Offspring number in a livebearing fish (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, Poeciliidae): reduced fecundity and reduced plasticity in a population of cave mollies

Life history traits within species often vary among different habitats. We measured female fecundity in mollies (Poecilia mexicana) from a H₂S-rich cave and from a neighbouring surface habitat, as well as in laboratory-reared individuals of both populations raised in either light or continuous darkness. Compared to conspecifics from surface habitats, cave-dwelling P. mexicana had reduced fecundity (adjusted for size) in the field. In the laboratory, the fecundity of surface mollies was higher in light than in darkness, whereas fecundity in the cave mollies was almost unaffected by the ambient light conditions. Our results suggest a heritable component to the reduction in fecundity in female cave mollies. Moreover, the reduced plasticity in fecundity of cave mollies in response to light conditions might be an example of genetic assimilation or channelling of a life history trait in a population invading a new environment.     

Effects of temperature and salinity on the life history of the sailfin molly (Pisces: Poeciliidae): lipid storage and reproductive allocation

While the life history traits of animals usually exhibit substantial phenotypic plasticity, such plasticity might reflect either a simple alteration in the level of energy accrual and use or a genuine shift in energy allocation tactics between environmental conditions. The latter would represent genuine plasticity in the life history itself, and thus it is important to distinguish which of these two processes underlies the observed plasticity of life history traits. We investigated this issue by examining the effects of temperature and salinity variation during ontogeny on the allocation of biomass and lipid storage in male sailfin mollies, Poecilia latipinna. We raised males from four natural populations from birth to maturity in controlled laboratory conditions. Neither distinct temperatures (23 or 29°C) nor different salinity regimes (2, 12, or 20 parts per thousand) affected body mass, although males from different populations differed substantially in body mass. However, males raised at the higher temperature had a greater allocation of biomass to testis and a lower allocation to viscera mass. The amount of stored lipid was altered by temperature variation but the direction and magnitude of the effect varied substantially among males from the different populations. Salinity variation affected neither biomass allocation nor the level of lipid storage. These results indicate that male mollies possess a flexible developmental program with respect to temperature that canalizes body size and alters the allocation of biomass among competing demands for reproductive readiness and capacity for energy storage. Received: 25 November 1996 / Accepted: 1 December 1997     

An experimental study of the effects of weed invasion on lizard phenotypes

We examined how a weed affected the basking and activity of a diurnal lizard, and the potential cascading effects of these shifts for life history strategies and expression of morphology. Hatchlings of the diurnal lizard Lampropholis delicata were raised to maturity in outdoor enclosures that mimicked high, moderate and low invasion by a sprawling plant (blue periwinkle, Vinca major). Skinks depend on sunlight for growth and maintenance. Periwinkle differs from displaced grassland by being structurally complex and blocking sunlight. Lizards restricted to the enclosure floor achieved preferred body temperatures only when exposure to periwinkle was moderate or low. However, lizards in high invasion enclosures could reach preferred body temperatures by climbing plants and basking on exposed canopy. This shift in basking strategy resulted in lizards growing longer hind limbs compared with animals that rarely (moderate invasion) and never (low invasion) climbed plants. Consequently, lizards reared in high invasion enclosures sprinted faster than conspecifics reared in lower invasion environments. Throughout the study there was no significant variation among treatments in the tendency of animals to be moving when they were not hidden. However, lizards in high invasion treatments hid more often during the day, were lighter in body mass, and females had lighter clutch masses and offspring than did those from moderate and low invasion enclosures. Thus, microhabitat degradation can drive a cascade of changes to an animal’s ecology.     

