Discrete Clutch Sizes, Local Mate Competition, and the Evolution of Precise Sex Allocation

Optimal sex allocation under a population structure with local mate competition has been studied mainly in deterministic models that are based on the assumption of continuous clutch sizes; Hamilton's (1967) model is the classic example. When clutch sizes are small, however, this assumption is not appropriate. When taking the discrete nature of eggs into account it becomes critically important whether females control only the mean sex ratio (“binomial” females) or the variance as well (“precise” females). As both types of sex ratio control have been found, it is of interest to investigate their evolutionary stability. In particular, it may be questioned whether perfect control of the sex ratio is always favoured by natural selection when mating groups are small. Models based on discrete clutch sizes are developed to determine evolutionarily stable (ES) sex ratios. It is predicted that when all females are of the binomial type they should produce a lower proportion of daughters than predicted by Hamilton's model, especially when clutch size and foundress number are small. When all females are of the precise type, the ES number of sons should generally be either a stable mixed strategy or a pure strategy, but there are special cases (for two foundresses and particular clutch sizes) where the ES number of sons lies in a trajectory of neutrally stable mixed strategies; the predicted mean sex ratios can be either higher or lower than predicted by Hamilton's model. The existence of ES mixed strategies implies that individual females do not necessarily have to produce sex ratios with perfect precision; some level of imperfection can be tolerated (i.e., will not be selected against). When the population consists of both binomial and precise females, the latter always have a selective advantage. This advantage of precision does not disappear when precision approaches fixation in the population. The latter result contradicts the conclusions of Taylor and Sauer (1980) which is due to their way of expressing selective advantage; they define selective advantage as the between-generation increase per allele, which will always become vanishingly small when an allele reaches fixation, irrespective of fitness differences.     

Female-biased Dispersal of the Emei Moustache Toad (Leptobrachium boringii) under Local Resource Competition

Dispersal is an important area of ecological and evolutionary research.Although many studies have been conducted in mammals and birds,dispersal pattern in amphibians is still unclear.To verify dispersal patterns of amphibians,an endangered species the Emei Moustache Toad(Leptobrachium boringii)was selected.In this study,we analyzed six genetic parameters:inbreeding coefficient(F_(IS)),gene diversity(H_S),the mean of corrected assignment index(mAI_C),the variance of corrected assignment index(vAI_C),relatedness(r)for all three years together and each year separately based on eight highly polymorphic microsatellite loci.Data in totaled across years and each year for 581 individuals captured during 2013–2015 revealed a significant female-biased dispersal pattern.Significantly higher F_(IS) and H_S in females,and lower mAI_C and r for each year separately in females support that L.boringii displays femalebiased dispersal,although r for the total dataset and vAI_C tests did not show significant differences between the sexes.Female-biased dispersal patterns may be explained by the local resource competition hypothesis.     

Social Mating System and Sex-Biased Dispersal in Mammals and Birds: A Phylogenetic Analysis

The hypothesis that patterns of sex-biased dispersal are related to social mating system in mammals and birds has gained widespread acceptance over the past 30 years. However, two major complications have obscured the relationship between these two behaviors: 1) dispersal frequency and dispersal distance, which measure different aspects of the dispersal process, have often been confounded, and 2) the relationship between mating system and sex-biased dispersal in these vertebrate groups has not been examined using modern phylogenetic comparative methods. Here, we present a phylogenetic analysis of the relationship between mating system and sex-biased dispersal in mammals and birds. Results indicate that the evolution of female-biased dispersal in mammals may be more likely on monogamous branches of the phylogeny, and that females may disperse farther than males in socially monogamous mammalian species. However, we found no support for a relationship between social mating system and sex-biased dispersal in birds when the effects of phylogeny are taken into consideration. We caution that although there are larger-scale behavioral differences in mating system and sex-biased dispersal between mammals and birds, mating system and sex-biased dispersal are far from perfectly associated within these taxa.     

