Cryptic maternal effects in Hippodamia convergens vary with maternal age and body size

Maternal effects can mold progeny phenotypes in various ways and may constitute ecological adaptations. By examining the effect of oviposition sequence on progeny produced by different size classes of female ladybird beetles (produced by controlling larval access to food), we show that maternal signals can change through adult life and alter the developmental programs of progeny, ostensibly to synchronize their life histories with predictable resource dynamics, thus maximizing maternal fitness. We also show that female body size, as determined by larval food supply, interacts with female age to influence progeny fitness. When fed ad libitum as adults, small females reared with limited food access laid fewer, smaller eggs than large females reared with ad libitum food access. Maternal body size interacted with oviposition sequence to influence progeny development, but the latter had greater impact. Eggs laid later by medium and large females hatched faster than those laid earlier, larvae fed longer in the fourth instar, their pupation period was shorter, total developmental time was reduced, and adults emerged with greater mass, most notably daughters. Oviposition sequence effects on progeny from small mothers were non‐significant for total developmental time and progeny mass. Only large mothers increased egg size over time and egg mass was not consistently correlated with developmental parameters, indicating that progeny phenotype was impacted by other, more cryptic, maternal signals. Such signals appear costly, as food limitation during development constrained not only fecundity and egg size but also maternal ability to manipulate progeny phenotype. The production of faster‐developing offspring that mature to larger sizes late in the oviposition cycle may be adaptive for exploitation of ephemeral aphid outbreaks with predictable dynamics of prey abundance and competition.     

The control of morph development in the parasitic nematode Strongyloides ratti.

The parasitic nematode Strongyloides ratti has a complex life cycle. The progeny of the parasitic females can develop into three distinct morphs, namely directly developing infective third-stage larvae (iL3s), free-living adult males and free-living adult females. We have analysed of the effect of host immune status (an intra-host factor), environmental temperature (an extra-host factor) and their interaction on the proportion of larvae that develop into these three morphs. The results are consistent with the developmental decision of larvae being controlled by at least two discrete developmental switches. One is a sex-determination event that is affected by host immune status and the other is a switch between alternative female morphs that is affected by both host immune status and environmental temperature. These findings clarify the basis of the life cycle of S. ratti and demonstrate how such complex life cycles can result from a combination of simple developmental switches.     

Embryogenesis and oögenesis in alate virginoparae,gynoparae, and oviparae of the aphid Myzus persicae, in relation to photoperiod

The study reports on the effects of prenatal and/or postnatal exposures to short-night or long-night conditions, and of crowding, on embryogenesis and oögenesis in alate virginoparae, gynoparae, and oviparae of a holocyclic strain of the green peach aphid, Myzus persicae, from Yakima, Washington State.In alate virginoparae raised at a density of 10–20 per radish seedling in a short-night regime (8 hr darkness per diem), 3–4 embryos occurred in each of their 10 ovarioles, when the aphids attained adulthood. More than 30 larvae were deposited by most of these alatae. However, in young adult gynoparae, raised at these densities in a long-night regime (15 hr darkness per diem), only one viable embryo (a presumptive ovipara) occurred per ovariole. The follicle containing this embryo was followed by 1–2 abnormal follicles in each ovariole, and the number of larvae deposited by a gynopara was generally less than 10. In young adult oviparae similarly raised under a long-night regime, only one egg typically occurred in each of their 10 ovarioles, and the eggs deposited by an ovipara (only after it had mated) generally numbered less than 10. Alate virginoparae and gynoparae contained an additional embryo in some of their ovarioles when these morphs were raised at a lower density (1–5 per plant).Presumptive gynoparae partially developed the reproductive features of alate virginoparae when transferred to a short-night regime at birth; the converse was true when presumptive alate virginoparae were transferred to a long-night regime early in larval life. Oviparae maintained in short nights from before birth developed the appearance of apterous virginoparae but still produced eggs rather than embryos. However, their oögenesis was enhanced and eggs (10–20) were deposited by them without prior mating. Under all regimes tested, oviparae were always deposited early in the larviposition sequence of their alate mothers, and the number of oviparae deposited never exceeded 15.The possible involvement of juvenile hormone in the regulation of these events and the ecological significance of the results are discussed.     

