A comparison of nest building by three-spined sticklebacks Gasterosteus aculeatus from still and flowing waters

The composition of nests, built under standardized, static laboratory conditions by male three-spined sticklebacks Gasterosteus aculeatus from two rivers and two lakes in mid-Wales (U.K.) was quantified. After transfer to nesting tanks, fish from the rivers began building nests more quickly than fish from the lakes. The number of threads used in nest construction and the mass of substrate deposited co-varied, but there were no consistent population- or habitat-related differences in the aspects of nest composition.     

The evolution of reaction norms: simple models for age and size at maturity

This paper presents a simple model for the evolution of reaction norms for age and size at maturity that predicts reaction norms with a variety of shapes. Using realistic parameter values the model predicts reaction norms close to those observed in Drosophila. The major assumptions of the model are: 1) that net reproductive rate is maximized, 2) that growth is determinate, and 3) that mortality rates are independent of age and size at maturity. If, additionally, juvenile mortality is uncorrelated with a growth coefficient, k, the model predicts that selection favors maturation later at a smaller size when k is reduced by environmental factors and that decreased juvenile mortality leads to delayed maturity. These two predictions conform with those found by previous models using other measures of fitness. Correlations between k and juvenile mortality can change the shape of the predicted reaction norm. Depending on the precise form of the correlation, the model can predict done- or bowl-shaped reaction norms and can predict delayed or earlier maturity as k decreases. These shapes are qualitatively different from those predicted by previous models that used different fitness measures. Systematic estimates of the parameter values for this and for related models are required to determine the appropriate fitness measure for models of reaction norms.     

How feeding performance and energy intake change with a small increase in the body size of the three-spined stickleback

Changes in the foraging behaviour due to variation in the body size of the three-spined stickleback Gasterosteus aculeatus were investigated. All sizes of fish had a high probability of attacking prey whenever encountered. The probability of eating the prey increased with the size of the fish, as the larger fish had larger jaws and a greater stomach capacity. Therefore, as fish increased in size there was an increase in the probability of successful prey capture. The level of satiation did not have an effect on the prey handling time, which is contrary to other studies and is probably a result of the large prey sizes. The physical size of the prey meant that the handling times were long regardless of the motivational level of the fish. The larger fish took in more energy and at a faster rate, although the time to reach satiation was similar for all fish sizes. The advantage that large fish appear to have in successfully gaining large prey is negated by their greater metabolic requirement. The changes in feeding performance induced by small increases in body size could have important consequences for intraspecific competition, habitat Use and risk of predation.     

Eutrophication and predation risk interact to affect sexual trait expression and mating success

Sexual traits are especially sensitive to low food resources. Other environmental parameters (e.g., predation) should also affect sexual trait expression by favoring investment in viability traits rather than sexual traits. We know surprisingly little about how predators alter investment in sexual traits, or how predator and resource environments interact to affect sexual trait investment. We explored how increasing phosphorous (P) availability, at a level mimicking cultural eutrophication, affects the development of sexual, nonsexual, and viability traits of amphipods in the presence and absence of predators. Sexual traits and growth were hypersensitive to low P compared to nonsexual traits. However, a key sexual trait responded to low P only when predator cues were absent. Furthermore, investment trade-offs between sexual traits and growth only occurred when P was low. The phenotypic changes caused by predator cues and increased P availability resulted in higher male mating success. Thus, eutrophication not only affects sexual trait expression but also masks the trade-off between traits with similar P demand. Sensitivity of sexually selected traits to changes in P, combined with the important roles these traits play in determining fitness and driving speciation, suggests that human-induced environmental change can greatly alter the evolutionary trajectories of populations.     