Evolution of Dispersal Toward Fitness

It is widely believed that the slowest dispersal strategy is selected in the evolutional if the environment is temporally invariant but spatially heterogeneous. Authors claim in this paper that this belief is true only if random dispersals with constant motility are considered. However, if a dispersal strategy with fitness property is included, the size of the dispersal is not such a crucial factor anymore. Recently, a starvation driven diffusion has been introduced by Cho and Kim (Bull. Math. Biol., 2013), which is a random dispersal strategy with a motility increase on starvation. The authors show that such a dispersal strategy has fitness property and that the evolutional selection favors fitness but not simply slowness. Such a conclusion is obtained from a stability analysis of a competition system between two phenotypes with different dispersal strategies of linear and starvation driven diffusions.     

Evolution of Complex Density-Dependent Dispersal Strategies

The question of how dispersal behavior is adaptive and how it responds to changes in selection pressure is more relevant than ever, as anthropogenic habitat alteration and climate change accelerate around the world. In metapopulation models where local populations are large, and thus local population size is measured in densities, density-dependent dispersal is expected to evolve to a single-threshold strategy, in which individuals stay in patches with local population density smaller than a threshold value and move immediately away from patches with local population density larger than the threshold. Fragmentation tends to convert continuous populations into metapopulations and also to decrease local population sizes. Therefore we analyze a metapopulation model, where each patch can support only a relatively small local population and thus experience demographic stochasticity. We investigated the evolution of density-dependent dispersal, emigration and immigration, in two scenarios: adult and natal dispersal. We show that density-dependent emigration can also evolve to a nonmonotone, “triple-threshold” strategy. This interesting phenomenon results from an interplay between the direct and indirect benefits of dispersal and the costs of dispersal. We also found that, compared to juveniles, dispersing adults may benefit more from density-dependent vs. density-independent dispersal strategies.     

Evolution of Inbreeding and Outcrossing in Ferns and Fern-Allies

Abstract Mating systems of 18 species of homosporous ferns follow a bimodal distribution, similar to that observed for seed plants (Schemske and Lande, 1985). Most species are highly outcrossing, a few are inbreeding, and two species examined to date have mixed mating systems. Equisetum arvense and several species of lycopods are also highly outcrossing. Several mechanisms, including inbreeding depression, antheridiogen, and ontogenetic sequences that result in effectively unisexual gametophytes, promote outcrossing in homosporous ferns and perhaps other homosporous pteridophytes as well. In some species of homosporous ferns, selection has favored the evolution of inbreeding as an adaptation for colonization. High levels of intra- and interpopulational gene flow via spore dispersal, coupled with high levels of intergametophytic crossing, generally lead to genetically homogeneous populations and species of homosporous ferns. However, rock-dwelling ferns and ferns from xeric habitats may exhibit significant population genetic structure due to physically patchy habitats. Reticulate evolution in homosporous ferns may be enhanced by high levels of intergametophytic crossing.     

The Correlated Evolution of Dispersal and Mating-System Traits

The existence of an evolutionary syndrome linking dispersal and mating-system traits has been discussed in both plants and animals. In animals, dispersal as a means of inbreeding-avoidance has been cited as an ultimate cause of sex-biased dispersal. In plants, self-compatibility is widespread, which is often cited as a mechanism for reproductive assurance in organisms that have limited control of dispersal. Limited dispersal has also been hypothesized to minimize outbreeding depression and increase local adaptation. Here, we compare and contrast the various evolutionary hypotheses that link dispersal and the mating system in both plants and animals. We conclude that the majority of theoretical evidence supports the existence of two evolutionary syndromes: (1) outcrossing and dispersing; (2) inbreeding and not dispersing. In the light of the evidence compiled, we advocate for a redefinition of Baker’s law, which we consider to be an exception rather than the rule. As environmental heterogeneity is common in nature, the role of bet-hedging in the evolution of these strategies is likely to be very strong, albeit difficult to prove empirically. Lastly, we argue that exceptions to these two general syndromes (e.g., inbreeding and dispersing) should not be viewed as contradictory; instead they merit exceptional attention because they represent unusual biological adaptations. Throughout, we refer to specific empirical examples to illustrate these scenarios and conclude by suggesting that a meta-analysis of the available data would be a useful next step.     

