Interpopulation variation in growth and life-history traits of the introduced sunfish, pumpkinseed Lepomis gibbosus, in southern England

We examined attributes of growth and reproduction in 19 populations of pumpkinseed (Lepomis gibbosus) introduced into southern England in order to: (i) assess variability of these traits in a northern European climate; (ii) assess inter‐relationships among these variables; and (iii) compare these attributes with populations from other parts of Europe where pumpkinseeds have been introduced. Growth rates varied considerably among populations, but juvenile growth rates and adult body sizes were generally among the lowest in Europe. Mean age at maturity ranged from 2.0 to 3.9, and was strongly predicted by the juvenile growth rate (earlier maturity with faster juvenile growth). Other population parameters that also displayed significant negative associations with mean age at maturity were gonadosomatic index, body condition, and adult body size (total length, TL at age 5). Mean TL at maturity and the adult growth increment showed no significant associations with any of the other growth or life‐history variables. Pumpkinseed populations in England matured significantly later than those introduced into warmer, more southerly areas of the continental Europe. All of these data suggest that a combination of cool summer temperatures and resource limitation is the cause of slow growth, small adult body size and delayed maturity relative to introduced populations on the European mainland.     

Interpopulation variation in Chrysoperla carnea reproduction: implications for mass-rearing and storage

To determine whether diverse populations and biotypes of Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae) can be mass-produced and stored under uniform conditions, we assessed reproduction in five North American populations, four from the carnea biotype and one from the mohave biotype. Non-diapause adults from the carnea biotype varied significantly in only one aspect of reproduction, viz., the preoviposition period, whereas the duration and rate of oviposition, as well as fecundity, did not vary significantly. Relative to the carnea populations, the mohave population typically had delayed oviposition and a lower rate of oviposition; however, its oviposition period was long, and thus its fecundity differed only from that of the most productive carnea populations.The shelf-life (= survival during storage, and subsequent survival and reproduction after storage) was evaluated for four populations of the carnea biotype. Survival of diapausing adults during the first 18 weeks of storage was high among all four populations; adults from three populations also showed high survival after 35 weeks of storage. A high incidence of females from all four populations laid some fertile eggs after being stored for up to 31 weeks or longer. After six or more weeks of storage, average fecundity was high and ranged from 400–900 eggs/female. Duration of storage (up to 39 weeks) was a significant determinant of fecundity in two of the four populations.In practice, diverse populations of C. carnea can be efficiently mass-produced (with or without storage) under uniform conditions in the insectary. Thus, the choice of which populations to mass-produce and use in augmentation can be based primarily on the characteristics (e.g., climate, habitat, crop) of the intended area of release, rather than on considerations concerning mass-rearing.     

Consequences of hierarchical allocation for the evolution of life-history traits

Resource allocation within individuals may often be hierarchical, and this may have important effects on genetic correlations and on trait evolution. For example, organisms may divide energy between reproduction and somatic growth and then subdivide reproductive resources. Genetic variation in allocation to pathways early in such hierarchies (e.g., reproduction) can cause positive genetic correlations between traits that trade off (e.g., offspring size and number) because some individuals invest more resources in reproduction than others. We used quantitative-genetic models to explore the evolutionary implications of allocation hierarchies. Our results showed that when variation in allocation early in the hierarchy exceeds subsequent variation in allocation, genetic covariances and initial responses to selection do not reflect trade-offs occurring at later levels in the hierarchy. This general pattern was evident for many starting allocations and optima and for whether traits contributed multiplicatively or additively to fitness. Finally, artificial selection on a single trait revealed masked trade-offs when variation in early allocation was comparable to subsequent variation in allocation. This result confirms artificial selection as a powerful, but not foolproof, method of detecting trade-offs. Thus, allocation hierarchies can profoundly affect life-history evolution by causing traits to evolve in the opposite direction to that predicted by trade-offs.     