Changes in size and age at maturity in a population of kokanee Oncorhynchus nerka during a period of declining growth conditions

Trends in size distributions and age at maturity of spawning kokanee Oncorhynchus nerka during a 5 year period of declining growth conditions at Bucks Lake, California, U.S.A. were consistent with the hypothesis that reductions in growth rates in successive cohorts induce a shift to an older age at maturity. This forestalls decreases in size at maturity during a transitional period characterized by an increasing proportion of individuals that delay maturation. During the course of the study, kokanee first began declining in size at maturity, and then shifted from a 3 year to a 4 year egg to adult cycle. Individuals that spawned during their fourth year (age 3 years) were significantly larger, on average, than members of their cohort that spawned during their third year (age 2 years). This difference was greatest when age 2 year adults were smallest. The shift to an older age at maturity prevented a steady decline in size at maturity, even though age‐specific size was steadily declining over time. Size at maturity, however, began to decline again once the transition to a 4 year cycle was complete. In addition, there was a general trend of decreasing length‐specific mass. The data indicate that there is a range of growth trajectories over which delayed maturity can prevent a temporal pattern of decreasing size at maturity as growth rates decline.     

Phenotypic plasticity in age and size at maturity and its effects on the integrated phenotypic expressions of life history traits of Cardamine flexuosa (Cruciferae)

We analyzed variation in phenotypic plasticity of life history traits between two Cardamine flexuosa populations based on differences in plasticity of age and size at maturity. C. flexuosa (Cruciferae) is a facultative, vernalization-sensitive, long-day annual, and its phenology and the phenotypic expressions of many life history traits are largely controlled by photoperiod and vernalization in natural populations. We used plants from two populations which differed in their responses to chilling and photoperiod treatments. The timing of developmental processes was changed by controlling temperature and photoperiod regimes in growth chambers. Plasticity in size at maturity was analyzed as changes in a growth trajectory using two parameters, age at maturity (Δt) and growth rate (k). Both traits showed plasticity, but differences between the populations were found mostly for Δt. Distinctive differences in size at maturity of individuals in the two populations were mainly due to different amounts of plasticity in Δt. Variations in plasticity of nine other life history traits and their associations to age and size at maturity were also analyzed. Variation for eight of the traits can be described, at least in part, as a function of age and size at maturity for both populations, and most of the variation in the total number of seeds was explained by age and size at maturity. Only age at maturity had any effect on changes in resource allocation. The nine life history traits were integrated through associated character expressions with age and size at maturity. Changes in the association between a trait and age and/or size at maturity were rather conservative compared to changes in the plasticity of a trait between the two populations. Associations with age and size at maturity are mostly explicable in terms of inherent relationships in the developmental processes, and they may limit the ecological range expansion and the adaptive evolution of plasticity in C. flexuosa. The negative correlation between reproductive allocation and age at maturity can be a cost of delaying maturation in C. flexuosa.     

Parasites may contribute to 'magic trait' evolution in the adaptive radiation of three-spined sticklebacks, Gasterosteus aculeatus (Gasterosteiformes: Gasterosteidae)

In the past decade, there has been a new effort to understand the ecology that drives population divergence and speciation. It is well established in theory that speciation is most likely to occur when a trait that is under divergent natural selection in different populations is also used in mate choice. Such traits have been dubbed 'magic traits' ( Gavrilets, 2004 ) and, although there appears to be good evidence that they exist, the ecological mechanisms that underlie their divergence are not well understood. Size at maturity in three-spined sticklebacks is an archetypal example of a magic trait. The present study documents for the first time that differences in body size at maturity in sympatric species pairs of lacustrine three-spined sticklebacks in British Columbia, Canada, are caused by differences in age at maturity. It is also shown that there are differences between the sympatric species in the patterns of infection with a virulent cestode, Schistocephalus solidus . Although the evidence is circumstantial, these differences in infection are consistent with the hypothesis that they have contributed to the observed divergence in age and size at maturity in these populations.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 425–433.     