The effects of water availability on the life history of the desert snail,Trochoidea seetzeni

Summary In the Negev desert of Israel, the pulmonate land snailTrochoidea seetzeni is active, grows and reproduces in the month following the torrential winter rains. Thereafter, these snails estivate until the following year's rains. By experimental supplementation of water in the field, we examined the ability of these snails to alter their life histories. Specifically, we measured changes in feeding activity, overall activity levels, movement patterns and reproductive output. We found that snails on artificially-watered plots stopped feeding on higher plants and fed on soil-surface algae growing as a result of increased water availability. Most snails moved onto the watered plots, and reduced the total amount of movement once there, in response to increased food availability. Snails on the watered plots grew more and remained active for a longer period into the summer months than did snails on control plots. However, we found little evidence for the ability of these snails to alter their reproductive outputs in response to increased water (and hence food) availability. There was also no carry-over effect of high water availability in one season enhancing reproduction in the following season. We postulate that the low level of phenotypic plasticity recorded here is a consequence of an adaptive conservative strategy to maintain a high reproductive output under extremely unpredictable desert conditions.     

Plasticity of grasshopper vitellogenin production in response to diet is primarily a result of changes in fat body mass

Life history plasticity is the developmental production of different phenotypes by similar genotypes in response to different environments. Plasticity is common in early post-embryonic or adult development. Later in the developmental stage, the transition from developmentally plastic to canalized (i.e., inflexible) phases is often associated with the attainment of a threshold level of storage. Thresholds are often described simply as total body mass or cumulative consumption of food. The physiological characteristics of thresholds, such as the contributions of the growth of particular organs or the production rate of proteins, are largely unstudied. To address the physiology underlying a threshold-induced developmental transition, total vitellogenin production in response to diet quality in the lubber grasshopper was studied. For individuals that differed in age or dietary protein, somatic mass, ovarian mass, fat body mass, mass-specific vitellogenin production, vitellogenin titer, and storage protein titer were measured. Age and diet strongly affected these parameters, with ovarian mass and fat body mass contributing most to the differences. During mid vitellogenesis, females were highly plastic in response to changing food quality. Only during late vitellogenesis were females unresponsive to changes in food quality. Fat body mass was a more important component of plasticity than was mass-specific vitellogenin production. Because these two variables together make up total vitellogenin production, the greater contribution of fat body mass than mass-specific vitellogenin production suggests that growth factors may be more important than tissue stimulators in producing developmental changes in total vitellogenin production. To our knowledge, this is the first study to demonstrate that mass gain of an organ is more important to developmental plasticity than is the output of that same organ.     

Maternal thermoregulation influences offspring viability in a viviparous lizard

Gravid females of the viviparous scincid lizard Eulamprus heatwolei were maintained in the laboratory, with some females allowed to bask for 8 h/day and others for only 2 h/day. Maternal basking regimes influenced the gestation period, and significantly affected the body shapes, activity levels and running speeds of the offspring born to these females. Neonates from females with lesser basking opportunities were relatively short and fat, were very active, and were relatively fast runners. Effects of the embryos' thermal regime on the young lizards' morphology and running speed were still detectable two months after birth. Thus, much of the morphological and behavioral variation among neonatal reptiles may arise from phenotypically plastic responses to the thermal environments experienced during embryonic development, rather than from heritable genetic differences among individuals. Hence, selection on maternal thermoregulatory behavior may be an important avenue for adaptive modifications to neonatal phenotypes in reptiles.     

Predator-induced delayed maturity in bluegill sunfish (Lepomis macrochirus): variation among populations

Previous studies suggested that differences in age at maturity among populations of bluegill sunfish (Lepomis macrochirus) were not genetically based, but rather were a phenotypic response to the presence of predators. I conducted two experiments to determine if the presence of largemouth bass affected age at maturity in bluegill sunfish. Bluegills from three populations were tested to see if the response to the threat of predation varied among source populations. Juvenile bluegills were maintained in the presence of predators or in controls with no contact with predators. Refuge use and growth were monitored during the experiments and reproductive activity was evaluated when bluegills reached age 1. Bluegills from one population exhibited delayed maturity in the presence of predators. Individuals from the other two populations showed no significant differences between predator and control treatments. The population that responded to the presence of predators had a history of high predation levels over the past 30–40 years. The other populations had a history of low levels of predation. This study suggests that presence of predators can induce phenotypic shifts in age at maturity of bluegills, but that the magnitude of response varies among populations in a manner consistent with historical patterns of coexistence. Received: 7 August 1996 / Accepted: 8 August 1997     

