LABORATORY EVOLUTION OF LIFE-HISTORY TRAITS IN THE BEAN WEEVIL (ACANTHOSCELIDES OBTECTUS): THE EFFECTS OF DENSITY-DEPENDENT AND AGE-SPECIFIC SELECTION

Four types of laboratory populations of the bean weevil (Acanthoscelides obtectus) have been developed to study the effects of density-dependent and age-specific selection. These populations have been selected at high (K) and low larval densities (r) as well as for reproduction early (Y) and late (O) in life. The results presented here suggest that the r- and K-populations (density-dependent selection regimes) have differentiated from each other with respect to the following life-history traits: egg-to-adult viability at high larval density (K > r), preadult developmental time (r > K), body weight (r > K), late fecundity (K > r), total realized fecundity (r > K), and longevity of males (r > K). It was also found that the following traits responded in statistically significant manner in populations subjected to different age-specific selection regimes: egg-to-adult viability (O > Y), body weight (O > Y), early fecundity (Y > O), late fecundity (O > Y), and longevity of females and males (O > Y). Although several life-history traits (viability, body weight, late fecundity) responded in similar manner to both density-dependent and age-specific selection regimes, it appears that underlying genetic and physiological mechanisms responsible for differentiation of the r/K and Y/O populations are different. We have also tested quantitative genetic basis of the bean weevil life-history traits in the populations experiencing density-dependent and age-specific selection. Among the traits traded-off within age-specific selection regimes, only early fecundity showed directional dominance, whereas late fecundity and longevity data indicated additive inheritance. In contrast to age-specific selecton regimes, three life-history traits (developmental time, body size, total fecundity) in the density-sependent regimes exhibited significant dominance effects. Lastly, we have tested the congruence between short-term and long-term effects of larval densities. The comparisons of the outcomes of the r/K selection regimes and those obtained from the low- and high-larval densities revealed that there is no congruence between the selection results and phenotypic plasticity for the analyzed life-history traits in the bean weevil.     

The Strengths of r- and K-Selection Shape Diversity-Disturbance Relationships

Disturbance is a key factor shaping species abundance and diversity in plant communities. Here, we use a mechanistic model of vegetation diversity to show that different strengths of r- and K-selection result in different disturbance-diversity relationships. R- and K-selection constrain the range of viable species through the colonization-competition tradeoff, with strong r-selection favoring colonizers and strong K-selection favoring competitors, but the level of disturbance also affects the success of species. This interplay among r- and K-selection and disturbance results in different shapes of disturbance-diversity relationships, with little variation of diversity with no r- and no K-selection, a decrease in diversity with r-selection with disturbance rate, an increase in diversity with K-selection, and a peak at intermediate values with strong r- and K-selection. We conclude that different disturbance-diversity relationships found in observations may reflect different intensities of r- and K-selection within communities, which should be inferable from broader observations of community composition and their ecophysiological trait ranges.     

DENSITY-DEPENDENT EVOLUTION OF LIFE-HISTORY TRAITS IN DROSOPHILA MELANOGASTER

Populations of Drosophila melanogaster were maintained for 36 generations in r- and K-selected environments in order to test the life-history predictions of theories on density-dependent selection. In the r-selection environment, populations were reduced to low densities by density-independent adult mortality, whereas populations in the K-selection environment were maintained at their carrying capacity. Some of the experimental results support the predictions or r- and K-selection theory; relative to the r-selected populations, the K-selected populations evolved an increased larval-to-adult viability, larger body size, and longer development time at high larval densities. Mueller and Ayala (1981) found that K-selected populations also have a higher rate of population growth at high densities. Other predictions of the thoery are contradicted by the lack of differences between the r and K populations in adult longevity and fecundity and a slower rate of development for r-selected individuals at low densities. The differences between selected populations in larval survivorship, larval-to-adult development time, and adult body size are strongly dependent on larval density, and there is a significant interaction between populations and larval density for each trait. This manifests an inadequacy of the theory on r- and K-selection, which does not take into account such interactions between genotypes and environments. We describe mechanisms that may explain the evolution of preadult life-history traits in our experiment and discuss the need for changes in theories of density-dependent selection.     

