Life history characteristics of Daphnia magna clones differing in phototactic behaviour

In an analysis of a life table experiment involving positively, intermediately and negatively phototactic Daphnia magna clones, life history traits such as the average duration of the adult instar, neonate and adult body size were found to be correlated with phototactic behaviour. The size of the eggs and neonates was positively correlated with adult body size, and with egg development time. Adult body size was positively correlated with the size of the second and subsequent clutches. I argue that the intrinsic positive correlation between offspring size and egg development time is a key factor structuring the differences in life history patterns observed between the positively and intermediately phototactic Daphnia genotypes, and that the two life history patterns are to be considered alternatives suited for different environmental conditions (e.g. habitats with and without strong predation pressure on adults).     

Evidence for sex-related differences in phototactic bahaviour ofStreptocephalus proboscideus (Crustacea: Anostraca)

The phototactic behaviour of adults of the Sudanese fairy shrimpStreptocephalus probiscideus was studied under laboratory conditions. Males were less negatively phototactic than females. This was also evident when colour filters were used. females only became little less negatively phototactic under yellow light, whereas males showed a strong positively phototactic response. The response to the positioning of a yellow filter was stronger than to the use of a red or blue filter for both sexes. The laboratory findings were compared with casual field observations onStreptocaphalus torvicornis that indicate differential vertical distribution between the sexes and a nocturnal vertical migration. Migratory behaviour with ascent starting at dusk is also predicted forS. proboscideus. This behaviour may reduce common stress factors in desert pools such as photodamage, visual predation pressure, and high surface temperatures.     

An analysis of the phototactic behaviour of Daphnia magna clones and their sexual descendants

Daphnia magna clones, isolated from different natural populations, were inbred and crossbred. Some aspects of the heredity of the phototactic behaviour, quantified by an index, were analysed. Heritability in the broad sense was determined for several assemblages of clones through calculation of the clonal repeatability. Heritability sensu strictu was determined by offspring-parent regressions. Frequency distributions of a phototactic index of several inbred and crossbred families are given.The results indicate a significant contribution of additive genetic variance to the total phenotypic variance. Comparison of heritability estimates suggests an important genetic interaction component.Some implications of these results on the variability of vertical migration patterns in natural populations are discussed.     

Climate and mammalian life histories

Mammals display considerable geographical variation in life history traits. To understand how climatic factors might influence this variation, we analysed the relationship between life history traits – adult body size, litter size, number of litters per year, gestation length, neonate body mass, weaning age and age at sexual maturity – and several environmental variables quantifying the seasonality and predictability of temperature and precipitation across the distribution range of five terrestrial mammal groups. Environmental factors correlated strongly with each other; therefore, we used principal components analysis to obtain orthogonal climatic predictors that could be used in multivariate models. We found that in bats, primates and even‐toed ungulates adult body size tends to be larger in species inhabiting cold, dry, seasonal environments, whereas in carnivores and rodents a smaller body size is characteristic of warm, dry environments, suggesting that low food availability might limit adult size. Species inhabiting cold, dry, seasonal habitats have fewer, larger litters and shorter gestation periods; however, annual fecundity in these species is not higher, implying that the large litter size of mammals living at high latitudes is probably a consequence of time constraints imposed by strong seasonality. On the other hand, the number of litters per year and annual fecundity were greater in species inhabiting environments with higher seasonality in precipitation. Lastly, we found little evidence for specific effects of environmental variability. Our results highlight the complex effects of environmental factors in the evolution of life history traits in mammals. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 719–736.     

Inter-clonal variation in phototactic behaviour and key life-history traits in a metapopulation of the cyclical parthenogen Daphnia ambigua: the effect of fish kairomones

We quantified genetic variation for ecological relevant traits in the presence and the absence of fish chemicals of eleven Daphnia ambigua clones that were isolated from six interconnected ponds that differed in water transparency. In a cohort life table experiment, we tested whether genetic variation for a set of key life history traits was present among these clones. In addition the phototactic behaviour of these clones cultured in the presence and the absence of fish kairomones was quantified using a biotest. We detected a significant effect of fish kairomones on the phototactic behaviour and a highly significant genetic variation among clones for this trait in D. ambigua clones isolated from ponds in De Maten. Differences in size at maturity among D. ambigua clones in De Maten were highly significant, whereas differences in spina length among D. ambigua clones in De Maten were not significant. The presence of fish chemicals did not affect the studied life-history traits. We observed a significant positive relationship between average phototactic behaviour for each population and size at maturity both in the presence as in the absence of fish kairomones. Most of the genetic differences could be attributed to a clone isolated from one clearwater pond that is not directly connected to the remainder of the pond complex.     

