Dispersal duration mediates selection on offspring size

Offspring size varies at all levels of organisation, among species, mothers and clutches. This variation is thought to be the result of a tradeoff between offspring quality and quantity, where larger offspring perform better but are more costly to produce. Local environmental conditions alter the benefits of increased offspring size and thereby mediate selection on this trait. For sessile organisms, dispersal is a crucial part of the offspring phase, and in animals, bigger offspring tend to better endure longer dispersal distances than smaller offspring because they have more energy. Theory predicts that increasing distances between suitable habitats strengthens selection for larger offspring. We manipulated the dispersal duration of offspring of different sizes in the bryozoan Watersipora subtorquata and then examined the relationship between offspring size and post‐metamorphic performance in the field. We found that selection on offspring size is altered by larval experience. Larger offspring had higher post‐settlement performance if the larval period was short but, contrary to current theory, performed worse when the larval period was extended. The reversal of the relationship between offspring size and performance by extending the larval phase in Watersipora may be due to the way in which offspring size affects growth in this species. Regardless of the mechanism, it appears that experiences in one life‐history stage alter selection on offspring size in another stage, even when they occupy identical habitats as adults.     

Chick growth rates in Charadriiformes: comparative analyses of breeding climate, development mode and parental care

The rate at which a young bird grows is highly diverse across taxa. We investigated the influences of ecological variables on growth rates of shorebirds, gulls and their allies (order Charadriiformes) using comparative analyses of 68 species. We investigated three hypotheses: (1) exposure to cold temperatures results in reduced growth rate due to the increased energy expenditure required for thermoregulation, (2) fast growth rates allow offspring to complete development in habitats with short periods of fair ecological conditions, and (3) parental feeding allows the offspring to grow faster than self-feeding offspring. Charadriiform species are suitable for testing these hypotheses, because they breed in diverse environmental conditions that include Arctic and Antarctic habitats, temperate zones and tropics, and in some species the offspring feed themselves (precocial) whereas in others they are fed by the parents (semi-precocial). First, we tested the influence of ambient temperature on growth rate and we found that species breeding in cold habitats had faster growth rates than species breeding in warm temperatures. The relationship between growth rate and ambient temperature was not significantly different between precocial and semi-precocial offspring. Second, we tested the influence of the length of the breeding season on growth rate, and we found that species with shorter breeding seasons had faster growth rates than species with longer breeding seasons. Finally, we show that precocial offspring grew slower than semi-precocial offspring, and this relationship remains significant when the influences of ambient temperature and breeding season length are statistically controlled. In conclusion, our work, using phylogenetically corrected models, confirms that ambient environment and developmental mode of young together influence the growth of Charadriiform chicks, and their effects are additive.     

The effect of previous host plant on the fecundity of Myzus persicae and its ability to transmit beet yellows virus

Myzus persicae transferred to rape produced offspring with higher survival, larger weight and faster development than aphids transferred to beet, irrespective of whether they had been reared on rape or beet. Aphids transferred from rape to beet produced fewer offspring than those reared continuously on rape. On beet, rape-reared adults produced progeny with lower survival, slower development, smaller adult size and lower fecundity than the progeny of beet-reared individuals, but there were no differences in these characteristics associated with host plant after one generation. Aphids from rape were less than half as efficient in transmitting beet yellows virus as aphids reared continuously on beet but after one subsequent generation on beet this difference had disappeared completely. These results are discussed in relation to field conditions.     

Life-stage specific environments in a cichlid fish: implications for inducible maternal effects

Through environmentally induced maternal effects females may fine-tune their offspring’s phenotype to the conditions offspring will encounter after birth. If juvenile and adult ecologies differ, the conditions mothers experienced as juveniles may better predict their offspring’s environment than the adult females’ conditions. Maternal effects induced by the environment experienced by females during their early ontogeny should evolve when three ecological conditions are met: (1) Adult ecology does not predict the postnatal environmental conditions of offspring; (2) Environmental conditions for juveniles are correlated across successive generations; and (3) Juveniles occasionally settle in conditions that differ from the juvenile habitat of their mothers. By combining size-structured population counts, ecological surveys and a genetic analysis of population structure we provide evidence that all three conditions hold for Simochromis pleurospilus, a cichlid fish in which mothers adjust offspring quality to their own juvenile ecology. In particular we show (1) that the spatial niches and the habitat quality differ between juveniles and adults, and we provide genetic evidence (2) that usually fish of successive generations grow up in similar habitats, and (3) that occasional dispersal in populations with a different habitat quality is likely to occur. As adults of many species cannot predict their offspring’s environment from ambient cues, life-stage specific maternal effects are likely to be common in animals. It will therefore be necessary to incorporate parental ontogeny in the study of parental effects when juveniles and adults inhabit different environments.     