NATURAL GENETIC VARIATION OF LIFE SPAN,REPRODUCTION, AND JUVENILE GROWTH IN DAPHNIA

The evolutionary theory of senescence predicts that high extrinsic mortality in natural populations should select for accelerated reproductive investment and shortened life span. Here, we test the theory with natural populations of the Daphnia pulex-pulicaria species complex, a group of freshwater zooplankton that spans an environmental gradient of habitat permanence. We document substantial genetic variation in demographic life-history traits among parent and hybrid populations of this complex. Populations from temporary ponds have shorter life spans, earlier and faster increases of intrinsic mortality risk, and earlier and steeper declines in fecundity than populations from permanent lakes. We also examine the age-specific contribution to fitness, measured by reproductive value, and to expected lifetime reproduction; these traits decline faster in populations from temporary ponds. Despite having more rapid senescence, pond Daphnia exhibit faster juvenile growth and higher early fitness, measured as population growth rate (r). Among populations within this species complex we observed negative genetic correlations between r and indices of life-history timing, suggesting trade-offs between early- and late-life performance. Our results cannot be explained by a trade-off between survival and fecundity or by nonevolutionary theories of senescence. Instead, our data are consistent with the evolutionary theory of senescence because the genetic variation in life histories we observed is roughly congruent with the temporal scale of environmental change in the field.     

Density-independent mortality and increasing plant diversity are associated with differentiation of Taraxacum officinale into r- and K-strategists

Background

Differential selection between clones of apomictic species may result in ecological differentiation without mutation and recombination, thus offering a simple system to study adaptation and life-history evolution in plants.

Methodology/Principal Findings

We caused density-independent mortality by weeding to colonizer populations of the largely apomictic Taraxacum officinale (Asteraceae) over a 5-year period in a grassland biodiversity experiment (Jena Experiment). We compared the offspring of colonizer populations with resident populations deliberately sown into similar communities. Plants raised from cuttings and seeds of colonizer and resident populations were grown under uniform conditions. Offspring from colonizer populations had higher reproductive output, which was in general agreement with predictions of r-selection theory. Offspring from resident populations had higher root and leaf biomass, fewer flower heads and higher individual seed mass as predicted under K-selection. Plants grown from cuttings and seeds differed to some degree in the strength, but not in the direction, of their response to the r- vs. K-selection regime. More diverse communities appeared to exert stronger K-selection on resident populations in plants grown from cuttings, while we did not find significant effects of increasing species richness on plants grown from seeds.

Conclusions/Significance

Differentiation into r- and K-strategists suggests that clones with characteristics of r-strategists were selected in regularly weeded plots through rapid colonization, while increasing plant diversity favoured the selection of clones with characteristics of K-strategists in resident populations. Our results show that different selection pressures may result in a rapid genetic differentiation within a largely apomictic species. Even under the assumption that colonizer and resident populations, respectively, happened to be r- vs. K-selected already at the start of the experiment, our results still indicate that the association of these strategies with the corresponding selection regimes was maintained during the 5-year experimental period.     

Effects of local density on fecundity and mating speed for periodical cicadas

Summary This paper catalogues several different dichotomies that have all been termed r- and K-selection. The status of the concept of r- and K-selection is discussed and a more restricted usage of the terms is recommended.     