Environmental networks,compensating life histories and habitat selection by white-footed mice

Summary Analysis of 6 years' data on a population of free-living white-footed mice documents both phenotypic and environmental control of litter size. Litter size was positively correlated with maternal body size. Maternal size depended upon both seasonal and annual variation. Paradoxically, the proportion of small versus large litters varied among habitats independently of the effects of body size. The result is an influence of habitat on life history that yields patterns of reproduction and survival opposite to the predictions of demographic theory. The habitat producing the largest litters had a relatively high ratio of adult/juvenile survival. Litter size was small in the habitat where the adult/juvenile survival ratio was smallest. All of these anomalous patterns can be explained through density-dependent habitat selection by female white-footed mice. Life-history studies that ignore habitat and habitat selection may find spurious correlations among traits that result in serious misinterpretations about life history and its evolution.     

Paravortex karlingi sp. nov. from Cerastoderma edule L., in Britain

Paravortex karlingi sp. nov. collected from the intestine of the bivalve mollusc Cerastorderma edule from the Ythan Estuary, N. E. Scotland, and elsewhere, is distinguished from a closely related species, P. cardii, also occurring in this host, on the basis of differences in habitat occupied by the two species as well as behavioural and morphological differences. P. karlingi is smaller, has fewer embryos in the gravid adult and shows a different behaviour pattern when released from the host intestine. It is also negatively phototactic whereas P. cardii is initially positively phototactic, only later becoming negatively phototactic. The occurrence of both species in Britain is briefly described.Abbreviations bc brood capsules - c core of indigestible particles - ca common atrium - ch chamber - e eye - fa female atrium - fol fibre optic light - gc gut caecum - gp genital pore - j juvenile worm - m mouth - o ovary - oe oesophagus - p pharynx - pe penis - sg shell gland - sv seminal vesicle - t testis - u uterus - vg vitelline gland - w window     

Variation and covariation of life history traits in aphids are related to infection with the facultative bacterial endosymbiont Hamiltonella defensa

Host–symbiont associations play an important role in insects. In aphids, facultative symbionts affect host plant use and increase thermal tolerance and resistance to natural enemies. In spite of these beneficial effects on aphid fitness, the frequency of facultative symbionts in aphids ranges from low to intermediate. Tradeoffs induced by symbionts could prevent the fixation of symbionts in aphid populations. Therefore, we studied the life history traits and correlations between them in 21 clones of the black bean aphid, Aphis fabae, seven of which were infected with the facultative endosymbiont Hamiltonella defensa. We found that clones harbouring H. defensa exhibited significantly higher body mass at maturity and offspring production, and a marginally higher intrinsic rate of increase. However, development time and offspring body size did not differ between symbiont‐free and infected clones. In addition, body mass at maturity was positively correlated with offspring production, offspring body size and intrinsic rate of increase, whereas development time was negatively correlated with body mass at maturity, offspring production and offspring body size. Excluding infected clones had little effect on these correlations; only correlations between body mass at maturity and offspring production, and between development time and offspring body size, became nonsignificant. Therefore, we did not find any evidence for tradeoffs between life history traits induced by symbiont infection. In fact, infected clones had higher overall fitness than symbiont‐free clones under the conditions of our experiment, suggesting that symbionts do not impose costs on aphids harbouring them. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 237–247.     

Evolution of predator avoidance in a Daphnia population: evidence from the egg bank

Dormant propagule pools (egg banks) of zooplankton populations that accumulate in sediments provide biological archives of past conditions and enable the investigation of evolutionary changes in populations over relatively long periods of time (many decades). This study examined the egg bank of a Daphnia pulicaria population in a lake that has been stocked annually with rainbow trout (a zooplanktivore) since 1961. Resting eggs from sediments from the 1920s to 2001 (determined by ²¹⁰Pb dating) were hatched and established as isofemale clonal lines. The phototactic behavior (a proxy for vertical migration behavior) of clones was assessed in the presence and absence of fish kairomones to evaluate the hypothesis that the heavy and consistent level of predation imposed by the stocked trout would have selected for clones that are more negatively phototactic. In addition, exposure to kairomones was expected to induce stronger negative phototaxis for all clones relative to control conditions. The behavior of clones derived from resting eggs from the trout era (after 1961) was significantly more negatively phototactic than it was for pre-trout era clones. Kairomone exposure induced a more negative phototactic response in clones from both eras, but the response was much greater for the pre-trout era clones. These results suggest that the consistently high level of predation by trout over several decades has selected for a conservative (less plastic) vertical migration strategy in which Daphnia maintain a deep daytime distribution with or without the presence of chemical cues from fish predators.     

