Phenotypic Plasticity In Life-history Traits Research Articles

Phenotypic plasticity in reproductive and somatic efforts of female Columbian ground squirrels

Phenotypic plasticity of life-history traits is well known among vertebrate species. We estimated reproductive and somatic efforts of female Columbian ground squirrels ( Urocitellus columbianus (Ord, 1815)) to test for plasticity of these important resource allocations. We examined a 27-year dataset of life-history traits on these long-living (8–10 years), hibernating, montane-living mammals. Environmental variation was estimated from two important traits of mothers, their relative timing of breeding and spring maternal body mass (initial “capital” for use in subsequent reproduction). Results from 183 known-aged mothers and 508 litters revealed considerable variation in the relative timing of breeding, initial maternal mass, and reproductive and somatic efforts, as well as significant variation among ages and years. Results from 125 mothers that reproduced more than once (and 450 litters) revealed significant plasticity of reproductive and somatic efforts with respect to relative timing of breeding and spring maternal mass. A within-subject centering statistical approach showed that phenotypically plastic reproductive and somatic efforts were due to variation within individuals, but were not always reflected by the pattern of responses among individuals in the population. The plastic responses of different mothers appeared to be similar in strength.

Shallow seamounts represent speciation islands for circumglobal yellowtail Seriola lalandi

Phenotypic plasticity in life-history traits in response to heterogeneous environments has been observed in a number of fishes. Conversely, genetic structure has recently been detected in even the most wide ranging pelagic teleost fish and shark species with massive dispersal potential, putting into question previous expectations of panmixia. Shallow oceanic seamounts are known aggregation sites for pelagic species, but their role in genetic structuring of widely distributed species remains poorly understood. The yellowtail kingfish (Seriola lalandi), a commercially valuable, circumglobal, epipelagic fish species occurs in two genetically distinct Southern Hemisphere populations (South Pacific and southern Africa) with low levels of gene-flow between the regions. Two shallow oceanic seamounts exist in the ocean basins around southern Africa; Vema and Walters Shoal in the Atlantic and Indian oceans, respectively. We analysed rare samples from these remote locations and from the South African continental shelf to assess genetic structure and population connectivity in S. lalandi and investigated life-history traits by comparing diet, age, growth and maturation among the three sites. The results suggest that yellowtail from South Africa and the two seamounts are genetically and phenotypically distinct. Rather than mere feeding oases, we postulate that these seamounts represent islands of breeding populations with site-specific adaptations.

Germination time influences post-germination life-history traits and progeny seed germination patterns in the desert annual Erodium laciniatum (Geraniaceae)

Annual plants in arid regions germinate at different times within a growing season, from early in the season to late, and this may affect post-germination traits. For this study, I tested the effect of germination timing on post-germination life-history traits, including progeny seed germination in the desert annual Erodium laciniatum var. pulverulentum (Cav.) Boiss. Traits of November- and February-germinated individuals were studied in a field survey carried out in northwestern Saudi Arabia, and the germination of freshly matured and after-ripened seeds from both early- and late-germinated plants was assessed. Overall, E. laciniatum showed significant phenotypic plasticity in life-history traits arising from different germination times. Density, survivorship and reproductive success of early-germinated plants were all significantly greater than for those that germinated later. Late-germinated plants flowered earlier, bolted at smaller size and allocated more biomass to reproduction than did early-germinated individuals. Delayed germination shortened both flowering period and life span. Seeds produced by late-germinated plants had greater germination percentage than did seeds from early-germinated plants.