SEX RATIO VARIATION IN A PARASITIC WASP II. DIALLEL CROSS

Sex ratio has been studied from many theoretical and empirical perspectives, but a general assumption in sex ratio research is that changes in sex ratio occur because of selection on sex ratio itself. I carried out a quantitative genetic experiment—a diallel cross among three strains—on a parasitic wasp, Muscidifurax raptor (Hymenoptera: Pteromalidae), to measure genetic variation for sex ratio. I also tested whether sex ratio may change as a consequence of selection on other life-history traits by estimating genetic covariances between sex ratio, fecundity, longevity, and development time. Most of the variation among strains could be accounted for by a maternal effect, likely caused by a microsporidian parasite that was transmitted through the West Germany (WG) strain. Genetic variation was small by comparison, but almost all traits were affected by dominance. The only significant additive genetic effect was for fecundity early in life. Upon crossing, all traits displayed heterosis: more female-biased sex ratio, greater fecundity, longer life, and faster development time. All life-history traits were correlated phenotypically, but the correlations were mainly the result of decreased performance in crosses with the WG strain that carried the microsporidian parasite. Dominance genetic correlations were also found between sex ratio, fecundity, and longevity. How the correlation between sex ratio and other life-history traits would affect sex ratio evolution depends upon the frequencies of sex-ratio genotypes within a population as well as the signs of the correlations, because sex ratio is under frequency-dependent selection whereas other traits are generally under directional selection. Although the results from crosses among laboratory populations should be approached with caution, the inbreeding depression (the difference between inbred and outcrossed progeny) found in M. raptor implies that the evolution of a female-biased sex ratio could be affected by selection for inbreeding avoidance.     

The life-history traits of widespread freshwater fish species: the case of Roach Rutilus rutilus (Linnaeus, 1758) (Pisces: Teleostei) in water bodies at the southern limits of distribution

The Roach Rutilus rutilus (Linnaeus, 1758) (Pisces: Teleostei) is a widespread Eurasian cyprinid freshwater fish. Although numerous studies have investigated the species’ life history traits across its main native range of distribution, little is known from water bodies located beyond the southern limits of the latter, especially where the species has been introduced. Between July 2010 and April 2011, a total of 949 R. rutilus specimens was collected from Porsuk Reservoir (NW Turkey), and life-history traits of age, growth, fecundity and egg size were studied and compared to six other R. rutilus populations (including Caspian Roach R. r. caspicus) from the southern limits of distribution. The Porsuk population was characterised by relatively shorter lifespan, slower growth and lower fecundity. These findings support previous studies emphasising the importance of local environmental factors in shaping the life-history traits of widespread fish species. These must be taken into consideration especially in view of the potential value of R. rutilus as an economic resource for in-land fisheries.     

Anhydrobiosis of <Emphasis Type="Italic">Adineta ricciae:</Emphasis> Costs and Benefits

To study the effect of anhydrobiosis on the rotifer life cycle, we dried a bdelloid species, Adineta ricciae, and determined the life-history traits of 1) the rehydrated animals and 2) the offspring produced after a period of dormancy. In the first experiment, a cohort was dried when 8-days-old, kept dry for 7 days and then rehydrated. Recovery was about 75%. The recovered rotifers had similar longevity and significantly higher fecundity than the hydrated controls. The time spent dry was completely disregarded by the anhydrobiotic rotifers, as predicted by the ‘Sleeping Beauty’ model. In the second experiment, we recorded the life-history traits of the orthoclones produced by recovered mothers aged 11 days, and 18 days. These orthoclones were coupled to controls that had been established from hydrated mothers aged 11 days and 18 days. The orthoclones produced after dormancy had significantly higher fecundity and longevity than the controls of same maternal age. Maternal age had a marked effect on the life-history traits of the orthoclones of both lines, causing the same loss of fitness in both.     

Eco-physiological Characterization of Soil Bacterial Populations in Different States of Growth

The method based on characterization of microbial populations in terms of their growth rate in agar plates has been used for testing the prediction of the theory of r- and K-selection in a microbial community from a tropical soil. Conditions which could lead bacterial populations to grow exponentially or to enter into a stationary phase were obtained by growing soil microbial populations in a chemostat and in a chemostat with recycle, respectively. Significant differences in population distribution patterns were observed by comparing results from the two growth systems. When soil community was grown in a chemostat and subjected specifically to well-defined r- and K-conditions, stable associations of organisms with r- and K-type characteristics developed as a consequence of environmental pressure. In contrast, when cultivated in chemostat with recycle under the same r- and K-conditions imposed on chemostat cultures, distribution patterns of r- and K-selected populations appeared very little affected by changes in substrate availability.     