Immigration,Local Dispersal Limitation,and the Repeatability of Community Composition under Neutral and Niche Dynamics

Repeatability of community composition has been a critical aspect for community structure, which is closely associated with community stability, predictability, conservation biology and ecological restoration. It has been shown that both immigration and local dispersal limitation can affect the community composition in both neutral and niche model. Hence, we use a spatially explicit individual-based model to investigate the potential influence of immigration rate and strength of local dispersal limitation on repeatability in both neutral and niche models. Similarity measures are used to quantify repeatability. We examine the repeatability of community composition among replicate communities (which means the same community repeats many times), and between niche and neutral replicate communities. We find the correlation between repeatability and immigration rate is positive in the neutral model and an inverted unimodal in the niche model. The correlation between repeatability and local dispersal distance is positive in the niche model and negative in the neutral model. High repeatability between niche communities and neutral communities is observed with high immigration rates or when high local dispersal distance appears in the niche model or low local dispersal distance in the neutral model. Our results show that repeatability of community composition is not only dependent on the types of community models (niche vs. neutrality) but also strongly determined by immigration rates and local dispersal limitation.     

Biological Invasions Across Spatial Scales: Intercontinental, Regional, and Local Dispersal of Cladoceran Zooplankton

The frequency of dispersal of invertebrates among lakes depends upon perspective and spatial scale. Effective passive dispersal requires both the transport of propagules and the establishment of populations large enough to be detected. At a global scale, biogeographic patterns of cladoceran zooplankton species suggest that effective dispersal among continents was originally rare, but greatly increased in the past century with expanded commerce. Genetic analysis allows some reconstruction of past dispersal events. Allozyme and mitochondrial DNA comparisons among New World and Old-World populations of several exotic cladocerans have provided estimates for likely source populations of colonists, their dispersal corridors, and timing of earlier dispersal events. Detecting the Old-World tropical exotic Daphnia lumholtzi early in its invasion of North America has allowed detailed analysis of its spatial spread. Twelve years of collection records indicate a rapid invasion of reservoirs in the United States, by both regional spread and long-distance jumps to new regions. Combining landscape features with zooplankton surveys from south-central US reservoirs revealed higher colonization rates of D. lumholtzi at lower landscape positions, a result which can be explained by either greater propagule load or by higher susceptibility of these downstream reservoirs. Because invaded reservoirs provide a source of propagules for nearby floodplain ponds, the rarity of this species in ponds suggests limitation by local environments. Such analyses of invading species over multiple spatial scales allow a better understanding of ecological processes governing invasion dynamics.     

Sex-Biased Dispersal of a Frog (Odorrana schmackeri) Is Affected by Patch Isolation and Resource Limitation in a Fragmented Landscape

Sex-biased dispersal is widespread in the animal kingdom and is affected by numerous factors including mating system, social factors and environmental conditions. Unlike birds and mammals, there is no common trend in amphibians and explaining the direction and degree of sex-biased dispersal in species-specific cases is difficult. We conducted a study on dispersal of the Chinese piebald odorous frog (Odorrana schmackeri) in a fragmented landscape associated with dam construction. Ten microsatellite loci were used to analyze 382 samples sourced from 14 fragmented ‘islands’. Assignment tests indicated a significant pattern of female-biased dispersal on one island with inconsistencies in the strength and direction of this pattern between nearby islands. The effects of four island attributes and two potential impact factors on the pattern of sex-biased dispersal were examined. We found that the extent of isolation from the mainland and the number of breeding sites both showed a negative correlation with female biased dispersal, such that the closer an island is to the mainland the more likely it is to display female biased dispersal, and the more breeding sites on an island the more male immigrants. Based on these results, we conclude that geographic isolation and limited breeding resources are the most likely explanation for the patterns of dispersal observed in this fragmented population of amphibians.     

