle origins. Most adult helminths benefit by sexual reproduction in vertebrates, often high up food chains, but direct infection is commonly constrained by a trophic vacuum between free‐living propagules and definitive hosts. Intermediate hosts fill this vacuum, facilitating transmission to definitive hosts. The central question concerns why sexual reproduction, and sometimes even larval growth, is suppressed in intermediate hosts, favouring growth arrest at larval maturity in intermediate hosts and reproductive suppression until transmission to definitive hosts? Increased longevity and higher growth in definitive hosts can generate selection for larger parasite body size and higher fecundity at sexual maturity. Life cycle length is increased by two evolutionary mechanisms, upward and downward incorporation, allowing simple (one‐host) cycles to become complex (multihost). In downward incorporation, an intermediate host is added below the definitive host: models suggest that downward incorporation probably evolves only after ecological or evolutionary perturbations create a trophic vacuum. In upward incorporation, a new definitive host is added above the original definitive host, which subsequently becomes an intermediate host, again maintained by the trophic vacuum: theory suggests that this is plausible even under constant ecological/evolutionary conditions. The final cycle is similar irrespective of its origin (upward or downward). Insights about host incorporation are best gained by linking comparative phylogenetic analyses (describing evolutionary history) with evolutionary models (examining selective forces). Ascent of host trophic levels and evolution of optimal host taxa ranges are discussed.     

Life history strategies in two coexisting agamospecies of dandelion

Populations of Taraxacum , dandelion, on a Northumberland sand dune system contain 27 agamospecies. Demographic parameters for subpopulations containing a preponderance of two closely related agamospecies, Taraxacum lacistophyllum and T. brachyglossum are described. High and low density populations were stable over a 3 year period. The estimated turnover times differed between densities and species. The early growth characteristics of the two species are described. The plastochrons of the two species are distinct, as are the growth patterns of the leaves, the total leaf length per plant, the number of leaves and rate of leaf birth, and the patterns of allocation of dry weight to the root, shoot, and reproductive organs of the plant. Differences between the species were observed in fruit, scape height, the wind speed required to achieve fruit dispersal, and the pattern of fruit dispersal at different wind speeds. The origin and maintenance of agamospecies diversity is discussed in relation to the observed life history variation.     

Different life history strategies in lake-outlet blackflies (Diptera: Simuliidae)

The life histories of Simulium noelleri Fried. and S. truncatum Ldstr. were studied at a lake-outlet site in Finland. A detailed study was made of the second summer generation of both species and this consisted of a single, synchronized cohort in S. noelleri, and overlapped cohorts in S. truncatum. Larvae of the two species also show differences in their habitat preference: those of S. noelleri predominate on the sluice which marks the beginning of the river, and S. truncatum larvae just downstream from the sluice. In winter, the latter site was dominated by immature individuals of two other species, S. erythrocephalum de Geer and S. equinum L., and these were found downstream from the two ‘lake-outlet’ species during the summer. Adult females of S. noelleri are able to mature a first batch of eggs without taking a blood meal but those of S. truncatum need to feed on blood before eggs can be fully developed. The consequences of these different reproductive strategies on the ecology of individuals of the two species is discussed.     

THE JOINT EVOLUTION OF DISPERSAL AND DORMANCY IN A METAPOPULATION WITH LOCAL EXTINCTIONS AND KIN COMPETITION

Dispersal and dormancy are two strategies that allow recolonization of empty patches and escape from kin competition. Because they presumably respond to similar evolutionary forces, it is tempting to consider that these strategies may substitute for each other. Yet in order to predict the outcome of the evolution of dispersal and dormancy, and to characterize the emerging covariation between both traits, it is necessary to consider models where dispersal and dormancy evolve jointly. Here, we analyze the evolution of dispersal and dormancy as a function of direct fitness costs, environmental variation, and competition among relatives. We consider two scenarios depending on whether the rates of dormancy for philopatric and dispersed individuals are constrained to be the same (unconditional dormancy) or allowed to be different (conditional dormancy). We show that only philopatric individuals should enter dormancy, at a rate increasing with increasing rates of local extinction and decreasing population sizes. When dormancy and dispersal evolve jointly, we observe a wide range of evolutionary outcomes. In particular, we find that the pattern of covariation between the evolutionarily stable rates of dispersal and dormancy is molded by the rate of extinction and the local population size.     