Correlation between Olfactory Responses, Dispersal Tendencies, and Life-history Traits of the Predatory Mite Neoseiulus Womersleyi (Acari: Phytoseiidae) of Eight Local Populations

To investigate the relationship between foraging behavior and life-history traits of the predatory mite Neoseiulus womersleyi, the olfactory responses, dispersal ratios from a prey patch, predation rates, fecundity, and developmental times in eight local populations of N. womersleyi were investigated. Significant differences among local populations were found in all these traits except fecundity. None of the life-history traits correlated with foraging behavior. A significant positive correlation was found only between the olfactory response and the dispersal ratio. These results suggested that predatory mites with low olfactory responses would stay in a prey patch longer than predatory mites with high olfactory responses.     

HYBRID ZONE DYNAMICS ARE INFLUENCED BY GENOTYPE-SPECIFIC VARIATION IN LIFE-HISTORY TRAITS: EXPERIMENTAL EVIDENCE FROM HYBRIDIZING GAMBUSIA SPECIES

Results from two experiments are presented that contrast differences in life-history traits and population dynamics between two species of live bearing fishes (Gambusia affinis and G. holbrooki) that hybridize across portions of the southeastern United States. Progeny from parental holbrooki and holbrooki-affinis F₁ crosses exhibited larger lengths at birth, at 15 days, and matured earlier, and at larger size than did progeny from parental affinis and affinis-holbrooki F₁ crosses. Comparisons of experimental populations of affinis, holbrooki, and mixed (affinis + holbrooki) species composition followed over two years revealed that affinis populations consistently exhibited smaller population size, lower carrying capacity, lower recruitment, and larger over-winter mortality than did holbrooki or mixed populations. Evidence for density-dependent reductions in fecundity and concomitant increases in juvenile mortality rates were observed in all populations, but were most pronounced for affinis populations. Genotype-specific differences in life-history traits appear to confer differential advantage to offspring of parental holbrooki origin and F, progeny of holbrooki maternal parentage given the resource availability and the age structure and densities experienced during these experiments. Results have direct implications regarding the rate and direction of evolution within hybrid zones formed by these two species.     

ON THE LOW HERITABILITY OF LIFE-HISTORY TRAITS

Life-history traits such as longevity and fecundity often show low heritability. This is usually interpreted in terms of Fisher's fundamental theorem to mean that populations are near evolutionary equilibrium and genetic variance in total fitness is low. We develop the causal relationship between metric traits and life-history traits to show that a life-history trait is expected to have a low heritability whether or not the population is at equilibrium. This is because it is subject to all the environmental variation in the metric traits that affect it plus additional environmental variation. There is no simple prediction regarding levels of additive genetic variance in life-history traits, which may be high at equilibrium. Several other patterns in the inheritance of life-history traits are readily predicted from the causal model. These include the strength of genetic correlations between life-history traits, levels of nonadditive genetic variance, and the inevitability of genotype-environment interaction.     

Life-history phenotypes in populations of Brachyrhaphis episcopi (Poeciliidae) with different predator communities

Variation among populations in extrinsic mortality schedules selects for different patterns of investment in key life-history traits. We compared life-history phenotypes among 12 populations of the live-bearing fish Brachyrhaphis episcopi. Five populations co-occurred with predatory fish large enough to prey upon adults, while the other seven populations lacked these predators. At sites with large predatory fish, both sexes reached maturity at a smaller size. Females of small to average length that co-occurred with predators had higher fecundity and greater reproductive allotment than those from populations that lacked predators, but the fecundity and reproductive allotment of females one standard deviation larger than mean body length did not differ among sites. In populations with large predatory fish, offspring mass was significantly reduced. In each population, fecundity, offspring size and reproductive allotment increased with female body size. When controlling for maternal size, offspring mass and number were significantly negatively correlated, indicating a phenotypic trade-off. This trade-off was non-linear, however, because reproductive allotment still increased with brood size after controlling for maternal size. Similar differences in life-history phenotypes among populations with and without large aquatic predators have been reported for Brachyrhaphis rhabdophora in Costa Rica and Poecilia reticulata (a guppy) in Trinidad. This may represent a convergent adaptation in life-history strategies attributable to predator-mediated effects or environmental correlates of predator presence.     

