Life-history plasticity in the butterfly Lycaena hippothoe: local adaptations and trade-offs

Life-history theory predicts some cost to be associated with short development time, the most frequently assumed being small adult size. Alternatively, insects may increase developmental rates and grow fast to a larger size. Seasonal environments should select for phenotypic plasticity in growth and development, based on the need to complete development up to the diapausing stage before the onset of unfavourable season. Nevertheless, there must be some limit beyond which a compensation for a shorter development cannot be achieved. By comparing three geographically isolated populations of Lycaena hippothoe in common environments we show that in the Hungarian population development time seems to be traded off against size at maturity. This population is the only bivoltine one within this principally monovoltine species. Thus, realization of an additional generation per year, achieved through largely reduced development times, appears to carry the cost of substantially lower adult weights compared with other populations. In contrast, differences in development time in two monovoltine populations were not accompanied by a trade-off between development time and size. These results suggest that clear trade-offs are restricted to stressful situations, when compensation by an increase in growth rates is no longer feasible. We suggest the particularly short development time in the Hungarian population (facilitating a second generation), as well as the shorter development in an alpine (short vegetation period) compared with a western German population, to be adaptations to local climatic conditions. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 173–185.     

Correlated contemporary evolution of life history traits in New Zealand Chinook salmon, Oncorhynchus tshawytscha

Size at age and age at maturity are important life history traits, affecting individual fitness and population demography. In salmon and other organisms, size and growth rate are commonly considered cues for maturation and thus age at maturity may or may not evolve independently of these features. Recent concerns surrounding the potential phenotypic and demographic responses of populations facing anthropogenic disturbances, such as climate change and harvest, place a premium on understanding the evolutionary genetic basis for evolution in size at age and age at maturity. In this study, we present the findings from a set of common-garden rearing experiments that empirically assess the heritable basis of phenotypic divergence in size at age and age at maturity in Chinook salmon (Oncorhynchus tshawytscha) populations introduced to New Zealand. We found consistent evidence of heritable differences among populations in both size at age and age at maturity, often corresponding to patterns observed in the wild. Populations diverged in size and growth profiles, even when accounting for eventual age at maturation. By contrast, most, but not all, cases of divergence in age at maturity were driven by the differences in size or growth rate rather than differences in the threshold relationship linking growth rate and probability of maturation. These findings help us understand how life histories may evolve through trait interactions in populations exposed to natural and anthropogenic disturbances, and how we might best detect such evolution.     

The quantitative genetics of growth in a field cricket

Because of its relationship with both development time and adult size, the rate of growth in determinately growing organisms is an important aspect of their life histories. We reared sixty-nine families of Gryllus pennsylvanicus derived from a natural population and found significant genetic variation in growth rate as estimated by the slope of linearized growth trajectories. We found no evidence for a genetic tradeoff between rate of growth and survival, nor rate of growth and fecundity. In principle, adult size may be determined both by the rate of growth and the time taken by the nymphs to develop. Our data indicate that variation in adult size is explained by variation in growth rate, not by variation in development time. We conclude with a discussion of the plausible explanations for the presence of genetic variation in growth rate in this natural population.     

Reaction norms for developmental time and weight at eclosion in Drosophila mercatorum

We determined reaction norms for developmental time and weight at eclosion for 2 isozygous and 11 genetically mixed strains of Drosophila mercatorum in four culture media differing in yeast concentration. With decreasing yeast concentration, development was delayed, the weight of emerging flies decreased, and the phenotypic variance of both variables increased. Differences among stocks and significant stock × yeast interactions indicated genetic variance for both variables within environment and different phenotypic responses of stocks across environments. The phenotypic correlation between developmental time and weight was negative at low yeast concentrations and disappeared gradually with increasing yeast. The comparison of completely homozygous with genetically heterogenous stocks showed that most of the increase of variability with deteriorating environment was due to the changing expression of genetic variance. The genetic correlation between developmental time and weight turned from negative in poor to positive in rich medium, while the environmental covariance was negative in all media. Plotting the reaction norms in the developmental time-weight plane rather than separately for each trait reveals most of these results at a glance. It also suggests that much of the genetic variance might be additive, because an effect of the half-sib family structure inherent in the design is clearly visible in the plot. We interpret the pattern of changing variances and covariances, pointing out that the special growth physiology of Drosophila and the way environmental factors affect it must be taken into account. We briefly discuss the implications of changing genetic correlations among traits for the evolution of phenotypic plasticity in general.     

Reaction norms for age and size at maturity in response to temperature: a test of the compound interest hypothesis

In ectotherms, temperature induces similar developmental and evolutionary responses in body size, with larger individuals occurring or evolving in low temperature environments. Based on the occasional occurrence of opposite size clines, showing a decline in body size with increasing latitude, an interaction between generation time and growing season length was suggested to account for the patterns found. Accordingly, multivoltine species with short generation times should gain high compound interest benefits from reproducing early at high temperatures, indicating potential for extra generations, even at the expense of being smaller. This should not apply for obligatorily monovoltine populations. We explicitly test the prediction that monovoltine populations (no compound interest) should be selected for large body size to maximise adult fitness, and therefore size at maturity should respond only weakly to temperature. In two monovoltine populations (an Alpine and a Western German one) of the butterfly Lycaena hippothoe, increasing temperatures had no significant effect on pupal weight and caused a slight decrease in adult weight only. In contrast, two closely related, yet potentially multivoltine Lycaena populations showed a greater weight loss at increasing temperature (in protandrous males, but not in females) and smaller adult sizes throughout. Thus, the results do support our predictions indicating that the compound interest hypothesis may yield causal explanations for the relationship between temperature and insect size at maturity. At all temperatures, the alpine population had higher growth rates and concomitantly shorter development times (not accompanied by a reduction in size) than the other, presumably indicating local adaptations to different climates.     