Phenotypic plasticity and local adaptation in freeze tolerance: Implications for parasite dynamics in a changing world

Marshallagia marshalli is a multi-host gastrointestinal nematode that infects a variety of artiodactyl species from temperate to Arctic latitudes. Eggs of Marshallagia are passed in host faeces and develop through three larval stages (L1, L2, and L3) in the environment. Although eggs normally hatch as L1s, they can also hatch as L3s. We hypothesised that this phenotypic plasticity in hatching behaviour may improve fitness in subzero and highly variable environments, and this may constitute an evolutionary advantage under current climate change scenarios. To test this, we first determined if the freeze tolerance of different free-living stages varied at different temperatures (−9 °C, −20 °C and −35 °C). We then investigated if there were differences in freeze tolerance of M. marshalli eggs sourced from three discrete, semi-isolated, populations of wild bighorn and thinhorn sheep living in western North America (latitudes: 40°N, 50°N, 64°N). The survival rates of eggs and L3s were significantly higher than L1s at −9 °C and −20 °C, and survival of all three stages decreased significantly with increasing freeze duration and decreasing temperature. The survival of unhatched L1s was significantly higher than the survival of hatched L1s. There was no evidence of local thermal adaptation in freeze tolerance among eggs from different locations. We conclude that developing to the L3 in the egg may result in a fitness advantage for M. marshalli, with the egg protecting the more vulnerable L1 under freezing conditions. This phenotypic plasticity in life-history traits of M. marshalli might be an important capacity, a potential exaptation capable of enhancing parasite fitness under temperature extremes.

Phenotypic plasticity of life-history traits of a calanoid copepod in a tropical lake: Is the magnitude of thermal plasticity related to thermal variability?

According to the Climatic Variability Hypothesis [CVH], thermal plasticity should be wider in organisms from temperate environments, but is unlikely to occur in tropical latitudes where temperature fluctuations are narrow. In copepods, food availability has been suggested as the main driver of phenotypic variability in adult size if the range of temperature change is less than 14°C. Leptodiaptomus garciai is a calanoid copepod inhabiting Lake Alchichica, a monomictic, tropical lake in Mexico that experiences regular, narrow temperature fluctuations but wide changes in phytoplankton availability. We investigated whether the seasonal fluctuations of temperature and food produce phenotypic variation in the life-history traits of this tropical species. We sampled L. garciai throughout a year and measured female size, egg size and number, and hatching success, along with temperature and phytoplankton biomass. The amplitude of the plastic responses was estimated with the Phenotypic Plasticity Index. This index was also computed for a published dataset of 84 copepod populations to look if there is a relationship between the amplitude of the phenotypic plasticity of adult size and seasonal change in temperature. The temperature annual range in Lake Alchichica was 3.2°C, whereas phytoplankton abundance varied 17-fold. A strong pattern of thermal plasticity in egg size and adult female size followed the inverse relationship with temperature commonly observed in temperate environments, although its adaptive value was not demonstrated. Egg number, relative reproductive effort and number of nauplii per female were clearly plastic to food availability, allowing organisms to increase their fitness. When comparing copepod species from different latitudes, we found that the magnitude of thermal plasticity of adult size is not related to the range of temperature variation; furthermore, thermal plasticity exists even in environments of limited temperature variation, where the response is more intense compared to temperate populations.

Phenotypic plasticity in growth and fecundity induced by strong population fluctuations affects reproductive traits of female fish.

Fish are known for their high phenotypic plasticity in life-history traits in relation to environmental variability, and this is particularly pronounced among salmonids in the Northern Hemisphere. Resource limitation leads to trade-offs in phenotypic plasticity between life-history traits related to the reproduction, growth, and survival of individual fish, which have consequences for the age and size distributions of populations, as well as their dynamics and productivity. We studied the effect of plasticity in growth and fecundity of vendace females on their reproductive traits using a series of long-term incubation experiments. The wild parental fish originated from four separate populations with markedly different densities, and hence naturally induced differences in their growth and fecundity. The energy allocation to somatic tissues and eggs prior to spawning served as a proxy for total resource availability to individual females, and its effects on offspring survival and growth were analyzed. Vendace females allocated a rather constant proportion of available energy to eggs (per body mass) despite different growth patterns depending on the total resources in the different lakes; investment into eggs thus dictated the share remaining for growth. The energy allocation to eggs per mass was higher in young than in old spawners and the egg size and the relative fecundity differed between them: Young females produced more and smaller eggs and larvae than old spawners. In contrast to earlier observations of salmonids, a shortage of maternal food resources did not increase offspring size and survival. Vendace females in sparse populations with ample resources and high growth produced larger eggs and larvae. Vendace accommodate strong population fluctuations by their high plasticity in growth and fecundity, which affect their offspring size and consequently their recruitment and productivity, and account for their persistence and resilience in the face of high fishing mortality.