Microgeographic variation in the effects of larval temperature environment on juvenile morphology and locomotion in the pool frog

In animals with complex life cycles, the environment experienced early during the development may have strong effects on later performance and fitness. We investigated the intraspecific variation in the effects of larval temperature environment on the morphology and locomotory performance of juvenile pool frogs Rana lessonae originating from three closely located populations of the northern fringe metapopulation in central Sweden. Tadpoles were raised individually at two temperatures (20 and 25 °C) until metamorphosis. We measured the morphology of the metamorphs and tested the jumping performance of the froglets after complete tail absorption. We found that early temperature environment affected juvenile morphology, metamorphs from high-temperature environments having relatively longer hindlimbs (tibiofibulas) and longer tails when weight at metamorphosis was accounted for. In absolute terms, froglets from low temperature jumped significantly longer; however, after correcting for size differences the relationship was reversed, individuals raised at high temperature performing better. In both temperatures, relative jumping performance was positively associated with tibiofibula and body length. Populations differed both in metamorphic traits and in jumping capacity, especially at low temperature, suggesting microgeographical variation in temperature sensitivity within the metapopulation. Our results indicate that the temperature environment experienced during the early aquatic stages can influence the morphology and performance of juvenile frogs, and that these effects can be population specific.     

洪水条件下湿地植物的生存策略

洪水是自然界存在的一种普遍现象。湿地植物由于所处生境的特殊性,会经常受到周期性或永久性的洪水胁迫。在长期的适应进化过程中,湿地植物形成了一些特殊的生存策略,以适应水文条件的大幅度变化。主要的生存策略如下:1)生活史方面,植物可通过改变生长时间、繁殖方式、种子特征等避免洪水的直接伤害或利用洪水的流动起到传播扩散的作用;2)形态学特征方面,植物可通过调整根系形态、分布等将根系生长到氧气相对充足的土壤表层或形成不定根增强根系通气功能;3)解剖学方面,植物可通过改善组织孔隙度形成通气组织等改善空气传导到根系的"气体通道";4)生理生化方面,植物可通过增加碳水化合物含量以延长生存时间,释放出一些生长激素(乙烯等)以调节植物缺氧条件下的生理活动或形态、解剖方面的变化。在今后的研究中,不定根的形成机理、乙烯在通气组织形成中的作用及其过程、放射氧损失(ROL)的形成机理及其释放速率的调控等一些机理性的工作还需进一步加强。     

Effects of embryo energy,egg size,and larval food supply on the development of asteroid echinoderms

Organisms have limited resources available to invest in reproduction, causing a trade‐off between the number and size of offspring. One consequence of this trade‐off is the evolution of disparate egg sizes and, by extension, developmental modes. In particular, echinoid echinoderms (sea urchins and sand dollars) have been widely used to experimentally manipulate how changes in egg size affect development. Here, we test the generality of the echinoid results by (a) using laser ablations of blastomeres to experimentally reduce embryo energy in the asteroid echinoderms (sea stars), Pisaster ochraceus and Asterias forbesi and (b) comparing naturally produced, variably sized eggs (1.7‐fold volume difference between large and small eggs) in A. forbesi. In P. ochraceus and A. forbesi, there were no significant differences between juveniles from both experimentally reduced embryos and naturally produced eggs of variable size. However, in both embryo reduction and egg size variation experiments, simultaneous reductions in larval food had a significant and large effect on larval and juvenile development. These results indicate that (a) food levels are more important than embryo energy or egg size in determining larval and juvenile quality in sea stars and (b) the relative importance of embryo energy or egg size to fundamental life history parameters (time to and size at metamorphosis) does not appear to be consistent within echinoderms.     

Fitness consequences of larval traits persist across the metamorphic boundary

Metamorphosis is thought to provide an adaptive decoupling between traits specialized for each life-history stage in species with complex life cycles. However, an increasing number of studies are finding that larval traits can carry-over to influence postmetamorphic performance, suggesting that these life-history stages may not be free to evolve independently of each other. We used a phenotypic selection framework to compare the relative and interactive effects of larval size, time to hatching, and time to settlement on postmetamorphic survival and growth in a marine invertebrate, Styela plicata. Time to hatching was the only larval trait found to be under directional selection, individuals that took more time to hatch into larvae survived better after metamorphosis but grew more slowly. Nonlinear selection was found to act on multivariate trait combinations, once again acting in opposite directions for selection acting via survival and growth. Individuals with above average values of larval traits were most likely to survive, but surviving individuals with intermediate larval traits grew to the largest size. These results demonstrate that larval traits can have multiple, complex fitness consequences that persist across the metamorphic boundary; and thus postmetamorphic selection pressures may constrain the evolution of larval traits.