Juvenile exposure to predator cues induces a larger egg size in fish

When females anticipate a hazardous environment for their offspring, they can increase offspring survival by producing larger young. Early environmental experience determines egg size in different animal taxa. We predicted that a higher perceived predation risk by juveniles would cause an increase in the sizes of eggs that they produce as adults. To test this, we exposed juveniles of the mouthbrooding cichlid Eretmodus cyanostictus in a split-brood experiment either to cues of a natural predator or to a control situation. After maturation, females that had been confronted with predators produced heavier eggs, whereas clutch size itself was not affected by the treatment. This effect cannot be explained by a differential female body size because the predator treatment did not influence growth trajectories. The observed increase of egg mass is likely to be adaptive, as heavier eggs gave rise to larger young and in fish, juvenile predation risk drops sharply with increasing body size. This study provides the first evidence that predator cues perceived by females early in life positively affect egg mass, suggesting that these cues allow her to predict the predation risk for her offspring.     

Divergent phenotypic responses to predators and cyanobacteria in Daphnia lumholtzi

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Maturation patterns in female European eel: age and size at the silver eel stage

The maturation pattern in the female European eel Anguilla anguilla was studied by investigating age and size patterns of silver eels in different aquatic environments in Sweden, covering limnic, brackish and marine waters. The results neither supported the hypothesis that there is a critical size or age when eels enter the silvery stage, nor that size and age at maturity are positively related. Age at maturity, however, was observed to be negatively related to growth rate in all localities, i.e. the female reproductive tactic apparently is to become sexually mature at the earliest possible opportunity. Furthermore, it was recognized that a significant amount of variation was due to habitat differences, since the female eel maturation pattern deviated systematically between sampling sites, as it did also when the effect of growth rate was eliminated. Thus, the ability of the female eel to adjust maturation to an optimal size and age can be questioned, because the panmictic nature of the eel means local adaptations are unlikely Growth rate dependent differences suggest that variations in maturation patterns between eel environments are linked more to the opportunity for nutrient accumulation than to other aspects of growth.     

The effects of perceived predation risk on pre- and post-metamorphic phenotypes in the common frog

Where organisms undergo radical changes in habitat during ontogeny, dramatic phenotypic reshaping may be required. However, physiological and functional interrelationships may constrain the extent to which an individual's phenotype can be equally well adapted to their habitat throughout the life cycle. The phenotypic response of tadpoles to the presence of a predator has been reported for several species of anuran but the potential post-metamorphic consequences have rarely been considered. We reared common frog Rana temporaria tadpoles in the presence or absence of a larval odonate predator, Aeshna juncea , and examined the consequences of the resulting phenotypic adjustment in the aquatic larval stage of the life cycle for the terrestrial juvenile phenotype. In early development tadpoles developed deeper tail fins and muscles in response to the predator and, in experimental trials, swam further than those reared in the absence of a predator. While the difference in swimming ability remained significant throughout the larval period, by the onset of metamorphosis we could no longer detect any differences in the morphological parameters measured. The corresponding post-metamorphic phenotypes also did not initially differ in terms of morphology. At 12 weeks post-metamorphosis, however, froglets that developed from predator-exposed tadpoles swam more slowly and less far than those that developed from tadpoles reared in the absence of predators, the opposite trend to that observed in the larval stage of the life cycle, and had narrower femurs. These results suggest that there may be long-term costs for subsequent life-history stages of tailoring the larval phenotype to prevailing environmental conditions.     

Measuring probabilistic reaction norms for age and size at maturation

We present a new probabilistic concept of reaction norms for age and size at maturation that is applicable when observations are carried out at discrete time intervals. This approach can also be used to estimate reaction norms for age and size at metamorphosis or at other ontogenetic transitions. Such estimations are critical for understanding phenotypic plasticity and life-history changes in variable environments, assessing genetic changes in the presence of phenotypic plasticity, and calibrating size- and age-structured population models. We show that previous approaches to this problem, based on regressing size against age at maturation, give results that are systematically biased when compared to the probabilistic reaction norms. The bias can be substantial and is likely to lead to qualitatively incorrect conclusions; it is caused by failing to account for the probabilistic nature of the maturation process. We explain why, instead, robust estimations of maturation reaction norms should be based on logistic regression or on other statistical models that treat the probability of maturing as a dependent variable. We demonstrate the utility of our approach with two examples. First, the analysis of data generated for a known reaction norm highlights some crucial limitations of previous approaches. Second, application to the northeast arctic cod (Gadus morhua) illustrates how our approach can be used to shed new light on existing real-world data.     