Life history variation among low-arctic clones of obligately parthenogenetic Daphnia pulex: a diploid-polyploid complex

Summary Laboratory life table experiments were conducted using nine clones of obligately parthenogenetic Daphnia pulex that were collected from a site in the Canadian low-arctic. Two of the nine clones were diploids, while the other seven clones were polyploids. Significant clonal differences in age at first reproduction, size at first reproduction, number of offspring in each of the first three broods, offsrring sizes for the first two broods, and intrinsic rates of natural increase were detected. Differences in life histories were evident between polyploids and diploids. Generally, polyploid clones reached maturity at later ages, matured at larger sizes, produced smaller broods, and larger offspring than the diploid clones. The data are discussed in reference to potential biotic (i.e. invertebrate predation) and abiotic factors (i.e. physicochemical gradients) that may influence life history variation in this clonal assemblage.     

Phototactic responses in Haematococcus lacustris and its modification by light intensity and the carotenoid biosynthesis inhibitor Norflurazon

At fluence rates below 45 W· m^-2 cells of the flagellate stage of Haematococcus lacustris react only positively phototactically with a rather high degree of orientation (indicated by r values up to 0.66 with the Rayleigh test). The directedness of orientation decreases with decreasing irradiance. The degree of directedness of the phototactic response depends on the intensity of preirradiation: Low light intensity applied after strong light application results in a dark reaction (low r values), low light given after darkness stimulates a rather high degree of directedness of positive phototaxis. Weak blue light (=483 nm; 0.4 W · m^-2) stimulates positive phototactic response, whereas comparable red light (=658 nm; 0.5 W · m^-2) does not.Cells which were grown in a medium containing 10^-4 M Norflurazon (effective in inhibition of carotenoid biosynthesis) although maintaining motility completely lose the ability to react positively phototactically. The possible role of carotenoids in the phototactic orientation is discussed.     

The Rate-Size Trade-Off Structures Intraspecific Variation in Daphnia ambigua Life History Parameters

The identification of trade-offs is necessary for understanding the evolution and maintenance of diversity. Here we employ the supply-demand (SD) body size optimization model to predict a trade-off between asymptotic body size and growth rate. We use the SD model to quantitatively predict the slope of the relationship between asymptotic body size and growth rate under high and low food regimes and then test the predictions against observations for Daphnia ambigua. Close quantitative agreement between observed and predicted slopes at both food levels lends support to the model and confirms that a ‘rate-size’ trade-off structures life history variation in this population. In contrast to classic life history expectations, growth and reproduction were positively correlated after controlling for the rate-size trade-off. We included 12 Daphnia clones in our study, but clone identity explained only some of the variation in life history traits. We also tested the hypothesis that growth rate would be positively related to intergenic spacer length (i.e. the growth rate hypothesis) across clones, but we found that clones with intermediate intergenic spacer lengths had larger asymptotic sizes and slower growth rates. Our results strongly support a resource-based optimization of body size following the SD model. Furthermore, because some resource allocation decisions necessarily precede others, understanding interdependent life history traits may require a more nested approach.     

Life history responses of the cladoceran Ceriodaphnia cf. dubia to variation in food concentration

The effect of five different food concentrations on the life history of Ceriodaphnia cf. dubia was examined by following cohorts of 25 individuals from <24 hours until the death of all individuals. The food concentrations used in the study were chosen to reflect densities found in lentic freshwater systems and those commonly used in toxicity testing, and ranged from 1 × 10⁴ cells mL^-1 to 15 × 10⁴ cells mL^-1. Food concentration was found to have a significant effect (p<0.05) on several life history parameters, with a decrease in food concentration leading to a decrease in brood sizes and population growth rate, and an increase in longevity. Population growth rates varied from approximately 0.39 neonates d^-1 to 0.54 neonates d^-1, while mean lifespan ranged from 16.7 days to 42.9 days. A decrease in food concentration also led to an increase in the mean generation time.     

