Effects of Overlapping Generations on Linkage Disequilibrium Estimates of Effective Population Size

Use of single-sample genetic methods to estimate effective population size has skyrocketed in recent years. Although the underlying models assume discrete generations, they are widely applied to age-structured species. We simulated genetic data for 21 iteroparous animal and plant species to evaluate two untested hypotheses regarding performance of the single-sample method based on linkage disequilibrium (LD): (1) estimates based on single-cohort samples reflect the effective number of breeders in one reproductive cycle (N_b), and (2) mixed-age samples reflect the effective size per generation (N_e). We calculated true N_e and N_b, using the model species’ vital rates, and verified these with individual-based simulations. We show that single-cohort samples should be equally influenced by N_b and N_e and confirm this with simulated results: N^b was a linear (r² = 0.98) function of the harmonic mean of N_e and N_b. We provide a quantitative bias correction for raw N^b based on the ratio N_b/N_e, which can be estimated from two or three simple life history traits. Bias-adjusted estimates were within 5% of true N_b for all 21 study species and proved robust when challenged with new data. Mixed-age adult samples produced downwardly biased estimates in all species, which we attribute to a two-locus Wahlund effect (mixture LD) caused by combining parents from different cohorts in a single sample. Results from this study will facilitate interpretation of rapidly accumulating genetic estimates in terms of both N_e (which influences long-term evolutionary processes) and N_b (which is more important for understanding eco-evolutionary dynamics and mating systems).     

OPTIMAL REPRODUCTIVE EFFORT IN STOCHASTIC,DENSITY-DEPENDENT ENVIRONMENTS

The amount of effort organisms should put into reproducing at any given time has been a matter of debate for many years. Early models suggested a simple rule of thumb: iteroparity should be favored when juvenile survival is relatively variable and semelparity when adult survival is relatively variable. When more mathematically complex models were developed, these simple conclusions were found to be special cases. Variability can select toward iteroparity or semelparity depending on a number of factors irrespective of relative adult/juvenile survival (e.g, the density-independent models of Orzack and Tuljapurkar). Using new techniques, we estimate the ESS reproductive effort for stage-structured models in density-dependent and stochastic conditions. We find that variability causes significant changes in reproductive effort, these changes are often small (± 10% of determinstic ESS effort, but up to 50% change in some instances), and the amount that effort increases or decreases depends on many factors (e.g., the deterministic population dynamics, the vital rates affected by density, the amount of variation, the correlations between the vital rates, the distribution from which the variation is drawn, and the deterministic ESS effort). In a variable environment, semelparity is the ESS in only 3.5% of cases; iteroparity is the rule.     

Pattern of gonad maturation and the question of semelparity in the paedomorphic goby Aphia minuta

At variance with previous reports, the paedomorphic goby Aphia minuta does not show semelparity, but abbreviate iteroparity like other small Mediterranean gobiid species. In spring, the mature ovary contains several batches of eggs at different stages of vitellogenesis. This indicates that the breeding season of A. minuta is quite long and that spawning takes place at least twice during its short lifespan, involving in each ovulatory wave different batches of oocytes which undergo vitellogenesis at different times. These observations agree with the testicular cycle data. In males, which in spring exhibit active spermiation coinciding with deposition of the first batch of oocytes, spermatogonial mitosis also resumes, eventually resulting in the production of new sperm in summer.     

Number and size of clutch in Ptomascopus morio (Coleoptera: Silphidae: Nicrophorinae), with comparisons to Nicrophorus

Ptomascopus morio displays simpler parental care than Nicrophorus species. The effects of carcass size and clutch number on clutch size in P. morio were examined. Clutch size was related to carcass size. There was a negative correlation between number of clutch and clutch size for most sizes of carcass. Longevity of females was shorter when the carcass size was larger, such that total lifetime fecundity did not differ among carcasses of different sizes. The clutch size of P. morio was larger than that of Nicrophorus quadripunctatus. The clutch size of P. morio declined rapidly with repeated clutch production, but that of N. quadripunctatus was rather constant. This indicates that N. quadripunctatus maintains a more constant clutch size than P. morio over several reproductive attempts, although the former displays more complex parental care.     

TO AGE, TO DIE: PARITY, EVOLUTIONARY TRACKING AND COLE'S PARADOX

The existence of semelparity or "big bang" reproduction (reproducing only once in a lifetime) and iteroparity (reproducing more than once in a lifetime) has led to many questions investigating the evolution or persistence of these strategies. Here we investigate semelparity and iteroparity for their evolutionary importance. A mathematical model is used to illustrate how a population's ability to evolve depends on this life-history trait, and how this rate of evolution impacts the individual. We find that the ability of a trait to evolve is greater toward a semelparous strategy and this expresses a fitness advantage. This leads to an optimality between survival, population tracking ability, and lifetime fecundity.     

Setting priorities for possum control at a regional level

Abstract

Because funds for possum control are always insufficient to carry out control in all the areas that may require it, it is important that the funds available are allocated to those areas that most deserve it. This can be achieved only by implementing a process that first identifies the resource types likely to be degraded by possums, and secondly ranks areas for control within each resource type. If necessary, high priority areas between resource types can also be ranked. A method for undertaking such a process is described.     