Gestation length variation in domesticated horses and its relation to breed and body size diversity

The domestication process and selective breeding reportedly alter some life history variables. In horses, it has been claimed that gestation length is particularly variable. Some of the factors influencing gestation length are already known and can be grouped into environmental and genetic factors, but the effects of breed and body size have rarely been evaluated. In this study we tested the influence of breed and body size on gestation length for 25 horse breeds from Central Europe. The mean gestation length for all breeds was 342.3 ± 10.2 days and we found significant differences among breeds with a variation of up to 11 days. Body size did not show a significant correlation with gestation length. Our data suggest that breed affiliation explains part of the large variability of gestation length in horses.     

To hatch and hatch not: similar selective trade-offs but different responses to egg predators in two closely related, syntopic treefrogs

Risk-sensitive hatching is adaptive for species facing a trade-off between egg-stage and post-hatching risks, and environmental variation in one or both stages. Such plasticity has been found in amphibians, fishes, reptiles and spiders, with red-eyed treefrogs (Agalychnis callidryas) being the best-studied case. We assessed hatching plasticity and egg- and larval-stage risks in a closely related, syntopic species, the gliding leaf-frog (Agalychnis spurrelli). We found a lower hatching response to egg-eating snakes in A. spurrelli (9–28% of embryos escaped) than in A. callidryas (59–80% escaped). Levels of snake predation were similarly high for clutches of both species monitored at a pond in Costa Rica, and in fish predation experiments early-hatched A. spurrelli tadpoles were more vulnerable than later hatchlings, as has been shown for A. callidryas. A. spurrelli thus face a risk trade-off similar to A. callidryas, and likely would benefit from predator-induced hatching; their lower responsiveness to snakes appears nonadaptive. A. spurrelli embryos showed a stronger hatching response (57% hatched in 1 h) to submergence underwater than to snake attacks even though submergence is a less frequent risk. This suggests they have a greater capacity for early hatching than is expressed in the context of snake attacks, but have much lower sensitivity to snake cues than to flooding cues. Development in A. spurrelli is accelerated compared to syntopic A. callidryas, and spontaneous hatching is earlier and more synchronous. This is congruent with predictions based on selection by egg predators in the absence of a strong escape hatching response.     

Age specific fecundity and size of offspring in the prosobranch snail,Viviparus ater

Summary Age specific fecundity and size of offspring were compared in two populations of Viviparus ater, one from Lake Zürich (Switzerland) and one from Lake Maggiore (Italy). Females were caged and their offspring sampled at weekly intervals. Average fecundity per female over one year was 19.6 and 14.7 offspring in Lake Zürich and Lake Maggiore, respectively. Average shell widths of offspring at birth in the two lakes were 8.5 mm and 7.4 mm. Fecundity did not change with age in Lake Zürich but decreased with increasing age of the snails in Lake Maggiore. Fecundity and offspring size were positively correlated with the size of the females in both lakes, but fecundity was higher and offspring were larger in Lake Zürich than in Lake Maggiore irrespective of the size of the females. An analysis of covariance, correcting for differences between the two lakes in the total fresh weight of the offspring produced annually, showed that females from the two lakes distributed their reproductive investment differently. Females in Lake Zürich produced relatively fewer but larger offspring than females in Lake Maggiore. This trade-off was hidden because females in Lake Zürich produced twice as much offspring biomass as females in Lake Maggiore. The possibility that different selection regimes might account for the observed differences is discussed.     