Per-stage growth rates of head and body sizes in scarab larvae (Coleoptera: Scarabaeidae) decrease across ontogeny following a species-specific pattern

It has been widely assumed that the stepwise increase in the exoskeleton size of larval insects approximately follows a geometric progression from instar to instar, known as Dyar's Rule. However, it is not clear whether the per-instar increase in body size follows this rule. In insects, Dyar's Rule has been identified either by regressing the log-scaled size on the instar number (log-linear regression analysis) or by comparing the postmolt/premolt size ratio between instars (growth rate analysis). A previous study on the body mass of caterpillars showed the methodological pitfall that Dyar's Rule was statistically supported by log-linear regression analysis, but not at all by growth rates analysis. I considered this concern here by examining the per-stage growth rates of head and body sizes for larvae of the beetle Trypoxylus dichotomus using both methods and compared the resulting growth rates for body size within and between taxonomic orders. Dyar's Rule was statistically supported by the log-linear regression analysis but not by growth rate analysis for both the head and body sizes in T. dichotomus. The body size growth rate in T. dichotomus decreased as the instar progressed. This developmental pattern was also found in reported data for the other six scarabs, but not in data for Lepidoptera or Hymenoptera. These findings confirm that the per-stage growth rate of body size does not follow Dyar's Rule in a wide range of insects, and suggest that developmental change in the body size growth rate varies among insect groups.     

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.     

Annual variations in the reproductive traits of Pomatoschistus microps in a Mediterranean lagoon undergoing environmental changes: evidence of phenotypic plasticity

The spawning period of the common goby Pomatoschistus microps from 1993 to 1997 in the Vaccarès lagoon did not vary, except in 1997 when it was longer due to the reproduction of the young-of-the-year. Egg size and number, and reproductive allocation varied greatly with one year to another. Female common gobies increased both their fecundity per spawning act and their egg size from 1993 to 1995. The annual variation in the reproductive effort suggests a high phenotypic plasticity of reproductive traits in P. microps , in the face of environmental perturbations. In winter 1993–1994, a centennial flood of the Rhône River caused major hydrological changes in the lagoon in less than 1 week, affecting many invertebrates and fish for several years. The reproductive investment of the common goby increased, possibly as a consequence of those environmental changes.     

Habitat-specific differences in thermal plasticity in natural populations of a soil arthropod

When populations experience substantial variation in environmental conditions, they may evolve phenotypic plasticity in response to these varying selection pressures. Evolutionary theory predicts differentiation in the level of phenotypic plasticity among different habitats. We evaluated temperature-induced phenotypic responses in juvenile growth rate in natural populations of the springtail Orchesella cincta , inhabiting forest and heathland. These habitats typically co-occur but differ strongly with respect to, for example, thermal regime, relative humidity, and structure. Offspring of females from the two habitats were reared at different temperatures in climate rooms and the temperature response of juvenile growth rate and egg size was measured. We found a habitat-specific difference in plasticity of juvenile growth rate. The reaction norms of the forest populations were steeper than the reaction norms for heath populations at two replicated sampling sites. Egg weight itself was demonstrated to be a plastic trait with a higher egg weight at low temperatures, but the thermal response did not differ between habitats. We conclude that these populations have diverged due to strong local natural selection. Our results support the argument that the level of phenotypic plasticity itself can be under selection and that differentiation in reaction norms can occur even in neighbouring habitats with no barrier to gene flow.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 265–271.     

Proximate causes of sexual size dimorphism in Colorado pikeminnow, a long‐lived cyprinid

Evidence for sexual size dimorphism (SSD) and its possible causes were examined in the endangered Colorado pikeminnow Ptychocheilus lucius, a large, piscivorous, cyprinid endemic to the Colorado River system of North America. Individuals representing 18–24% of the upper Colorado River population were captured, measured, sexed and released in 1999 and 2000. Differing male and female total length‐(L_T) frequency distributions revealed SSD with females having greater mean and maximum sizes than males. Although both sexes exhibit indeterminate post‐maturity growth, growth trajectories differed. The point of trajectory divergence was not established, but slowed male growth might coincide with the onset of maturation. Differing growth rate was the dominant proximate cause of SSD, accounting for an estimated 61% of the observed difference in mean adult L_T. The degree of SSD in adults, however, was also related to two other factors. Evidence suggests males become sexually active at a smaller size and earlier age than females; a 2 year difference, suggested here, accounted for an estimated 12% of the between‐sex difference in mean adult L_T. Temporal shifts in gender‐specific survival accounted for an additional 27% of the observed between‐sex difference in mean adult L_T. Estimated age distributions indicated a higher number of older females than older males and more younger males than younger females in the population during the period of sampling. Dissimilarity of age distributions was an unexpected result because the male : female population sex ratio was 1 : 1 and estimates of long‐term annual survival for adult males and females were equal (88%). Future assessments of SSD in this population are apt to vary depending on the prior history of short‐term gender‐specific survival. Without recognizing SSD, non‐gender‐specific growth curves overestimate mean age of adult females and underestimate mean age of adult males of given L_T. Assuming age 8 years for first reproduction in males and age 10 years for females, the adult male : female ratio was estimated as 1·1 : 1 and mean adult age, or generation time, was estimated as 16·4 years for males and 18·4 years for females.