Gene-Culture Coevolution and Sex-Ratios: The Effects of Infanticide, Sex-Selective Abortion, Sex Selection, and Sex-Biased Parental Investment on the Evolution of Sex Ratios

The evolutionary consequences of culturally transmitted practices that cause differential mortality between the sexes, thereby distorting the sex ratio (e.g., female infanticide and sex-selective abortion), are explored using dynamic models of gene-culture coevolution. We investigate how a preference for the sex of offspring may affect the selection of genes distorting the primary sex ratio. Sex-dependent differences in mortality have been predicted to select for a male- or female-biased primary sex ratio, to have no effect, or to favor either under different circumstances. We find that when a mating pair′s behavior modifies mortality rates in favor of one sex, but does not change the number of offspring produced in the mating, the primary sex ratio will evolve a bias against the favored sex However, when the total number of offspring of a mating pair is significantly seduced as a consequence of their prejudice, the primary sex ratio will evolve to favor the preferred sex. These results hold irrespective of whether the sex ratio is distorted by the mother′s, the father′s or the individual′s own autosomal genes. The use of dynamic models of gene-culture coevolution allows us to explore the evolution of alleles which distort the sex ratio, as well as the final equilibrium states of the system. Gene-culture interactions can provide equilibria different from those in purely genetic systems, slow the approach to these equilibria by orders of magnitude, and move the primary (PSR) and the adult sex ratio (ASR) away from any stable equilibrium for hundreds of generations.     

Inbreeding,inbreeding depression and extinction

Inbreeding is unavoidable in small, isolated populations and can cause substantial fitness reductions compared to outbred populations. This loss of fitness has been predicted to elevate extinction risk giving it substantial conservation significance. Inbreeding may result in reduced fitness for two reasons: an increased expression of deleterious recessive alleles (partial dominance hypothesis) or the loss of favourable heterozygote combinations (overdominance hypothesis). Because both these sources of inbreeding depression are dependent upon dominance variance, inbreeding depression is predicted to be greater in life history traits than in morphological traits. In this study we used replicate inbred and control lines of Drosophila simulans to address three questions:1) is inbreeding depression greater in life history than morphological traits? 2) which of the two hypotheses is the major underlying cause of inbreeding depression? 3) does inbreeding elevate population extinction risk? We found that inbreeding depression was significantly greater in life history traits compared to morphological traits, but were unable to find unequivocal support for either the overdominance or partial dominance hypotheses as the genetic basis of inbreeding depression. As predicted, inbred lines had a significantly greater extinction risk.     

Fitness Traits and Dispersal Ability in the Herb Tragopogon pratensis (Asteraceae): Decoupling the Role of Inbreeding Depression and Maternal Effects

Abstract: Inbreeding depression can decrease several fitness traits and maternal effects can strongly influence the amount of inbreeding depression. Understanding the effects of inbreeding depression on plant fitness is especially important in the context of habitat fragmentation, where plant populations become smaller and more isolated, exhibiting increasing levels of inbreeding depression. We examined the joint influence of inbreeding depression and maternal effects on life cycle traits and dispersal ability in the herb Tragopogon pratensis that grows in fragmented populations in Europe. We conducted experimental crosses to obtain selfed and outcrossed progeny in two contrasted environments. In particular, we produced a first generation of seeds and plants that were self-pollinated again to produce a second generation of seeds. Individual seeds were weighed and their pappuses measured to estimate the dispersal potential. Pollination treatment only had a significant effect on seed mass and dispersal ability. Coefficients of inbreeding depression did not differ between selfed and outcrossed plants. Seed mass had a significant effect on germination date. Environment had a significant effect on mass of the second generation of seeds and the interaction between pollination treatment and family was significant for six traits, indicating the existence of strong maternal effects in T. pratensis. Results suggest population differentiation. Overall, T. pratensis populations exhibited a good performance under selfing, in terms of life cycle traits and dispersal ability, which would allow the species to cope with problems associated with fragmentation.     

Synchrony of Seed Dispersal, Hydrology and Local Climate in a Semi-arid River Reach in California