Genetic differentiation and phenotypic plasticity in life-history traits between native and introduced populations of invasive maple trees

Genetically based phenotypic differentiation between native and invasive populations of exotic plants has been increasingly documented and commonly invoked to explain the success of some invasive species. Nonetheless, this basic information is lacking for invasive trees although they currently represent a major concern worldwide. Reciprocal common gardens were therefore set up in both native and introduced ranges of two exotic maple trees to assess the contribution of genetic differentiation and phenotypic plasticity to tree invasiveness. Almost 3,000 native and invasive seedlings of Acer negundo and Acer platanoides were planted in Canada and in France and their performances were compared in various life-history traits related to growth, leaf phenology and ecophysiology over 2 and 3 year periods. Invasive populations of A. negundo exhibited strong genetic differentiation in all the traits examined. Compared to their native conspecifics, they grew significantly larger in the introduced range and showed lower survival, reduced maximum assimilation rate and increased leaf area in the two gardens. They also expressed greater plasticity for growth and greater phenological sensitivity to temperature. Native and invasive populations of A. platanoides were plastic across environments but in contrast did not exhibit any genetic differentiation. This cross-continental comparison provides evidence that both genetic differentiation and phenotypic plasticity contribute synergistically to tree invasiveness. The influence of these respective processes depends on stage of invasion and the life-history strategy of each species. Plastic effects are likely more important during colonization and establishment whilst genetic effects may contribute more significantly during the spread of established populations.

Reproductive parameters of topmouth gudgeon (Pseudorasbora parva) from a heated Lake Licheńskie (Poland)

Abstract Temperature regime is known to have considerable influence on the life-histories of fish. Topmouth gudgeon is a very successful invader with high biological flexibility and phenotypic plasticity. The flexibility of topmouth gudgeon in a normal thermal regime has been well described, however there has been no information published about how it would cope with extreme conditions, e.g. permanently heated water. Several studies have demonstrated that phenotypic plasticity provides an important advantage to invasive species. To explain their invasion success, the theory of alternative ontogenies and invasive potential has been proposed. The aim of the present study was to examine if the reproductive parameters of a population living in permanently heated water differ from those living in habitats with a normal temperature regime, as well as to test a hypothesis derived from the theory of alternative ontogenies and invasive potential. Over a period of 8 years (2004–2011), a high variation in the reproductive parameters of topmouth gudgeon from Lake Licheńskie was found. With a small exception, all the observed results met the expectations predicted by the hypothesis tested. This indicates that topmouth gudgeon retains high phenotypic plasticity in life-history traits even in a habitat with extreme conditions, which is important for the species’ invasion success.

Life-history and thermal tolerance traits display different thermal plasticities and relationships with temperature in the marine polychaete Ophryotrocha labronica La Greca and Bacci (Dorvilleidae)

In order to gain insights into links between fitness and physiological function, the phenotypic plasticity of life-history traits and thermal tolerances has been investigated in a laboratory strain of the polychaete Ophryotrocha labronica acclimated to a broad range of temperatures. Reproductive pairs were kept in isolation for two months at one of six temperatures (10, 15, 20, 25, 30 and 35°C), and survival and a number of life-history traits recorded. Tolerance to heat and cold was assessed in individuals that survived the acclimation period by measuring upper and lower thermal limits. Life-history and thermal tolerance traits showed different relationships with temperature: generally non-monotonic for life-history and monotonic for thermal limits. The thermal history of the strain used in the study seemed to be partly responsible for this pattern, affecting the thermal responses of most life-history traits, but not those of the upper and lower thermal limits. Survival correlated positively with tolerance to cold but not with heat tolerance. This result supports the idea that tolerance to heat is under stronger selection in O. labronica, resulting in a lower level of plasticity for this trait. Some life-history features positively co-varied with tolerance to heat (e.g. body size), whilst in other cases reproductive traits showed apparent trade-offs with thermal limits. In all cases correlations amongst traits were apparently driven by their relationship with temperature. Even if we found no evolutionary trade-offs among the thermal responses of life-history traits and thermal limits, the strategies evolved by organisms to cope with thermal variability result from the mutual interaction of these traits, which plays a key role in defining organisms' responses to temperature changes and fluctuations.