Factors promoting maternal trophic egg provisioning in non-eusocial animals

The adaptive function of trophic egg-laying is generally regarded as extended parental investment to the offspring. However, the evolutionary factors promoting trophic egg-laying are still unclear, because the amount of maternal investment per offspring should be ideally equal between smaller offspring with trophic eggs and larger offspring without any additional investment. Several authors have suggested that trophic egg-laying should evolve only when egg size is constrained, but this hypothesis has not been evaluated. We investigated the evolutionary mechanisms of trophic egg-laying by two different approaches. First, we evaluated morphological constraints on egg size in two sibling ladybird species, Harmonia axyridis, which is known to produce trophic eggs, and H. yedoensis. Second, we theoretically predicted the optimal proportion of trophic eggs to total eggs and egg size in relation to environmental heterogeneity, predictability of environmental quality, and investment efficiency of trophic egg consumption. The intra- and interspecific morphological comparisons suggest that morphological constraints on the evolutionary determination of egg size are weak at best in the two ladybird species. Moreover, we theoretically showed that small egg size and trophic egg-laying are favoured in heterogeneous environments when mothers cannot adjust egg size plastically. We also showed that even a small reduction in investment efficiency makes a trophic egg strategy unlikely, despite relatively high environmental predictability. We conclude that trophic egg provisioning may be a flexible maternal adaptation to a highly heterogeneous environment rather than a response to a morphological constraint on egg size.     

Feeding while evading predators by a lotic mayfly: linking short-term foraging behaviours to long-term fitness consequences

Laboratory experiments were performed to quantify the combined effects of food abundance (low, high) and predation threat, imposed using a model fish (safe, risky) on fitness correlates (i.e. growth, time to emergence, adult body mass, fecundity, egg size) of the mayfly Baetis tricaudatus. These effects were determined by rearing larvae under different combinations of food abundance and predator threat. Fitness correlates were significantly affected by food abundance, predation threat or the interaction of these factors. High food abundance and the absence of predation threat significantly increased larval growth rates, adult body mass, fecundity, egg size, and decreased time to emergence. Long-term effects of predator threat and food abundance on fitness correlates of B. tricaudatus were compared to previously conducted short-term patch choice trials to test for concordance between short-term measures of patch choice and their potential longer-term fitness consequences. This comparison indicated that patches that were utilized the most were those that yielded the highest fitness benefits. These results suggest that behavioural costs of balancing mortality risk due to predation against food acquisition by B. tricaudatus can have strong fitness consequences.     

Thermal dependence of reproductive allocation in a tropical lizard

In ectotherms, environmental temperature is the most prominent abiotic factor that modulates life-history traits. We explored the influence of environmental temperature on reproduction in the Madagascar ground gecko (Paroedura picta) by measuring reproductive traits of females at constant temperatures (24, 27, 30 °C). Females of this species lay clutches of one or two eggs within short intervals. For each female, we measured egg mass for the first five clutches. For one clutch, we also measured the energetic content of eggs via bomb calorimetry. Temperature positively influenced the rate of egg production, but females at 30 °C laid smaller eggs than did females at either 24 or 27 °C. Dry mass of eggs scaled allometrically with wet mass, but this relationship was similar among thermal treatments. Females at all temperatures produced eggs with similar energy densities. Females at 24 °C allocated less energy per time unit (≈8 mW) to reproduction than did females from higher temperatures (≈12 mW). However, females at either 24 or 27 °C allocated significantly more energy per egg than did females at 30 °C. Our results demonstrate that a complex thermal sensitivity of reproductive rate can emerge from distinct thermal sensitivities of egg size, egg composition and clutch frequency.     