Fitness consequences of alternative life histories in water striders,Aquarius remigis (Heteroptera: Gerridae)

Using field and laboratory observations and experiments over 3 years, I investigated whether reproductive trade-offs shape individual life histories in two natural populations of the water strider, Aquarius remigis, in which univoltine and bivoltine life cycles coexist. Both later eclosion dates and food shortages, even after adult eclosion, induced diapause in females, thus deferring reproduction to the following spring. Adult body size was positively affected by food availability during juvenile development. Higher food levels also increased the reproductive output of females, but not their longevity or oviposition period. When compared to spring breeders (univoltine life cycle), direct (summer) breeders (bivoltine life cycle) experienced reduced lifetime egg numbers and longevity, as well as reduced survivorship of their second-summer-generation offspring; these reproductive costs offset, at least in part, the advantage in non-decreasing populations of having two generations per year. Fecundity was correlated with body size, and among summer-generation females direct breeders were larger than non-breeders. The time remaining before the onset of winter and/or the time since adult eclosion augmented cumulative energy uptake, and consequently the lipid reserves and winter survival probability of non-breeding (diapausing) summer adults approaching hibernation. Overwintered spring reproductives died at faster rates than non-reproductive summer individuals despite greater food availability in spring, indicating a mortality cost of reproduction. Body length correlated with absolute and not with proportional lipid content but showed no consistent relationship with survivorship in the field. These results are in agreement with current theory on the evolution of insect voltinism patterns, and further indicate high degrees of life history flexibility (phenotypic plasticity) in the study populations in response to variable environmental factors (notably photoperiod and food availability). This may be related to their location in a geographic transition zone from uni- to bivoltine life cycles.     

Patterns of life history among cyclopoid copepods of central Europe

• 1 Life history characters (body size of adults, egg diameter, egg sac length and breadth) of nineteen species of central European cyclopoid copepods were measured and sexual size dimorphism (adult female length x adult male length^?1), relative egg size (egg weight X body weight^?1), weight of adult females and of eggs, egg sac shape (egg sac length x egg sac breadth^?1), and reproductive effort (clutch weight produced per female weight per day) were calculated to detect trends in life history strategies.
• 2 Typical planktonic species exhibited the lowest reproductive effort. Among planktonic species, the value for egg sac shape increased with clutch size.
• 3 Large species and small species exhibited different trends in life history characters. Large species had larger clutches, larger eggs, and a greater sex size dimorphism than small species. However, small species had a greater relative egg size.
• 4 Large species live in cold water and reproduce during the spring bloom of phytoplankton where the production of large clutches with relatively small eggs is advantageous. Reserves are unnecessary for juveniles because food is abundant. Small species generally are most abundant during the warm season, when conditions are less predictable, and relatively large eggs, possibly provided with reserves, are advantageous.

     

An estimation of the heritability of phototaxis in Daphnia magna Straus

Summary The phototactic responses of four clones of Daphnia magna were experimentally analysed. Broad-sense heritability of this behavioural character was estimated through an analysis of variance, and it was very high under the standardised experimental conditions of this study.     

Offspring size–number tradeoffs and food quality feedbacks impact population dynamics in a Daphnia–algae system

Population fluctuations can be affected by both extrinsic (e.g. weather patterns, food availability) and intrinsic (e.g. life‐history) factors. A key life‐history tradeoff is the production of offspring size versus number, ranging from many small offspring to few large offspring. Models show that this life‐history tradeoff in offspring size and number, through maturation time, can have significant impacts on population dynamics. However, few manipulative experiments have been conducted that can isolate life‐history effects from impacts of extrinsic factors in consumer–resource systems. We experimentally tested the effect of an offspring size–number tradeoff on population stability and food availability in a consumer–resource system. Using Daphnia pulex, we created a shift from many, small offspring being produced to fewer, larger offspring. Two sets of experiments were performed to examine the interaction of an extrinsic factor (light levels) and intrinsic population structure on dynamics, and we controlled for the ingestion pressure on algal prey at the time of the manipulation. We predicted that the tradeoff would impact internal consumer population characteristics, including biasing the stage structure towards adults, increasing adult size, and increasing average population‐level reproduction. This adult‐dominated stage structure was predicted to then lead to instability and a low quantity–high quality food state. Under all light levels, the manipulated populations became dominated by large adults. Contrary to predictions, the amplitudes of fluctuations in Daphnia biomass were lower in populations shifted to few–large offspring, representing higher stability in these populations. Furthermore, in high light conditions, a stable low Daphnia – high algae biomass (low food quality) state was observed in few–large offspring treatments but not in control (many–small offspring) treatments. Our results show a strong link between light levels as an extrinsic factor and the life‐history tradeoff of consumer offspring size versus number that impacts consumer–resource population dynamics through feedbacks with resource quality.     