Within-stream variation in early life-history traits in brown trout

Significant additive genetic variance for most early life-history traits was found in brown trout Salmo trutta living in both allopatry above an impassable waterfall and sympatry (below the waterfall in the same stream) with alpine bullhead Cottus poecilopus. These traits included length, mass and yolk sac volume at hatching, and size at'button-up' (the time when yolk is enclosed within the body cavity). There were small differences in size at hatching and size at button-up among populations (adjusted for egg size). However, sympatric fry grew more rapidly and experienced lower mortality rates during the period of first feeding than allopatric fry. This might indicate behavioural differences between brown trout from the two populations. It is suggested that these phenotypic differences may be a result of adaptation to living in sympatry with alpine bullhead.     

Genetic variation for phenotypically invariant traits detected in teosinte: implications for the evolution of novel forms.

How new discrete states of morphological traits evolve is poorly understood. One possibility is that single-gene changes underlie the evolution of new discrete character states and that evolution is dependent on the occurrence of new single-gene mutations. Another possibility is that multiple-gene changes are required to elevate an individual or population above a threshold required to produce the new character state. A prediction of the latter model is that genetic variation for the traits should exist in natural populations in the absence of phenotypic variation. To test this idea, we studied traits that are phenotypically invariant within teosinte and for which teosinte is discretely different from its near relative, maize. By employing a QTL mapping strategy to analyze the progeny of a testcross between an F(1) of two teosintes and a maize inbred line, we identified cryptic genetic variation in teosinte for traits that are invariant in teosinte. We argue that such cryptic genetic variation can contribute to the evolution of novelty when reconfigured to exceed the threshold necessary for phenotypic expression or by acting to modify or stabilize the effects of major mutations.     

Interpopulation variation in reproductive traits of female masu salmon, Oncorhynchus masou

Several life history models predict that larger eggs and lower fecundity should be favored in a low-growth environment. We applied the model of Sibly et al. to seven Japanese populations of masu salmon ( Oncorhynchus masou ) in order to test the predictions that populations in which individual growth rate is low are characterized by larger eggs and lower fecundity. Two populations, Shumarinai and Shikaribetsu, had the lowest growth rates of individuals, largest egg sizes and lowest fecundities of all populations examined. In contrast, the Shiribetsu, Chitose and Uono populations had highest growth rates of individuals, smallest egg sizes and highest fecundities. The Shibetsu and Toya populations were intermediate between these two groups. The gonadsomatic index (GSI) of the Shumarinai and Shikaribetsu populations was smaller than that of all other populations. Correlation analysis indicated that populations with lower individual growth rates had larger eggs and lower fecundities. The results were consistent with the predictions of the Sibly et al. model: increased egg size which results in decreased fecundity is probably an adaptation to low-growth environments. Therefore, in masu salmon, growth differences among populations may explain interpopulation variation in egg size and fecundity.     

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.     

Large-scale intraspecific variation in life-history traits of European freshwater fish

Aims To test the magnitude and direction of the effects of large‐scale environmental factors (latitude and habitat type: lotic or lentic) on the intraspecific variations in multiple life‐history traits, across multiple European freshwater fish species. To test the relevance of defining species traits by quantifying the magnitude of interspecific vs. intraspecific variability in traits. Location Europe. Methods We obtained estimates of 11 fish traits from published sources for 1089 populations of 25 European freshwater fish species. Traits were: longevity, maximal length, growth rate, asymptotic length, mortality rate, age and length at maturation, fecundity, egg size, gonadosomatic index, and length of breeding season. We described population habitats by latitude and habitat type (lotic or lentic), when available. For each species we tested the combined effect of latitude and habitat type on the intraspecific variation of each trait using analysis of covariance (ancova ). We compared the intraspecific variation in traits with the variation between species using an analysis of variance (anova ) for each trait, all species pooled. Results We found a consistent effect in direction of latitude on six traits, but we showed that this effect is not always significant across species. Higher‐latitude populations often grew more slowly, matured later, had a longer life span and a higher maximal and asymptotic length, and allocated more energy to reproduction than populations at lower latitudes. By contrast, we noted only a slight effect of habitat type on the intraspecific variation in traits, except for Salmo trutta. All traits varied significantly between species. However, traits such as growth rate, mortality rate and length of breeding season varied more between populations than between species, whereas fecundity and traits associated with body length varied more between species. Main conclusions Latitude, in contrast to habitat type, is an important factor influencing several traits of geographically widely dispersed populations of multiple European freshwater fish species. Species traits that vary more between species than between populations are attractive variables for understanding and predicting the responses of stream fish communities to their environment.     