HOW TO MEASURE MATURATION: A COMPARISON OF PROBABILISTIC METHODS USED TO TEST FOR GENOTYPIC VARIATION AND PLASTICITY IN THE DECISION TO MATURE

Maturation is a developmental trait that plays a key role in shaping organisms’ life‐history. However, progress in understanding how maturation phenotypes evolve has been held back by confusion over how best to model maturation decisions and a lack of studies comparing genotypic variation in maturation. Here, we fitted probabilistic maturation reaction norms (PMRNs) to data collected from five clones of Daphnia magna and five of Daphnia pulex collected from within and between different populations. We directly compared the utility of modeling approaches that assume maturation to be a process with an instantaneous rate with those that do not by fitting maturation rate and logistic regression models, and emphasize similarities and differences between them. Our results demonstrate that in Daphnia, PMRNs using a logistic regression approach were simpler to use and provided a better fit to the data. The decision to mature was plastic across a range of growth trajectories and dependent upon both body size and age. However, the age effect was stronger in D. magna than D. pulex and varied considerably between clones. Our results support the idea that maturation thresholds can evolve but also suggest that the notion of a threshold based on a single fixed state is an oversimplification that underestimates the adaptability of these important traits.     

Life-history variations in the fluvial sculpin, Cottus nozawae (Cottidae), along the course of a small mountain stream

Life-history variations in male and female fluvial sculpins, Cottus nozawae, were studied in a small mountain stream in Hokkaido, Japan, primarily by using capture-mark-recapture methods. At three study areas established along the stream course, the majority of marked sculpins were recaptured in their original location over one or more years, indicating their long-term occupation of each restricted habitat area. Sculpin densities increased toward the upstream habitats, whereas individual growth rates were more rapid downstream. In both sexes, sculpins distributed downstream matured at a larger body size and later in life than upstream sculpins, clearly demonstrating a clinal variation in these respects. A comparison of life-history variations in C. nozawae with those in amphidromous C. hangiongensis suggests that intrapopulational life-history variations in the former might be environmentally induced, and that one of the most important determinants for the variations in Cottus species might be population density.     

Sex-specific plasticity of growth and maturation size in a spider: implications for sexual size dimorphism

Sex-specific plasticity in body size has been recently proposed to cause intraspecific patterns of variation in sexual size dimorphism (SSD). We reared juvenile male and female Mediterranean tarantulas (Lycosa tarantula) under two feeding regimes and monitored their growth until maturation. Selection gradients calculated across studies show how maturation size is under net stabilizing selection in females and under directional selection in males. This pattern was used to predict that body size should be more canalized in females than in males. As expected, feeding affected male but not female maturation size. The sex-specific response of maturation size was related to a dramatic divergence between subadult male and female growth pathways. These results demonstrate the existence of sex-specific canalization and resource allocation to maturation size in this species, which causes variation in SSD depending on developmental conditions consistent with the differential-plasticity hypothesis explaining Rensch's Rule.     