Does artificial selection on diapause incidence cause correlational changes in other life‐history traits? Case study in a spider mite population

Theoretical studies suggest that the timing of entering hibernation by arthropods has large effects on long-term fitness, incurring strong selection pressure on diapause attributes every year. On the other hand, diapause attributes are often genetically correlated with other important life-history traits such as fecundity or development time. To understand the evolutionary process of life cycle formation, there is a need to investigate not only diapause attributes themselves but also their genetic association with other life-history traits. The Kanzawa spider mite, Tetranychus kanzawai Kishida (Acari: Tetranychidae), is a small herbivore that lives on the undersurface of host plant leaves. This mite has been investigated for the mode of inheritance of diapause attributes, but scarcely for genetic correlations with other life-history traits. Here, I investigated whether diapause proneness, measured as the proportion of diapausing females under short-day conditions, is genetically correlated with fecundity or development time under long-day conditions using artificial selection experiments. Diapause incidence responded to the selection for both increasing and decreasing directions, suggesting that high genetic variance in diapause proneness is maintained in the study population. However, the change in proportion of diapausing females during the selection period was not associated with responses in fecundity or development time. These results suggest that diapause proneness and other life-history traits have different genetic backgrounds, and thus diapause proneness may freely evolve without being constrained by changes in other life-history traits.     

Maturation,growth and fecundity of Arctic charr, <Emphasis Type="Italic">Salvelinus alpinus</Emphasis> (L.), life-history variants co-existing in lake systems of Southern Baffin Island,Nunavut, Canada

Two life-history variants of Arctic charr (Salvelinus alpinus), anadromous and lake-resident, have been previously identified in lakes of Southern Baffin Island, Nunavut, Canada. In accordance with classical life-history theory, it is hypothesised that anadromous charr will delay maturation in both size and age, and have increased fecundity (per spawning event) in comparison with lake-resident charr. Sagittal otoliths and biological data were collected for both life-history variants within the three studied lakes: Iqalugaarjuit, Qasigiat and Qinngu. Sagittal otoliths were embedded in epoxy resin, cross-sectioned for age determination, and imaged for back-calculation (size and age). Back-calculated data in each lake were fit to von Bertalanffy growth models for each life-history variant and compared via analysis of residual sums of squares. Anadromous charr had greater mean size at maturity and experienced a delay in mean age at maturity in comparison with lake-residents. The relationship between size and fecundity or egg diameter did not differ between the two life-history variants. Growth models indicate that the overall growth coefficients of lake-resident and anadromous charr were different in all three studied lakes. The Brody growth coefficient for all lake-resident charr populations was greater than anadromous individuals indicating that maximum length was reached at a rapid rate, resulting in a smaller asymptotic length. Indirect evidence suggests that anadromous and lake-resident charr belong to one reproductive population. Future genetic analysis is necessary to further determine the degree of divergence between the life-history variants.     

Trade-offs in the life history and energy budget of the parthenogenetic collembolan Folsomia candida (Willem)

Three clones of Folsomia candida from different locations in Europe were compared in four experiments investigating genetic and phenotypic correlations between life-history traits. The first three experiments focused on the effects of food type, clone and temperature on traits associated with the first clutch. Differences in clutch size between clones and treatments were almost completely attributable to body size. Clones differed in length of the juvenile period, but the difference decreased at low temperatures. Age and weight at first reproduction were negatively correlated in the food type experiment and positively correlated in the temperature experiment, an often-encountered result for which no general explanation is as yet available. In the temperature experiment egg size variation was considerable, and was highest at low temperatures. The fourth experiment, with two clones at two feeding levels, aimed at finding trade-offs, in particular between reproduction and survival. It was hypothesized that higher fecundity led to increased scenescence through a higher metabolic rate. The trade-off was clearly present among the clones: one combined fast growth, late reproduction and high lifetime fecundity with lower survival, while in the other the relation between these traits was opposite. The proposed mechanism, however, was not confirmed, as no difference in metabolic rate was found. The effect of food level was too small to result in significant differences in the life-history traits in either of the clones.     