Mothers determine offspring size in response to own juvenile growth conditions

Through non-genetic maternal effects, mothers can tailor offspring phenotype to the environment in which young will grow up. If juvenile and adult ecologies differ, the conditions mothers experienced as juveniles may better predict their offspring's environment than the adult environment of mothers. In this case maternal decisions about investment in offspring quality should already be determined during the juvenile phase of mothers. I tested this hypothesis by manipulating juvenile and adult maternal environments independently in a cichlid fish. Females raised in a poor environment produced larger young than females raised without food limitations, irrespective of the feeding conditions experienced during adulthood. This maternal boost was due to a higher investment in eggs and to faster larval growth. Apparently, mothers prepare their offspring for similar environmental conditions to those they encountered as juveniles. This explanation is supported by the distribution of these fishes under natural conditions. Juveniles live in a different and much narrower range of habitats than adults. Therefore, the habitat mothers experienced as juveniles will allow them to predict their offspring's environment better than the conditions in the adult home range.     

Parasitoids induce production of the dispersal morph of the pea aphid, Acyrthosiphon pisum

In animals, inducible morphological defences against natural enemies mostly involve structures that are protective or make the individual invulnerable to future attack. In the majority of such examples, predators are the selecting agent while examples involving parasites are much less common. Aphids produce a winged dispersal morph under adverse conditions, such as crowding or poor plant quality. It has recently been demonstrated that pea aphids, Acyrthosiphon pisum, also produce winged offspring when exposed to predatory ladybirds, the first example of an enemy‐induced morphological change facilitating dispersal. We examined the response of A. pisum to another important natural enemy, the parasitoid Aphidius ervi, in two sets of experiments. In the first set of experiments, two aphid clones both produced the highest proportion of winged offspring when exposed as colonies on plants to parasitoid females. In all cases, aphids exposed to male parasitoids produced a higher mean proportion of winged offspring than controls, but not significantly so. Aphid disturbance by parasitoids was greatest in female treatments, much less in male treatments and least in controls, tending to match the pattern of winged offspring production. In a second set of experiments, directly parasitised aphids produced no greater proportion of winged offspring than unparasitised controls, thus being parasitised itself is not used by aphids for induction of the winged morph. The induction of wing development by parasitoids shows that host defences against parasites may also include an increased rate of dispersal away from infected habitats. While previous work has shown that parasitism suppresses wing development in parasitised individuals, our experiments are the first to demonstrate a more indirect influence of parasites on insect polyphenism. Because predators and parasites differ fundamentally in a variety of attributes, our finding suggests that the wing production in response to natural enemies is of general occurrence in A. pisum and, perhaps, in other aphids.     

Predator-induced morphological shift in the pea aphid

Aphids exhibit a polymorphism whereby individual aphids are either winged or unwinged. The winged dispersal morph is mainly responsible for the colonization of new plants and, in many species, is produced in response to adverse environmental conditions. Aphids are attacked by a wide range of specialized predators and predation has been shown to strongly influence the growth and persistence of aphid colonies. In two experiments, we reared two clones of pea aphid (Acyrthosiphon pisum) in the presence and absence of predatory ladybirds (Coccinella septempunctata or Adalia bipunctata). In both experiments, the presence of a predator enhanced the proportion of winged morphs among the offspring produced by the aphids. The aphid clones differed in their reaction to the presence of a ladybird, suggesting the presence of genetic variation for this trait. A treatment that simulated disturbance caused by predators did not enhance winged offspring production. The experiments indicate that aphids respond to the presence of a predator by producing the dispersal morph which can escape by flight to colonize other plants. In contrast to previous examples of predator-induced defence this shift in prey morphology does not lead to better protection against predator attack, but enables aphids to leave plants when mortality risks are high.     

Secondary sex ratio covaries with demographic trends and ecological conditions in the barn swallow

Parents are expected to invest more in the sex that benefits most from the local environment. When the quality of breeding sites varies spatially and natal dispersal of males and females differs, parents in high-quality habitats should skew their progeny sex ratio in favor of the less dispersing sex. We tested this prediction in the barn swallow (Hirundo rustica L.), by relating the proportion of male offspring around fledging (secondary sex ratio) of first and second broods to the ecological quality (presence of livestock farming and relative surface of hayfields in the foraging range) and local demographic trends of the farms where the colonies were located. Consistent with our predictions, the proportion of male offspring, which are more philopatric than females, increased with the extent of hayfields, which are high quality, preferred foraging habitats. Moreover, the proportion of male offspring in second broods was smaller in colonies with positive demographic trends, possibly indicating density-dependent effects on sex ratio. Independent of the mechanism generating uneven sex ratio (zygote sex ratio adjustment or sex-related pre-fledging mortality), barn swallows breeding under favorable conditions overproduced the sex that is more likely to benefit from such conditions.     