The components of life-history traits variation in Daphnia magna population

The variation of six growth and reproduction traits in the parthenogenetic clones of Daphnia magna and its relationships with the genotypes at 5 isozyme loci were studied under controlled conditions at three food concentrations. A significant genetic component of age at maturity, growth rate and fecundity variation was detected. Nested ANOVA revealed the influences of the genotypes of the isozyme loci, that are more pronounced at the high food concentration. In some cases the genotype-environment interactions were found.The variation of body length at 6 days and total number of offspring includes a significant heterozygosity — environment interaction. The most heterozygous clones show the least differences in life history traits between food conditions.Life-history traits heritability, genotype-environment interactions and different reproductive strategies of the clones are discussed.     

Quantitative genetic approach for assessing invasiveness: geographic and genetic variation in life-history traits

Predicting the spread of invasive species is a challenge for modern ecology. Although many invasive species undergo genetic bottlenecks during introduction to new areas resulting in a loss of genetic diversity, successful invaders manage to flourish in novel environments either because of pre-adaptations or because important traits contain adaptive variation enabling rapid adaptation to changing conditions. To predict and understand invasion success, it is crucial to analyse these features. We assessed the potential of a well-known invader, the Colorado potato beetle (Leptinotarsa decemlineata), to expand north of its current range in Europe. A short growing season and harsh overwintering conditions are apparent limiting factors for this species’ range. By rearing full-sib families from four geographically distinct populations (Russia, Estonia, Poland, Italy) at two fluctuating temperature regimes, we investigated (a) possible differences in survival, development time, and body size among populations and (b) the amount of adaptive variation within populations in these traits. All populations were able to complete their development in cooler conditions than in their current range. A significant genotype–environment interaction for development time and body size suggests the presence of adaptive genetic variation, indicating potential to adapt to cooler conditions. The northernmost population had the highest survival rates and fastest development times on both temperature regimes, suggesting pre-adaptation to cooler temperatures. Other populations had minor differences in development times. Interestingly, this species lacks the classical trade-off between body size and development time which could have contributed to its invasion potential. This study demonstrates the importance of considering both ecological and evolutionary aspects when assessing invasion risk.     

Comparisons of life-history traits between clonal and sexual fish (Poeciliopsis:Poeciliidae) raised in monoculture and mixed treatments

Summary Clonal and sexual co-existence is common in a number of vertebrate taxa, even though the cost of sex makes such co-existence theoretically unlikely. The frozen niche-variation (FNV) model explains this co-existence on the basis of differences in overall niche breadth and competition between clones and sexuals. In the present study I examined two predictions of the FNV model. First, I examined the prediction that genetically variable populations have higher relative fitness when compared with monoclonal populations by comparing the performances of clonal and outcrossed sexual strains ofPoeciliopsis in monocultures at two densities. The prediction of increased overall productivity for the sexuals was verified, with net reproductive rates for the sexuals being between two and four times as high as the clones. Second, I tested the prediction that derived clones will successfully compete with their sexual progenitor(s) in the narrow range to which the clones are adapted, while the sexuals should co-exist because of their ability to use a wider range of resources than any single clone. I examined this prediction by comparing performance variables (e.g. growth, fecundity and survival) of each strain in pure culture with their partitioned performance from the mixed treatments. Clonal performance increased in mixtures compared to monocultures, as expected. However, the expectation that the sexual's performance would be less affected by mixtures than the clones' performance, was not met. The sexuals had reduced growth and fecundity on a par with the increase in both variables in the clones. Therefore, support for the FNV model was mixed. Although the performance in monocultures suggests that the sexuals have a wider niche breadth than the clones, performances in mixtures do not indicate such a relationship. Switching of behaviours or resource-use patterns between mixed and pure cultures may have caused the equivocal results.     