Revisiting the homoiology hypothesis: the impact of phenotypic plasticity on the reconstruction of human population history from craniometric data

Homoiologies are homoplasies that are caused by nongenetic environmental factors. The homoiology hypothesis predicts that osseous regions subject to repeated biomechanical stress during growth should be more variable and, therefore, less reliable for the reconstruction of phylogeny compared with osseous regions relatively unaffected by stress. Previous studies based on the analysis of multiple primate species found that regions of the cranium subject to masticatory-induced stress were significantly more variable than non-masticatory regions, as predicted by the homoiology hypothesis. However, these studies also found that the masticatory regions were no less reliable for reconstructing primate phylogenetic relationships when subjected to parsimony analysis. It was suggested, therefore, that homoiology may be a more potent problem for the reconstruction of phylogeny at the intraspecific level rather than interspecific phylogenetics. This suggestion was tested here using matched molecular and craniometric data for 12 modern human populations. The results show that, as predicted by the homoiology hypothesis, regions of the human cranium related to mastication were more variable than non-masticatory regions. However, masticatory regions were no less reliable for inferring human population history. Therefore, the results match those found from the interspecific analysis of primate species and do not support the suggestion that homoiology is a greater problem for the analysis of intraspecific taxa. The results also suggest that within-taxon variability cannot be relied upon to predict the phylogenetic efficacy of morphometric characters.     

Phenotypic plasticity and nutrition in a phytophagous insect: consequences of colonizing a new host

The European rose-hip fruit fly Rhagoletis alternata (Diptera, Tephritidae) infests hips of Rosa species. This fly includes R. rugosa, an Asian species now cultivated all over Europe, in its host range. Differences in size and biomass of hips between the ancestral host R. canina and the new host translate into better growth, shorter larval development of larvae within hips of R. rugosa and larger body size and fertility of flies which developing in the new host. In turn this causes different interactions with other organisms of the food-web centred on the host plant. The importance of nutrition and phenotypic plasticity is twofold: they generate a considerable part of life-history diversity within a species and reinforce differences in the ecological context of the ancestral and new host.     

Offspring fitness in relation to population size and genetic variation in the rare perennial plant species Gentiana pneumonanthe (Gentianaceae)

Seeds were sampled from 19 populations of the rare Gentiana pneumonanthe, ranging in size from 5 to more than 50,000 flowering plants. An analysis was made of variation in a number of life-history characters in relation to population size and offspring heterozygosity (based on seven polymorphic isozyme loci). Life-his-tory characters included seed weight, germination rate, proportion of seeds germinating, seedling mortality, seedling weight, adult weight, flower production per plant and proportion of plants flowering per family. Principal component analysis (PCA) reduced the dataset to three main fitness components. The first component was highly correlated with adult weight and flowering performance, the second with germination performance and the third component with seed and seedling weight and seedling mortality. The latter two components were considered as being maternally influenced, since these comprised life-history traits that were significantly correlated with seed weight. Multiple regression analysis showed that variation in the first fitness component was mainly associated with heterozygosity and not with population size, while the third fitness component was only correlated with population size and not with heterozygosity. The latter relationship appeared to be non-linear, which suggests a stronger loss of fitness in the smallest populations. The second (germination) component was neither correlated with population size nor with genetic variation. There was only a weak association between population size, heterozygosity and the population coefficients of variation for each life history character. Most correlation coefficients were negative, however, which suggests that there is more variation among progeny from smaller populations. We conclude that progeny from small populations of Gentiana pneumonanthe show reduced fitness and may be phenotypically more variable. One of the possible causes of the loss of fitness is a combination of unfavourable environmental circumstances for maternal plants in small populations and increased inbreeding. The higher phenotypic variation in small populations may also be a result of inbreeding, which can lead to deviation of individuals from the average phenotype through a loss of developmental stability.     