The evolution of life histories in spatially heterogeneous environments: Optimal reaction norms revisited

Summary Natural populations live in heterogeneous environments, where habitat variation drives the evolution of phenotypic plasticity. The key feature of population structure addressed in this paper is the net flow of individuals from source (good) to sink (poor) habitats. These movements make it necessary to calculate fitness across the full range of habitats encountered by the population, rather than independently for each habitat. As a consequence, the optimal phenotype in a given habitat not only depends on conditions there but is linked to the performance of individuals in other habitats. We generalize the Euler-Lotka equation to define fitness in a spatially heterogeneous environment in which individuals disperse among habitats as newborn and then stay in a given habitat for life. In this case, maximizing fitness (the rate of increase over all habitats) is equivalent to maximizing the reproductive value of newborn in each habitat but not to maximizing the rate of increase that would result if individuals in each habitat were an isolated population. The new equation can be used to find optimal reaction norms for life history traits, and examples are calculated for age at maturity and clutch size. In contrast to previous results, the optimal reaction norm differs from the line connecting local adaptations of isolated populations each living in only one habitat. Selection pressure is higher in good and frequent habitats than in poor and rare ones. A formula for the relative importance of these two factors allows predictions of the habitat in which the genetic variance about the optimal reaction norm should be smallest.     

Adaptive life-history evolution in the livebearing fish Brachyrhaphis rhabdophora: genetic basis for parallel divergence in age and size at maturity and a test of predator-induced plasticity

I document a genetic basis for parallel evolution of life-history phenotypes in the livebearing fish Brachyrhaphis rhabdophora from northwestern Costa Rica. In previous work, I showed that populations of B. rhabdophora that co-occur with predators attain maturity at smaller sizes than populations that live in predator-free environments. I also demonstrated that this pattern of phenotypic divergence in life histories was independently repeated in at least five isolated drainages. However, life-history phenotypes measured from wild-caught fish could be attributed to environmental effects rather than to genetic differences among populations. In the present study, I reared male fish from four populations (two that co-occur with predators and two from predator-free environments) under four sets of environmental conditions. The pattern of phenotypic divergence in maturation size documented in the field between populations collected from different predation environments persisted after two generations in the laboratory. I also found a genetic basis for differences between populations in the age at which males attain maturity and in growth rates. By rearing fish in four different common environments, I tested for phenotypic plasticity in male life-history traits in response to nonlethal exposure to predators. There was a significant delay in the onset of sexual maturity in fish exposed to predators relative to those in the control, but no differences among treatments in size at maturity or growth rates. These results, coupled with previous work on B. rhabdophora, demonstrate a repeated pattern of parallel evolutionary divergence among genetically isolated populations that is strongly associated with predation.     

Correlated responses to artificial body size selection in growth, development, phenotypic plasticity and juvenile viability in yellow dung flies

Most life history traits are positively influenced by body size, whereas disadvantages of large body size are poorly documented. To investigate presumed intrinsic costs of large size in the yellow dung fly (Scathophaga stercoraria; Diptera: Scathophagidae), we established two replicates each of three body size laboratory selection lines (small, control and large; selection on males only), and subjected flies of the resulting extended body size range to various abiotic stresses. Response to selection was symmetrical in the small and large lines (realized h(2) = 0.16-0.18). After 24 generations of selection body size had changed by roughly 10%. Female size showed a correlated response to selection on male size, whereas sexual size dimorphism did not change. Development time also showed a correlated response as, similar to food limited flies, small line flies emerged earlier at smaller body size. At the lowest larval food limit possible, flies of all lines emerged at the same small body size after roughly the same development time; so overall phenotypic plasticity in body size and development time strongly increased following selection. Juvenile mortality increased markedly when food was extremely limited, large line flies showing highest mortality. Winter frost disproportionately killed large (line) flies because of their longer development times. Mortality at high temperatures was high but size-selective effects were inconsistent. In all environments the larger males suffered more. Initial growth rate was higher for males and at unlimited food. Small line individuals of both sexes grew slowest at unlimited larval food but fastest at limited larval food, suggesting a physiological cost of fast growth. Overall, extension of the natural body size range by artificial selection revealed some otherwise cryptic intrinsic juvenile viability costs of large size, mediated by longer development or faster growth, but only in stressful environments.