Sex-specific dispersal in spatially varying environments leads to habitat-dependent evolutionary stable offspring sex ratios

When the environment varies spatially, so that some habitatsare more favorable to reproduction than others, an individualshould attempt to increase the number of offspring establishingin high-quality habitats. Hence, if male and female dispersalbehavior differ, it may be adaptive to produce more offspringof the more dispersing sex in low-quality habitats, since these offspring are likely to disperse to another patch, and moreoffspring of the most philopatric sex in high-quality habitats,since these offspring are likely to remain in that patch. Sucha strategy is shown to be evolutionarily stable provided thatmale and female dispersal rates are different and that reproductivesuccess varies between habitats (lack of ideal free distribution).Highly biased sex ratios are predicted (1) in rare habitats, (2) in poor habitats, (3) when difference between habitat qualityis large, (4) when at least one sex disperses at a rate closeto random with respect to habitat availability, (5) when bothsexes disperse at a high rate, (6) when individuals are unableto select their reproducing habitat, and, presumably, (7) withmoderate temporal variation of habitat quality. The model appearsto be a good candidate to explain the pattern of sex ratiovariation in a variety of species : phytophagous arthropods,species with environmental sex determination, and territorialpasserines.     

The effect of experimental warming on a low‐latitude aphid,Myzus varians

Tropical invertebrates are currently living very close to their optimal temperature. Warming due to global climate change will likely exceed their physiological optima and have deleterious consequences for insects living at low latitudes. In this study, we assess the effects of various levels of summer warming predicted for the late 21st century (+1.2 and +3.7 °C, with the same diel oscillation as the current regime) on the physiology and demography of the aphid Myzus varians Davidson (Hemiptera: Aphididae) in subtropical and tropical Taiwan. Aphids subjected to a moderate (3.7 °C) increase in temperature did not reach adulthood and thus left no offspring, meaning that wild populations could go extinct during the summer. Slight (+1.2 °C) warming did not significantly affect development time and generation time. However, warming reduced nymphal survival, adult longevity, and reproduction, and, thus, reduced the fitness of aphid populations. Aphids have a number of adaptations for surviving or avoiding unfavorable conditions. However, predicted increases in global temperatures will likely decrease their survival and reproduction, which could increase the frequency of local extinction.     

The role of sink to source re-colonisation in the population dynamics of insects living in unstable habitats: an example of terrestrial chironomids

Certain species of terrestrial chironomids (Diptera) are specialised on open patches in initial stages of primary or secondary succession (early fallow, lichens and mosses on rocks, etc.). These "source" habitats provide good quality food for their larvae and most offspring are produced here, but they are sensitive to summer desiccation. This often results in extinction of the summer larval population, followed in winter by re-colonisation from less suitable, but more stable "sink" habitats in the surrounding landscape. Soil dwelling and long-lived larvae are poor migrants; short-lived, winged adult females select patches for their development. Proper choice of oviposition sites and consequent distribution of eggs among individual habitats is thus critical for the success of these species. A mathematical model was developed in order to find out whether this re-colonisation strategy could ensure population persistence at the landscape level. The model was verified using long-term data on Smittia atterima abundance in old fields. The results indicate that even a small proportion of eggs laid in a sink habitat can ensure a successful re-colonisation of the source habitat. Thus, re-colonisation of source habitats from sink habitats is concluded to be one of the reasons for persistence of the latter. The model indicates that this re-colonisation may ensure population persistence even in conditions when exclusive use of only one habitat leads to population extinction either due to environmental stress or to a negative growth rate.     

Body colour and genetic variation in winged morph production in the pea aphid

Aphids (Homoptera: Aphidoidea) produce a number of different phenotypes in their life-cycle, among which are winged (alate) and wingless (apterous) morphs. Lowe & Taylor (1964) and Sutherland (1969a, b) were the first to suggest that aphid clones differ in their propensity to produce the winged morph and that in the pea aphid (Acyrthosiphon pisum Harris), this propensity is linked to the colour of the phenotype. We tested for the occurrence of genetic variation in winged morph production by rearing individuals from red and green clones of pea aphid under wing-inducing (crowding) and control conditions, and scored the phenotypes of their offspring. Clones differed significantly in alate production and red clones produced on average a higher proportion of winged morphs than green clones. Importantly, however, there was considerable variation between clones of the same colour. Broad-sense heritabilities of winged morph production were 0.69 (crowding treatment) and 0.63 (control). Clones also differed in the number of offspring they produced. When exposed to the crowding stimulus, aphids deferred offspring production, resulting in a higher number of offspring produced in the crowding treatment than in the control.     