Divergent phenotypic responses to predators and cyanobacteria in Daphnia lumholtzi

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Evolution of feeding structure plasticity in marine invertebrate larvae: a possible trade-off between arm length and stomach size

The ability of an organism to alter its morphology in response to environmental conditions (phenotypic plasticity) occurs in several species of marine invertebrates. Examples are sea urchin and sand dollar larvae (plutei). When food is scarce, plutei produce longer food-gathering structures (larval arms and a ciliary band) and smaller stomachs than when food is abundant. However, it is unclear whether stomach size is actually induced through changes in morphogenesis or simply by food distending the stomach. Distinguishing between these two hypotheses is possible because plutei morphologically respond to food concentrations and change the length of their food-gathering structures before they are capable of feeding. More importantly, these two hypotheses provide insights to whether a trade-off exists between the response in food-gathering structures and the response in stomach size—a possible explanation for the evolution of feeding-structure plasticity in marine invertebrate larvae. In this study, I investigated whether sea urchin larvae (Strongylocentrotus purpuratus and S. franciscanus) reared in different amounts of food produced stomachs of different sizes before they were capable of feeding. Prior to having the ability to ingest food, larvae produced larger stomachs and shorter arms when food was abundant than when food was scarce, consistent with the hypothesis that food induced changes in morphogenesis. In addition, there was a strong negative correlation between the magnitude of plasticity in larval arm length and the magnitude of plasticity in stomach size. These results are consistent with the idea that a trade-off exists between the response in arm length and the response in stomach size, and at least in part, explains the evolution of feeding structure plasticity in plutei. This may also explain why feeding-structure plasticity has evolved in larvae of other taxa (e.g. other echinoderms and gastropods).     

Phenotypic variation in the breeding phenology of the woodlouse Armadillidium vulgare

Summary The breeding phenology of temperate wood-lice is strongly seasonal, the result of physiological constraints and precise environmental cues for reproduction. The adaptive value of such mechanisms is that the release of offspring coincides with favourable conditions for growth and survival (Willows 1984). We recorded the breeding phenology of Armadillidium vulgare (Latreille) on two grassland sites in Great Britain and found between-site and between-year variation in the onset of reproduction, the duration of reproductive activity, the release of offspring, the size of reproductive females and the number of broods per female. Between 82.7 and 97.7% of gravid females sampled were semelparous at 23 months, with the remainder iteroparous, producing a second brood after 35 months. On one site (Weeting Health) improved growth conditions during 1984 allowed some females (19.3% of gravid females sampled in that year) to produce a brood after 11 months. There was also an increase in the number of 3-year-old females found to be gravid. An experimental manipulation of the same habitat confirmed that such changes in life history tactics could be phenotypic responses. The observed phenotypic variation was sufficient to produce a range of life history tactics within a population. Mixtures of life history tactics within a population may be typical of invasive species and populations at the edge of the species range. Our results support the idea that phenotypic plasticity can be an appropriate tactic to maximise fitness in a fluctuating environment (Caswell 1983, 1989).     

Life-history traits of the lizard Sceloporus undulatus from two populations raised in a common laboratory environment

Hatchling Sceloporus undulatus elongatus from Washington Co., Utah and S. u. garmani from Woods Co., Oklahoma were raised to maturity and reproduction under identical laboratory conditions with ad libitum food availability. Growth, allometry, age and size of maturity, clutch size and egg mass were compared among lab-raised cohorts from the two populations, among lab-raised and field-caught animals (including their field-caught mothers) and, for growth, with values obtained by previously published field studies on the same or nearby populations. For all traits population differences observed in previous field studies and current field samples resulted from both a plastic response to proximate environmental conditions and intrinsic (possibly genetic) difference. The most plastic traits were growth and age of maturity. Cohorts from both populations expressed the ability to mature in less than 6 months in the laboratory but only the S.u. garmani express early maturity in the field. Allometric differences generated during growth in the lab were not observed in field samples but may reflect an adaptive physiological difference. The least plastic trait was egg mass. The only trait for which the rank order of the difference in the field was reversed in the lab was growth rate. S.u. elongatus grew significantly faster than S.u. garmani in the lab but much slower in the field. The tendency of S.u. garmani females to breed at minimum size of maturity may be greater than that of S.u. elongatus.