Seasonal Photoperiods Alter Developmental Time and Mass of an Invasive Mosquito,Aedes albopictus(Diptera: Culicidae), Across Its North-South Range in the United States

The Asian tiger mosquito, Aedes albopictus (Skuse), is perhaps the most successful invasive mosquito species in contemporary history. In the United States, Ae. albopictus has spread from its introduction point in southern Texas to as far north as New Jersey (i.e., a span of approximately 14 degrees latitude). This species experiences seasonal constraints in activity because of cold temperatures in winter in the northern United States, but is active year-round in the south. We performed a laboratory experiment to examine how life-history traits of Ae. albopictus from four populations (New Jersey [39.4 degrees N], Virginia [38.6 degrees N], North Carolina [35.8 degrees N], Florida [27.6 degrees N]) responded to photoperiod conditions that mimic approaching winter in the north (short static daylength, short diminishing daylength) or relatively benign summer conditions in the south (long daylength), at low and high larval densities. Individuals from northern locations were predicted to exhibit reduced development times and to emerge smaller as adults under short daylength, but be larger and take longer to develop under long daylength. Life-history traits of southern populations were predicted to show less plasticity in response to daylength because of low probability of seasonal mortality in those areas. Males and females responded strongly to photoperiod regardless of geographic location, being generally larger but taking longer to develop under the long daylength compared with short day lengths; adults of both sexes were smaller when reared at low larval densities. Adults also differed in mass and development time among locations, although this effect was independent of density and photoperiod in females but interacted with density in males. Differences between male and female mass and development times was greater in the long photoperiod suggesting differences between the sexes in their reaction to different photoperiods. This work suggests that Ae. albopictus exhibits sex-specific phenotypic plasticity in life-history traits matching variation in important environmental variables.

Effects of maternal and offspring environmental conditions on growth, development and diapause in latitudinal yellow dung fly populations

Extensive phenotypic plasticity can allow populations to persist in changing environ- ments. Maternal effects represent one important but often neglected source of phenotypic plasticity. Mothers and offspring of 2 high- (northern Norway and central Sweden) and 2 low- (northern and southern Spain) latitude yellow dung fly Scathophaga stercoraria (Diptera: Scathophagidae) popula- tions were exposed to cold (12°C) and warm (18°C) temperatures and to short (8 h light:16 h dark) and long (16 h light:8 h dark) photoperiods in a fully blocked, split-brood common garden design of 8 treatment combinations. We also considered the role of energy content and size of the eggs in pro- ducing cross-generational maternal effects on offspring diapause, development time and growth rate. The incidence of diapause strongly declined towards the south, and the northernmost popula- tion grew and developed faster in response to perceived seasonal time constraints. There was strong population-specific phenotypic plasticity of all traits in response to offspring temperature and, more weakly, to offspring photoperiod, indicating a genetic basis of plasticity as well as genetic differenti- ation among populations. There were additional subtle cross-generational maternal effects exerted primarily by the lipid content of the eggs, largely independent of maternal treatment and population. Phenotypic plasticity of life-history traits in the yellow dung fly is predominantly influenced by the growing conditions during larval development, but populations can also respond to changing envi- ronments via trans-generational maternal effects.