The gene fl(2)d is required for various Sxl-controlled processes in Drosophila females

Summary In Drosophila melanogaster, the gene Sex-lethal (Sxl) controls the processes of sex determination, dosage compensation, oogenesis and sexual behaviour. The control of Sxl is by alternative splicing of its primary RNA. We have identified a gene, female-lethal-2-d (fl(2)d), which is needed for the female-specific splicing of Sxl RNA and which also has a vital function independent of Sxl. Here we analyse other aspects of the gene fl(2)d. Specifically, we have analysed the effect of the temperature-sensitive mutation fl(2)d ¹ on the viability of adult flies homozygous for this mutation. We have found that the viability of the mutant females is reduced, while that of the mutant males is not affected. In addition, the capacity of the mutant females to be inseminated is considerably reduced, whilst all the mutant males are able to inseminate females. These effects on females are suppressed by Sxl ^M1. However, the fat body cells of fl(2)d ¹ homozygous females are able to synthesize yolk proteins at the restrictive temperature. We have also carried out, in males, a clonal analysis of fl(2)d ², a mutation lethal in both sexes. We have found that the clones are fully viable. We conclude that the gene fl(2)d seems to be necessary during the adult life of females for the processes that require Sxl ⁺ activity. Moreover, the Sxl-independent vital function of fl(2)d seems to be required in both sexes only during larval development. Offprint requests to: L. Sánchez     

Behaviour in a standardized assay,but not metabolic or growth rate,predicts behavioural variation in an adult aquatic top predator Esox lucius in the wild

This study tested for links among behaviour, state and life‐history variables as predicted by the pace‐of‐life hypothesis in adult pike Esox lucius. First, a standardized open‐field behavioural assay was developed to assess individual behaviour of wild‐captured adult E. lucius. Behaviour within the standardized assay predicted swimming behaviour in the lake, providing an ecological validation of the assay. There was no relationship between standardized behaviour and any of the life‐history and state variables, including metabolism, body condition, juvenile growth rate and adult growth rate in contrast to predictions from the pace‐of‐life hypothesis. This study demonstrates that it is possible to assess ecologically relevant behavioural variation in a large‐bodied top predator using a standard open‐field assay, but it is noteworthy that this standardized behaviour is not systematically related to standard metabolism or growth.     

Different growth responses to temperature and resource limitation in three fly species with similar life histories

Growth responses to temperature and resource limitation in three dipteran species with similar life histories were compared. With respect to current life history theory, two points are raised. First, growth rate in real time increased steeply with temperature in all species, following the standard pattern. However, when expressed in physiological time growth rate increased as temperature decreased in the yellow dung fly Scathophaga stercoraria, remained approximately constant in Sepsis cynipsea, and increased in Drosophila melanogaster. These responses can be understood as adaptations to climate and seasonality. It is concluded that some patterns of adaptation may be more easily interpreted if, and some may even go undetected unless, they are analysed in physiological time. Second, a decrease in body size, development rate and growth rate when resources are limited is believed to be nearly universal and generally predicted by life history models. Despite their similar life histories, the three species investigated showed qualitatively different growth responses to larval food shortage. At unlimited resources, yellow dung flies showed the fastest initial larval body mass gain per unit time, while those of S. cynipsea and D. melanogaster were lower and about equal. The period of no body mass gain at the end of larval development was longest in S. stercoraria and shortest in S. cynipsea. When facing resource limitation, S. stercoraria emerged smaller but earlier (thus nearly maintaining their growth rate), S. cynipsea smaller after the same development period, and D. melanogaster smaller and later (showing reduced and much reduced growth, respectively). It is concluded that whether growth really slows when resources are limited depends on the precise ecological circumstances of the species in question. More refined models, particularly those where mortality costs are independent of time, and more experiments are necessary to account for the variation in growth and size and age at maturity present in nature. This revised version was published online in July 2006 with corrections to the Cover Date.