High Wolbachia density correlates with cost of infection for insecticide resistant Culex pipiens mosquitoes

In the mosquito Culex pipiens, insecticide resistance genes alter many life-history traits and incur a fitness cost. Resistance to organophosphate insecticides involves two loci, with each locus coding for a different mechanism of resistance (degradation vs. insensitivity to insecticides). The density of intracellular Wolbachia bacteria has been found to be higher in resistant mosquitoes, regardless of the mechanism involved. To discriminate between costs of resistance due to resistance genes from those associated with elevated Wolbachia densities, we compared strains of mosquito sharing the same genetic background but differing in their resistance alleles and Wolbachia infection status. Life-history traits measured included strength of insecticide resistance, larval mortality, adult female size, fecundity, predation avoidance, mating competition, and strength of cytoplasmic incompatibility (CI). We found that: (1) when Wolbachia are removed, insecticide resistance genes still affect some life-history traits; (2) Wolbachia are capable of modifying the cost of resistance; (3) the cost of Wolbachia infections increases with their density; (4) different interactions occurred depending on the resistance alleles involved; and (5) high densities of Wolbachia do not increase the strength of CI or maternal transmission efficiency relative to low Wolbachia densities. Insecticide resistance genes generated variation in the costs of Wolbachia infections and provided an interesting opportunity to study how these costs evolve, a process generally operating when Wolbachia colonizes a new host.     

Experimental evolution of parasite life-history traits in Strongyloides ratti (Nematoda)

Evolutionary ecology predicts that parasite life-history traits, including a parasite's survivorship and fecundity within a host, will evolve in response to selection and that their evolution will be constrained by trade-offs between traits. Here, we test these predictions using a nematode parasite of rats, Strongyloides ratti, as a model. We performed a selection experiment by passage of parasite progeny from either early in an infection ('fast' lines) or late in an infection ('slow' lines). We found that parasite fecundity responded to selection but that parasite survivorship did not. We found a trade-off mediated via conspecific density-dependent constraints; namely, that fast lines exhibit higher density-independent fecundity than slow lines, but fast lines suffered greater reduction in fecundity in the presence of density-dependent constraints than slow lines. We also found that slow lines both stimulate a higher level of IgG1, which is a marker for a Th2-type immune response, and show less of a reduction in fecundity in response to IgG1 levels than for fast lines. Our results confirm the general prediction that parasite life-history traits can evolve in response to selection and indicate that such evolutionary responses may have significant implications for the epidemiology of infectious disease.     

Effects of founding genetic variation on adaptation to a novel resource

Population genetic theory predicts that adaptation in novel environments is enhanced by genetic variation for fitness. However, theory also predicts that under strong selection, demographic stochasticity can drive populations to extinction before they can adapt. We exposed wheat-adapted populations of the flour beetle (Tribolium castaneum) to a novel suboptimal corn resource, to test the effects of founding genetic variation on population decline and subsequent extinction or adaptation. As previously reported, genetically diverse populations were less likely to go extinct. Here, we show that among surviving populations, genetically diverse groups recovered faster after the initial population decline. Within two years, surviving populations significantly increased their fitness on corn via increased fecundity, increased egg survival, faster larval development, and higher rate of egg cannibalism. However, founding genetic variation only enhanced the increase in fecundity, despite existing genetic variation-and apparent lack of trade-offs-for egg survival and larval development time. Thus, during adaptation to novel habitats the positive impact of genetic variation may be restricted to only a few traits, although change in many life-history traits may be necessary to avoid extinction. Despite severe initial maladaptation and low population size, genetic diversity can thus overcome the predicted high extinction risk in new habitats.