Phenotypic plasticity in fish life-history traits in two neotropical reservoirs: Petit-Saut Reservoir in French Guiana and Brokopondo Reservoir in Suriname

Fish species are known for their large phenotypic plasticity in life-history traits in relation to environmental characteristics. Plasticity allows species to increase their fitness in a given environment. Here we examined the life-history response of fish species after an abrupt change in their environment caused by the damming of rivers. Two reservoirs of different age, both situated on the Guiana Shield, were investigated: the young Petit-Saut Reservoir in French Guiana (14 years) and the much older Brokopondo Reservoir in Suriname (44 years). Six life-history traits in 14 fish species were studied and compared to their value in the Sinnamary River prior to the completion of Petit-Saut Reservoir. The traits analyzed were maximum length, absolute and relative length at first maturation, proportion of mature oocytes in ripe gonad, batch fecundity and mean size of mature oocytes. The results revealed a general increase of reproductive effort. All species showed a decrease in maximum length. Compared to the values observed before the dam constructions, eight species had larger oocytes and three species showed an increased batch fecundity. These observed changes suggest a trend towards a pioneer strategy. The changes observed in Petit-Saut Reservoir also seemed to apply to the 30 years older Brokopondo Reservoir suggesting that these reservoirs remain in a state of immaturity for a long time.

DIVERGENT SELECTION, NOT LIFE-HISTORY PLASTICITY VIA FOOD LIMITATION, DRIVES MORPHOLOGICAL DIVERGENCE BETWEEN PREDATOR REGIMES INGAMBUSIA HUBBSI

We recently tested the hypothesis of ecological speciation in a post-Pleistocene radiation of Bahamas mosquitofish (Gambusia hubbsi) inhabiting inland blue holes (vertical, water-filled caves) on Andros Island, the Bahamas (Langerhans et al. 2007). Combining morphological, molecular, and behavioral data, our results suggested that enhanced premating isolation evolved as a byproduct of differences in body shape between predator regimes, due to assortative mating for body shape. Our results strongly supported the hypothesis of ecological speciation, and parallel results were observed across species within the genus, suggesting a past history of such ecological speciation within the Gambusia lineage. Downhower et al. (in press) question the morphological results described for G. hubbsi in our paper, contending that prior work on life-history variation in G. hubbsi (primarily Downhower et al. 2000, 2002) invalidates our conclusions. Here we show that this contention is based on a mischaracterization of our study, inaccurate quotation of our paper, and misinterpretation of the relevance of their life-history data for morphological results observed in our study. Downhower et al. (in press) suggest—without providing any direct evidence—that differences in body shape between predator regimes in G. hubbsi reflect correlative effects of phenotypic plasticity in life-history traits in response to variation in food availability. To make such a conclusion, we would have to set aside the following facts: (1) morphological differences were observed in adult males, who do not exhibit the life-history traits in question (e.g., number and size of offspring), (2) results matched well-supported a priori predictions of body shape variation based on divergent selection on locomotor performance, (3) results matched empirical results observed in other fish presumably experiencing similar forms of divergent selection between predator regimes, (4) the observed morphological differences are known to influence swimming performance in Gambusia fish and consequently influence endurance and survival with predators, (5) consistent results were observed for males both within and between Gambusia species, (6) any differences in food availability among blue holes are unknown, and (7) available evidence to date indicates that differences in body shape partly reflect genetic divergence, and not solely phenotypic plasticity. The criticisms in Downhower et al. (in press) can be summarized as three arguments: (1) incorrect attribution of evidence for divergent selection between predator regimes in G. hubbsi to several previous

Reaction norms of size and age at maturity of Pomacea canaliculata (Gastropoda: Ampullariidae) under a gradient of food deprivation