Maternal effects provide phenotypic adaptation to local environmental conditions

In outcrossing plants, seed dispersal distance is often less than pollen movement. If the scale of environmental heterogeneity within a population is greater than typical seed dispersal distances but less than pollen movement, an individual's environment will be similar to that of its mother but not necessarily its father. Under these conditions, environmental maternal effects may evolve as a source of adaptive plasticity between generations, enhancing offspring fitness in the environment that they are likely to experience. This idea is illustrated using Campanula americana, an herb that grows in understory and light-gap habitats. Estimates of seed dispersal suggest that offspring typically experience the same light environment as their mother. In a field experiment testing the effect of open vs understory maternal light environments, maternal light directly influenced offspring germination rate and season, and indirectly affected germination season by altering maternal flowering time. Results to date indicate that these maternal effects are adaptive; further experimental tests are ongoing. Evaluating maternal environmental effects in an ecological context demonstrates that they may provide phenotypic adaptation to local environmental conditions.     

Exploitation of patchy soil water resources by the clonal vine <Emphasis Type="Italic">Ficus tikoua</Emphasis> in karst habitats of southwestern China

The karst habitats of southwestern China are characterized by a highly heterogeneous distribution of water resources. We hypothesized that the clonal integration between connected ramets of the clonal vine Ficus tikoua was an important adaptive strategy to the patchy distribution of water resources in these habitats. We grew ramet pairs (each consisting of a parent and an offspring ramet) in both homogeneously and heterogeneously watered conditions. The offspring ramets were well-watered, whereas their connected parent ramets were randomly assigned to four water treatments: well-watered, mild water stress, moderate water stress, and severe water stress. Increasing water stress decreased leaf water potential, relative water content, net assimilation rate, maximum quantum yield of PSII (F _v/F _m), and biomass of the parent ramets. Subjecting the parents to water stress significantly increased root biomass and root mass ratio (RMR) of their offspring ramets. Exploitation of plentiful water resources through the increased adventitious roots connected to another soil patch permitted the complete restoration of water relations and photosynthetic capacity of offspring ramets after an initial depression. Water relations and gas exchange of the parents were not affected by the water supply to their connected offspring ramets, suggesting that offspring ramets hardly exported water to the stressed parents. However, net assimilation rate and proline content of the offspring ramets increased when they were connected to water-stressed parents. The compensatory photosynthetic responses of offspring ramets connected to stressed parents revealed an increasing trend as the experiment progressed. Morphological and physiological plasticity of F. tikoua in response to heterogeneous water resources allow them to adapt to karst habitats and be suitable candidates for vegetation restoration projects.     

Should females prefer to mate with low-quality males?

We analyse a model of mate choice when males differ in reproductive quality and provide care for their offspring. Females choose males on the basis of the success they will obtain from breeding with them and a male chooses his care time on the basis of his quality so as to maximise his long-term rate of reproductive success. We use this model to establish whether high-quality males should devote a longer period of care to their broods than low-quality males and whether females obtain greater reproductive success from mating with higher quality males. We give sufficient conditions for optimal care times to decrease with increasing male quality. When care times decrease, this does not necessarily mean that high-quality males are less valuable to the female because quality may more than compensate for the lack of care. We give a necessary and sufficient condition for high-quality males to be less valuable mates, and hence for females to prefer low-quality males. Females can prefer low-quality males if offspring produced and cared for by high-quality males do well even if care is short, and do not significantly benefit from additional care, while offspring produced and cared for by low-quality males do well only if they receive a long period of care.     

Poor maternal environment enhances offspring disease resistance in an invertebrate

Natural populations vary tremendously in their susceptibility to infectious disease agents. The factors (environmental or genetic) that underlie this variation determine the impact of disease on host population dynamics and evolution, and affect our capacity to contain disease outbreaks and to enhance resistance in agricultural animals and disease vectors. Here, we show that changes in the environmental conditions under which female Daphnia magna are kept can more than halve the susceptibility of their offspring to bacterial infection. Counter-intuitively, and unlike the effects typically observed in vertebrates for transfer of immunity, mothers producing offspring under poor conditions produced more resistant offspring than did mothers producing offspring in favourable conditions. This effect occurred when mothers who were well provisioned during their own development then found themselves reproducing in poor conditions. These effects likely reflect adaptive optimal resource allocation where better quality offspring are produced in poor environments to enhance survival. Maternal exposure to parasites also reduced offspring susceptibility, depending on host genotype and offspring food levels. These maternal responses to environmental conditions mean that studies focused on a single generation, and those in which environmental variation is experimentally minimized, may fail to describe the crucial parameters that influence the spread of disease. The large maternal effects we report here will, if they are widespread in nature, affect disease dynamics, the level of genetic polymorphism in populations, and likely weaken the evolutionary response to parasite-mediated selection.     