The temporal availability of propagules is a critical factor in sustaining pioneer riparian tree populations along snowmelt-driven rivers because seedling establishment is strongly linked to seasonal hydrology. River regulation in semi-arid regions threatens to decouple seed development and dispersal from the discharge regime to which they evolved. Using the lower Tuolumne River as a model system, we quantified and modeled propagule availability for Populus fremontii (POFR), Salix gooddingii (SAGO), and Salix exigua (SAEX), the tree and shrub species that dominate near-channel riparian stands in the San Joaquin Basin, CA. A degree-day model was fit to field data of seasonal seed density and local temperature from three sites in 2002–2004 to predict the onset of the peak dispersal period. To evaluate historical synchrony of seed dispersal and seasonal river hydrology, we compared peak spring runoff timing to modeled peak seed release periods for the last 75 years. The peak seed release period began on May 15 for POFR (range April 23–June 10), May 30 for SAGO (range May 19–June 11) and May 31 for SAEX (range May 8–June 30). Degree-day models for the onset of seed release reduced prediction error by 40–67% over day-of-year means; the models predicted best the interannual, versus site-to-site, variation in timing. The historical analysis suggests that POFR seed release coincided with peak runoff in almost all years, whereas SAGO and SAEX dispersal occurred during the spring flood recession. The degree-day modeling approach reduce uncertainty in dispersal timing and shows potential for guiding flow releases on regulated rivers to increase riparian tree recruitment at the lowest water cost.     

Modeling Within-Host Evolution of HIV: Mutation,Competition and Strain Replacement

Virus evolution during infection of a single individual is a well-known feature of disease progression in chronic viral diseases. However, the simplest models of virus competition for host resources show the existence of a single dominant strain that grows most rapidly during the initial period of infection and competitively excludes all other virus strains. Here, we examine the dynamics of strain replacement in a simple model that includes a convex trade-off between rapid virus reproduction and long-term host cell survival. Strains are structured according to their within-cell replication rate. Over the course of infection, we find a progression in the dominant strain from fast- to moderately-replicating virus strains featuring distinct jumps in the replication rate of the dominant strain over time. We completely analyze the model and provide estimates for the replication rate of the initial dominant strain and its successors. Our model lays the groundwork for more detailed models of HIV selection and mutation. We outline future directions and application of related models to other biological situations.     

Inbreeding,energy use and condition

In energetic terms, fitness may be seen to be dependent on successful allocation of energy between life‐history traits. In addition, fitness will be constrained by the energy allocation ability, which has also been defined as condition. We suggest here that the allocation ability, estimated as the difference between total energy budget and maintenance metabolism, may be used as a measure of condition. We studied this possibility by measuring the resting metabolic rate and metabolism during forced exercise in Gryllodes sigillatus crickets. To verify that these metabolic traits are closely related to fitness, we experimentally manipulated the degree of inbreeding of individuals belonging to the same pedigree, hence enabling analysis of both inbreeding depression and heritability of traits. We found that inbreeding increased maintenance metabolism, whereas total energy budget was rather insensitive to inbreeding. Despite this, inbreeding led to decreased allocation ability. Overall, metabolic traits exhibited strong inbreeding depression and rather low heritabilities, a pattern that is typical of traits under strong selection. However, traditionally used condition indices were not affected by inbreeding and did not covary with metabolic traits. Moreover, in contrast to the common, but largely untested, tenet, it seems that high resting metabolic rate is indicative of low rather than high quality.     

藓类植物传孢类型及其系统演化关系

藓纲包括３个亚纲（Ｂｒｏｔｈｅｒｕｓ,１９２４－１９２５）。泥炭藓亚纲、黑藓亚纲,真藓亚纲,全世界约１００个科,近８００属,约１５０００种。藓类的传孢类型和藓类植物系统演化有密切关系。我们认为传孢不同类型,直接反映了他们的演化程度。我们根据藓类孢子体的分化、蒴齿分化与其机能,分为５种传孢类型。即腐媒传孢型,风媒孢型,汽－风媒传孢型,水媒传孢型,和虫媒传孢型等５种。根据它们的演化程度比较,我们提出了     

Evolution of Dispersal in a Structured Metapopulation Model in Discrete Time

In this article, a structured metapopulation model in discrete time with catastrophes and density-dependent local growth is introduced. The fitness of a rare mutant in an environment set by the resident is defined, and an efficient method to calculate fitness is presented. With this fitness measure evolutionary analysis of this model becomes feasible. This article concentrates on the evolution of dispersal. The effect of catastrophes, dispersal cost, and local dynamics on the evolution of dispersal is investigated. It is proved that without catastrophes, if all population–dynamical attractors are fixed points, there will be selection for no dispersal. A new mechanism for evolutionary branching is also found: Even though local population sizes approach fixed points, catastrophes can cause enough temporal variability, so that evolutionary branching becomes possible.