Pomacea canaliculata, an apple snail native to South America, has become a serious pest of aquatic crops and a promoter of ecosystem changes in natural wetlands worldwide. Its success as an invader has been attributed to its great phenotypic plasticity in life-history traits. Our aims were to determine the reaction norms of size and age at maturity under a gradient of food deprivation. Full sibling experimental snails were reared in isolation from hatching and maintained until maturity under seven different levels of relative food deprivation based on size-specific ingestion rates. To detect the onset of sexual activity of experimental snails, fully mature virgin snails reared in the laboratory were used as consorts. The reaction norms for age and size at maturity of P. canaliculata showed marked sexual dimorphism. Shell length was the main component of variation in the male reaction norms for both copulation and egg-laying by female consorts, whereas age was the main component of variation for females. Irrespective of the intensity of food deprivation, males mature at the same age at the expense of size, since size is apparently irrelevant in the access to females and male fitness can be maximized through fast maturation. In contrast, a minimum size is required for females to reach maturity, perhaps as a result of their higher reproductive costs. The highly dimorphic reaction norms lead to an increasing lag between male and female maturity as deprivation increases; in temperate regions, males born early in the reproductive season would mature in the same season irrespective of food availability, while most females would have to overwinter before attaining sexual maturity in unproductive habitats or those dominated by unpalatable macrophytes. The great life-history plasticity reported in invaded areas could be a heritage from populations in the native range.

Phenotypic Plasticity in Life-history Traits and Feeding Appendages in Two Species of Daphnia Fed a Natural Phytoplankton Assemblage

The ability of two cladoceran species – Daphnia pulicaria and D. mendotae – to utilize a natural phytoplankton assemblage dominated by cyanobacteria was investigated experimentally. Reference animals were fed high quality green algae. The natural phytoplankton assemblage originated from Oneida Lake (NY, USA). Both cladoceran species occur in the lake, but their densities vary temporally. As expected, lake phytoplankton were a poor food source. Fecundity decreased, and age at first reproduction and offspring size increased. The magnitude of these effects varied between and within species. The increase in offspring size differs from studies using toxic cyanobacterial strains, but agrees with expectations on adaptive responses to low food availability. Both species performed equally on the lake phytoplankton, but D. pulicaria appeared to be the superior species in the high quality food treatment.

Plasticity in age and size at metamorphosis of Crinia georgiana tadpoles: responses to variation in food levels and deteriorating conditions during development

Adaptive phenotypic plasticity in life-history traits is predicted to evolve when populations occur in heterogeneous environments. Anuran larvae of many species cannot escape their aquatic environment until metamorphosis and therefore should show plasticity in response to conditions experienced as tadpoles. In this study, we manipulated the aquatic environments of Crinia georgiana tadpoles in the laboratory to mimic variation among ponds in resources and drying conditions in nature. This species breeds in very shallow water in winter and ponds frequently dry between bouts of rain, especially towards spring. Tadpoles kept in constant conditions at different levels of food metamorphosed at different body sizes but showed no plasticity in metamorphic timing. Tadpoles fed only lettuce metamorphosed at sizes similar to those of field-collected tadpoles, whereas tadpoles fed a more protein-rich food metamorphosed at unusually large sizes, indicating that the seeps where C. georgiana tadpoles occur are poor in nutrients. When we decreased food and water levels, tadpoles at later developmental stages were able to accelerate development and metamorphose earlier than tadpoles kept under constant conditions. Furthermore, tadpoles in very shallow water with no access to food metamorphosed earlier and at smaller body sizes than tadpoles with a more moderate decrease in depth that were able to continue feeding. Rapid development and the ability to accelerate metamorphosis in C. georgiana tadpoles are consistent with adaptation in a heterogeneous environment where larvae are under strong time constraints.

Survival of male Tengmalm's owls under temporally varying food conditions.

We analysed whether annual survival of male Tengmalm's owls in western Finland varies according to changes in the abundance of their main prey, Microtus voles. Our analyses were based on capture-recapture data over five 3-year population cycles of voles from 1981to 1995, each cycle consisting of consecutive years of low, increasing and decreasing vole abundance. Survival estimates of males in the increasing vole years (ca. 55%) were higher than in the decreasing vole years. In the latter case, males faced a drastic crash in Microtus vole numbers, and only about one-third of males survived over this crash. After the crash of vole populations, male survival increased rapidly (up to 76%) according to the recovery of Microtus vole populations. These results show that temporal variation in the abundance of their main prey modifies the survival of male Tengmalm's owls. In addition to survival, recapture rates also varied, largely due to the fact that in poor vole years a majority of males skipped breeding. The large among-cycle phase variation in survival (25-76%) probably creates selection for phenotypic plasticity in life-history traits related to survival and reproduction.