Local adaptation in brown trout early life-history traits: implications for climate change adaptability

Knowledge of local adaptation and adaptive potential of natural populations is becoming increasingly relevant due to anthropogenic changes in the environment, such as climate change. The concern is that populations will be negatively affected by increasing temperatures without the capacity to adapt. Temperature-related adaptability in traits related to phenology and early life history are expected to be particularly important in salmonid fishes. We focused on the latter and investigated whether four populations of brown trout (Salmo trutta) are locally adapted in early life-history traits. These populations spawn in rivers that experience different temperature conditions during the time of incubation of eggs and embryos. They were reared in a common-garden experiment at three different temperatures. Quantitative genetic differentiation (QST) exceeded neutral molecular differentiation (FST) for two traits, indicating local adaptation. A temperature effect was observed for three traits. However, this effect varied among populations due to locally adapted reaction norms, corresponding to the temperature regimes experienced by the populations in their native environments. Additive genetic variance and heritable variation in phenotypic plasticity suggest that although increasing temperatures are likely to affect some populations negatively, they may have the potential to adapt to changing temperature regimes.     

Parasites and host life-history traits: implications for community ecology and species co-existence

Most of the evidence for a key role of parasites in structuring communities is based on the idea of a differential susceptibility of host species to infection and its consequences. Recent advances in community ecology suggest that life-history traits of free-living species can be an important determinant of their co-existence within communities. On the other hand, parasites have the potential to indirectly alter the life-history traits of their hosts, such as developmental time or dispersal. We discuss the idea that these indirect effects could influence the structure of free-living and parasite communities. We explore this idea in relation to related concepts including 'parasitic arbitration' and engineering processes.     

Predation effects on the evolution of life-history traits in a clonal oligochaete

Although size at maturity and size and number of offspring are life-history traits widely studied in sexual and parthenogenetic reproduction, there is no such research on animals reproducing asexually without the involvement of gametes. Here we present an individual-based model in combination with experiments to study the clonal growth of Stylaria lacustris, an oligochaete reproducing through fission. We studied the effect of individual size at fission and fission ratio on clone fitness. Our results show that in benign environments without predators, fitness is higher when small worms produce small offspring. Then we included size-specific sublethal predation and found that the fitness of the clone is maximized when parental worms start fission at a large size and produce large descendants intercalated in the middle of the parental worm's body. These results agree with empirical findings. Furthermore, the results of our own laboratory experiment revealed that when S. lacustris is exposed to chemical alarm signals from injured conspecifics, it alters its life history in the same direction as predicted by the model. Our findings suggest that the effect of size-specific sublethal predation is similar to the effect of size-specific lethal predation because both modes of predation result in size-dependent prey mortality.     

Chemical defence in a sawfly: genetic components of variation in relevant life-history traits

Larvae of several tenthredinid sawfly species readily release droplets of haemolymph through their integument when attacked by predators. This defence mechanism via 'bleeding' is characterised by a low integument resistance and a high haemolymph deterrence. Both traits are variable, and negatively correlated among species. We sought to determine if such differences in the propensity to bleed also occur intraspecifically by studying the heritability of traits potentially associated with the bleeding phenomenon in the turnip sawfly Athalia rosae ruficornis Jakovlev (Hymenoptera: Tenthredinidae, Allantinae). For three European populations, heritabilities were estimated in the laboratory in a parent-offspring and a full-sib design for haemolymph deterrence (measured as concentration of sequestered glucosinolate), integument resistance, body mass of eonymph and adult, and developmental time. Within A. rosae, no significant negative phenotypic correlation was found between the two traits directly related to the defence mechanism: integument resistance and haemolymph deterrence. However, the significant heritabilities found for these traits in the full-sib analysis (0.39 and 0.35, respectively, for males in the Swiss population) show that the variation has a genetic component. While full-sib analysis revealed highly significant heritabilities for most traits in all the three populations, parent-offspring regression revealed little or no evidence of heritable variation. Effects of common environment for siblings and variation in the host-plant quality between insect generations are likely to be the main factors explaining these differences. A consequence of such host-plant variation in the wild might be that genetic variation of such chemical defensive traits is largely invisible to natural selection.     