Facultative bacterial endosymbionts benefit pea aphids Acyrthosiphon pisum under heat stress

Abstract 1. Natural populations of pea aphids in California contain at least two facultative bacterial secondary symbionts (pea aphid secondary symbiont, PASS, or pea aphid rickettsia, PAR) in a range of frequencies throughout the state.
2. Two pea aphid clones without either of these facultative associates failed to reproduce in the first 8 days after the final moult if they had been heat-stressed for a period of about 4 h at 39 °C as 1-day-old larvae in the laboratory.
3. Aphids infected artificially with PASS, however, were able to produce up to 48% of the normal complement of offspring produced by PASS-positive aphids that had not been heat-stressed. Clones infected artificially with PAR did not have the same advantage as those with PASS after heat stress.
4. In aphids without PASS or PAR, heat stress reduced the number of bacteriocytes (in which the obligate primary symbiont, Buchnera , resides) to 7% of non-heat-stressed aphids, while aphids with only PASS retained 70% of their bacteriocytes. Bacteriocytes in aphids with PAR but not PASS were reduced to 42% of controls.
5. When larvae were heat-stressed as older instars (5 days old), a similar pattern emerged, though the effect of heat stress was less extreme. Clones containing PASS produced the most offspring, three to 14 times as many as aphids without PASS or PAR. Aphids with PAR only, or PASS and PAR together, had reduced or no advantage over aphids without facultative symbionts.
6. Aphids of all clones that had been heat-stressed as later instars gave birth to a variable number of stillborn offspring. Aphids without facultative symbionts produced the most stillborn larvae.
7. Field studies showed a higher incidence of PASS in aphids collected in California in summer compared with aphids from the same sites collected 2–4 months earlier. The difference was significant in two of three widely dispersed locations.     

The effect of female polyandry and sperm precedence on the evolution of sexual difference in dispersal timing

Predispersal copulation and unpredictable environment facilitate the evolution of female-biased dispersal in species, where females are functionally monandrous. Females should migrate and reproduce over different habitats to spread their risks due to environmental fluctuation. On the other hand, males do not have to disperse because their risks are spread by their mating partners who produce their offspring in different habitats. However, when females are functionally polyandrous, it is expected that they will not contribute to spreading the male's risk extensively. Therefore, by simulation with the individual based model, the present study evaluated how female polyandry influences the sexual difference in dispersal timing. This model revealed that when females are polyandrous, the timing of female remating and sperm priority patterns have an important influence on the evolution of sex-biased dispersal. Particularly when female remating is not synchronized with dispersal or when last-male sperm precedence does not exist, female-biased dispersal is evolved.     

Environmental Change and Extended Phenotypes: Does Eutrophication Influence Nest Building in Sticklebacks?

Many species use extended phenotypes, such as purpose‐built nests, to increase their reproductive success. These traits have to be adjusted to local environmental conditions to maximize fitness. An important question is whether species are able to adjust their extended phenotypes to human‐induced rapid environmental changes. We investigated whether populations of threespine stickleback, Gasterosteus aculeatus, exposed to different degrees of human‐induced eutrophication during the last decades, have differentiated phenotypically in their nest‐building behaviour. Stickleback males build nests that they use both in mate attraction and for offspring protection and whose characteristics vary with environmental conditions. We allowed males from parallel pairs of mildly and severely eutrophied habitats to build nests under standardized conditions in the laboratory. We recorded the time it took the males to build a nest, the size and neatness of the completed nest and the use of nest ornaments. We found eutrophication at the site of capture to influence nest‐building time – males from eutrophied habitats built faster. However, eutrophication did not alter nest structure or the use of nest ornaments. This is probably because the nests are concealed in vegetation or under stones and females cannot evaluate them before they have followed the male to the nest, and predators cannot detect them before close to the nest. Thus, reduced long‐range visibility does not influence the use of nests as mate choice cues or for offspring protection. This contrasts with a recorded effect of eutrophication on courtship behaviour, whose efficiency depends on long‐range visibility. This suggests that traits are adjusted to eutrophication depending on the influence of eutrophication on the function of the traits.