DOES EVOLUTION OF ITEROPAROUS AND SEMELPAROUS REPRODUCTION CALL FOR SPATIALLY STRUCTURED SYSTEMS?

Abstract.— A persistent question in the evolution of life histories is the fitness trade-off between reproducing only once (semelparity) in a lifetime or reproducing repeated times in different seasons (iteroparity). The problem can be formulated into a research agenda by assuming that one reproductive strategy is resident (has already evolved) and by asking whether invasion (evolution) of an alternative reproductive strategy is possible. For a spatially nonstructured system, Bulmer (1994) derived the relationship v + PA < 1 (PA is adult survival; vbs and bs are offspring numbers for iteroparous and semelparous breeding strategies, respectively) at which semelparous population cannot be invaded by an iteroparous mutant. When the inequality is changed to v + PA > 1, invasion of a semelparous mutant is not possible. From the inequalities, it is easy to see that possibilities for evolutionary establishment of a novel reproductive strategy are rather narrow. We extended the evolutionary scenario into a spatially structured system with dispersal linkage among the subunits. In this domain, a rare reproductive strategy can easily invade a population dominated by a resident reproductive strategy. The parameter space enabling invasion is far more generous with spatially structured evolutionary scenarios than in a spatially nonstructured system.     

Climate and life-history evolution in evening primroses (Oenothera, Onagraceae): a phylogenetic comparative analysis

Evolutionary ecologists have long sought to understand the conditions under which perennial (iteroparous) versus annual (semelparous) plant life histories are favored. We evaluated the idea that aridity and variation in the length of droughts should favor the evolution of an annual life history, both by decreasing adult survival and by increasing the potential for high seedling survival via reduced plant cover. We calculated phylogenetically independent contrasts of climate with respect to life history in a clade of winter-establishing evening primroses (sections Anogra and Kleinia; Oenothera; Onagraceae), which includes seven annuals, 12 perennials, and two variable taxa. Climate variables were quantified from long-term records at weather stations near collection localities. To explicitly account for phylogenetic uncertainty, contrasts were calculated on a random sample of phylogenetic trees from the posterior distribution of a Bayesian analysis of DNA sequence data. Statements of association are based on comparing the per-tree mean contrast, which has a null expectation of zero, to a set of per-tree mean contrasts calculated on the same trees, after randomizing the climate data. As predicted, increased annual aridity, increased annual potential evapotranspiration, and decreased annual precipitation were associated with transitions to the annual habit, but these trends were not significantly different from the null pattern. Transitions to the annual habit were not significantly associated with increases in one measure of aridity in summer nor with increased summer drought, but they were associated with significantly increased maximum summer temperatures. In winter, increased aridity and decreased precipitation were significantly associated with transitions to the annual habit. Changes in life history were not significantly associated with changes in the coefficient of variation of precipitation, either on an annual or seasonal (summer vs. winter) basis. Though we cannot attribute causality on the basis of a correlational, historical study, our results are consistent with the idea that increased heat and drought at certain times of the year favor the evolution of the annual habit. Increased heat in summer may cause adult survival to decline, while increased aridity and decreased precipitation in the season of seedling recruitment (winter) may favor a drought-avoiding, short-lived annual strategy. Not all of the predicted patterns were observed: the capability for drought-induced dormancy may preclude change in habit in response to summer drought in our study group.     

Fire, rain and the selection of seeder and resprouter life-histories in fire-recruiting, woody plants

Several Cape species of the genus Erica are known to present seeder and resprouter phenotypes, and this variation seems to have a genetic basis. Therefore, this genus provides ideal model systems for using to elucidate the evolution of nonsprouting or seeder and resprouter life-histories in woody, fire-recruiting plants. A simple simulation model was developed to identify, under life-history optimality, the ecological conditions (viz. rainfall conditions and fire frequency) that conferred a selective advantage to the seeder phenotype over the resprouter in a given Cape Erica species. The model illustrated that the seeder life-history was able to invade and replace a resprouter population only under a mild mediterranean climate, with short, moderate summer droughts. This simulation approach will contribute to a better understanding of the biogeographical pattern of seeder and resprouter lineages of one of the paradigmatic fynbos woody taxa throughout the Cape floristic region.     

Simple life-history traits explain key effective population size ratios across diverse taxa

Effective population size (N_e) controls both the rate of random genetic drift and the effectiveness of selection and migration, but it is difficult to estimate in nature. In particular, for species with overlapping generations, it is easier to estimate the effective number of breeders in one reproductive cycle (N_b) than N_e per generation. We empirically evaluated the relationship between life history and ratios of N_e, N_b and adult census size (N) using a recently developed model (agene) and published vital rates for 63 iteroparous animals and plants. N_b/N_e varied a surprising sixfold across species and, contrary to expectations, N_b was larger than N_e in over half the species. Up to two-thirds of the variance in N_b/N_e and up to half the variance in N_e/N was explained by just two life-history traits (age at maturity and adult lifespan) that have long interested both ecologists and evolutionary biologists. These results provide novel insights into, and demonstrate a close general linkage between, demographic and evolutionary processes across diverse taxa. For the first time, our results also make it possible to interpret rapidly accumulating estimates of N_b in the context of the rich body of evolutionary theory based on N_e per generation.