Covariation in life-history traits, demographics and behaviour in ischnuran damselflies: the evolution of monandry

The apparent erratic variation in life history traits, coloration patterns, and behaviours that exists among species within the damselfly genus Ischnura is shown to be interpretable when the species are partitioned into three groups. One group consists of species whose males are missing a pair of stout basal spines on the penultimate segment of their accessory penes. These are the only ischnurans in which males, by tandem guarding females, prevent sperm displacement. The other two groups can be recognized by the relative frequency with which mating occurs: monandrous species mate infrequently, polyandrous species more often. Compared to polyandrous species, monandrous species contain smaller size individuals, have greater sexual size dimorphism, have shorter duration copulations, do not have male biased operational sex ratios at aquatic sites, and are more likely to contain monochromatic females. Females belonging to the monandrous species tend to develop a characteristic form of pruinescence at maturity that obscures their underlying colour, and mature at a younger age. We propose that copulation serves only for sperm addition in monandrous species, for both sperm addition and displacement in polyandrous tandem guarding ischnurans, and for contact guarding as well as sperm addition and displacement in polyandrous species that do not tandem guard.     

The evolution of life-history traits in parasitic and free-living platyhelminthes: a new perspective

Parasite life histories have been assumed to be shaped by their particular mode of existence. To test this hypothesis, we investigate the relationships between life-history traits of free-living and parasitic platyhelminthes. Using phylogenetically independent contrasts we examine patterns of interspecific covariation in adult size, progeny volume, daily fecundity, total reproductive capacity, age at first reproduction and longevity. The correlations obtained indicate a similar causal chain of life history variations for free-living and parasitic platyhelminthes. These results suggest that increased longevity favours delayed reproduction. Furthermore, growth pattern determines adult body size and age at maturity. For platyhelminthes, whether free-living or parasitic, the total reproductive capacity is found to be directly determined by the size of the worm. Within this group the parasitic way of life does not seem to influence the basic patterns of life history evolution. Received: 20 September 1997 / Accepted: 1 March 1998     

Interpopulation variation in reproductive traits of anadromous female brown trout, Salmo trutta L.

We studied reproductive traits in nine anadromous brown trout, Salmo trutta L., populations in seven Norwegian rivers. Within populations we found a positive significant correlation between fish length and fecundity in all populations, and between fish length and egg diameter in five populations. There were significant differences in these relationships between populations from different rivers, and between populations from different locations within rivers. When adjusted for variation in fish length, mean fecundity and mean egg diameter showed a negative significant correlation among populations. The ratio of gonadal weight to somatic weight (gonadosomatic index) varied significantly among populations but was not associated with variation in fish length. Comparatively few large eggs were found in brown trout populations co-existing with several other fish species.     

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.     

Population structure in a common Caribbean coral-reef fish: implications for larval dispersal and early life-history traits

Genetic population structure throughout the Caribbean Basin for one of the most common and widespread reef fish species, the bicolour damselfish Stegastes partitus was examined using microsatellite DNA markers. Spatial autocorrelation analysis showed a significant positive correlation between genetic and geographic distance (isolation by distance) over distances <1000 km, suggesting that populations are connected genetically but probably not demographically, i.e. over shorter time scales. A difference in spatial patterns of populations in the eastern v. the western Caribbean also raises the probability of an important role for meso-scale oceanographic features and landscape complexity within the same species. A comparison of S. partitus population structure and life-history traits with those of two other species of Caribbean reef fish studied earlier showed the findings to be concordant with a common hypothesis that shorter pelagic larval dispersal periods are associated with smaller larval dispersal scales.