GENETIC VARIATION FOR PHENOTYPIC PLASTICITY IN THE LARVAL LIFE HISTORY OF SPADEFOOT TOADS (SCAPHIOPUS COUCHII).

Phenotypic plasticity in life-history traits is common. The relationship between phenotype and environment, or reaction norm, associated with life-history plasticity can evolve by natural selection if there is genetic variation within a population for the reaction norm and if the traits involved affect fitness. As with other traits, selection on plasticity in a particular trait or in response to a particular environmental factor may be constrained by trade-offs with other traits that affect fitness. In this paper, I experimentally evaluated broad-sense genetic variation in the reaction norms of age and size at metamorphosis in response to two environmental factors, food level and temperature. Differences among full-sib families in one or both traits were evident in all treatments. However, variation among families in their responses to each treatment (genotype-environment interaction) resulted in variation among treatments in estimated heritabilities and genetic correlations. Age at metamorphosis was equally sensitive to temperature in all families, but size at metamorphosis was more sensitive to temperature in some families than in others. Size at metamorphosis was equally sensitive to food level in all families, but age at metamorphosis was sensitive to food in some families but not in others. At high temperature or low food, the genetic correlation between age and size at metamorphosis was positive, generating a potential trade-off between metamorphosing early to attain higher larval survival and metamorphosing later to achieve larger size. This trade-off extends across treatments: families with the largest average size at metamorphosis achieved larger size with the longest average and greatest plasticity in age at metamorphosis. Other families achieved shorter average larval periods by exhibiting greater plasticity in size at metamorphosis but had the smallest average size at metamorphosis. This trade-off may reflect an underlying functional constraint on the ability to respond optimally to all environments, resulting in persistent genetic variation in reaction norms.

Phenotypic plasticity of life-history traits in clonal and sexual fish (Poeciliopsis) at high and low densities.

Models of resource allocation strategies predict an array of life-history responses of individuals living in resource-stressed versus non-stressed environments. I tested a number of these predictions using three fish strains (a sexual and two clonal strains) in high and low density treatments. To examine the plasticity of life-history traits in females raised in these two environments, I measured survival, growth, egg production, egg size, and proportion mature at 10 weeks of age. Survival was not affected by density treatment. However, both growth and overall egg production were lower in females from the high density treatments, and reproductive maturity was significantly delayed at the high density for all strains. Egg production per unit size was not affected by density in any strain, signifying that differences in the numbers of eggs produced was merely a reflection of the differences in size of fish in the two density treatments. Egg size was also unaffected by density in all strains. These results are related to models of resource allocation in stressful environments. There was a consistent pattern of increased reproductive investment in the sexual strain relative to the two clonal strains. The sexual strain matured earlier, produced more eggs per unit body weight, and had larger eggs than either clone at both densities. These results are interpreted by considering the predicted adaptive responses of these three strains to the long-term environmental differences in their natural habitats.

Phenotypic plasticity in life-history traits of femaleThalassoma bifasciatum (Pisces: Labridae): 2. Correlation of fecundity and growth rate in comparative studies

The cost of reproduction is a central concept in theories of life-history evolution. One way to empirically examine the tradeoff between current reproduction and future reproductive prospects is to use natural intraspecific variation in life-history traits. However, this approach is complicated by the sensitivity of life-history traits to variation in the level of resources. We report here an attempt to measure the cost of increasing reproductive activity in populations of female bluehead wrasse,Thalassoma bifasciatum, a coral-reef fish. All of the significant correlations of fecundity and growth rate were positive, in contradiction to the tradeoff predicted by the cost concept. In one of two regions studied, the populations with relatively high mean growth rate had a relatively large mean fecundity. The trait means were also positively associated over time: in months of rapid growth, female reproductive activity was high. Even after removing the effects of habitat and time period in a comparison of individual traits, no growth cost to reproduction appears. Variation in the abundance of resources over space and time is likely to interfere with the measurement of the cost of reproduction in many natural systems.