Density reductions by predatory trout increase adult size and fecundity of surviving caddisfly larvae in a detritus-based stream food web

1. In some situations, individuals surviving in environments where predation is intense can grow faster because the benefits of release from intraspecific competition outweigh costs associated with anti-predator responses. Whether these 'thinning' effects of predation occur in detritus-based food webs where resource renewal occurs independently of consumption by consumers was studied. We investigated how effects of predatory brown trout ( Salmo trutta ) on the larvae of the detritivorous stream caddisfly, Zelandopsyche ingens , influenced the size and fecundity of the caddisfly adults.
2. Trout substantially reduced the abundance of Z. ingens larvae, but adult male and female Z. ingens were significantly larger in trout streams compared to fishless streams. Females in trout streams had 33% more eggs than fishless stream females, and egg sizes were not significantly different. In mesocosms, Z. ingens larvae in low density treatments reflecting trout stream abundances grew significantly faster than larvae in high density treatments that were characteristic of fishless stream abundances. Non-lethal trout presence did not influence case building behaviour, feeding rates or growth or Z. ingens larvae, indicating non-lethal effects of predators were negligible.
3. Increased adult size and fecundity associated with trout stream individuals were probably a result of predator thinning of larval density indirectly releasing surviving Z. ingens from intraspecific competition. Thus, predator thinning did influence interactions between larvae in this detritus-based food web as larval growth was strongly density-dependent. However, extrapolating the total number of eggs potentially produced indicates the increased fecundity of females in trout streams would not compensate for losses of larvae to trout predation.     

Life histories and habitat selection in Daphnia: divergent life histories of D. magna clones differing in phototactic behaviour

To test the hypothesis of co-adaptation of life histories and daytime vertical distribution (vertical migration behaviour) in Daphnia, life history characteristics were analysed for two positively, three negatively, and four intermediately phototactic Daphnia magna clones. Clones with different phototactic behaviour were found to have divergent life history strategies, with positively phototactic clones being good exploiters under the non-limiting conditions provided in the laboratory, i.e. low density (1 ind./1), high food concentration (6,5–7 10⁵ Scenedesmus cells/ml, restored daily) and high temperature (20° C). They realized a high intrinsic rate of increase at a small adult body size through rapid development, at a cost of producing small neonates. Negatively and intermediately phototactic clones had larger adult body sizes, and produced larger neonates that were more starvation-resistant than those of positively phototactic clones. Selection for high intrinsic rate of increase in intermediately phototactic clones was mediated through the production of large clutches.     

Effects of aggregation pheromone on individual behaviour and food web interactions: a field study on Drosophila

Abstract.  1. The effects of an aggregation pheromone on individual behaviour and food web interactions were investigated in two ecological communities, using Drosophila melanogaster and D. simulans as focal species.
2. Fruit substrates with aggregation pheromone were significantly more attractive to adult D. melanogaster and D. simulans than control fruit substrates, and the response was positively dose dependent. Competing species and natural enemies were also significantly attracted to substrates with the aggregation pheromone of D. melanogaster and D. simulans .
3. Significantly more eggs were deposited on pheromone-treated fruits than on control fruits, and the microdistribution of eggs within fruits was correlated to the microdistribution of the pheromone. The aggregation pheromone induced more females to share the breeding site.
4. The extremely high densities of fruit flies in the large aggregations appeared to reduce the oviposition rate of females. Physical interactions with conspecific and heterospecifics were frequently observed in the aggregations, and often led to patch leaving of the fruit flies.
5. Competition for food among larvae occurred at high densities and parasitism was density dependent. Aggregation pheromones can be directly responsible for these patterns through their effects on the con- and heterospecific behaviour.
6. The combined results show that aggregation pheromones affect a multitude of aspects in the ecology of interacting animals. The importance of incorporating the communication signals in ecological theory of aggregations is discussed.     

Size and fecundity in the Jamaican gall-midge Asphondylia boerhaaviae

ABSTRACT. 1. The gall-midge Asphondylia boerhaaviae Möhn galls weeds belonging to the Nyctaginaceae in Jamaica and other parts of the Caribbean. The species breeds throughout the year with a generation length of < 1 month.
2. In adult females, plots of wing length on hind tibial length, of maximum potential fecundity (m.p.f.) on wing length, and of m.p.f. on ovariole number, suggested the presence of a small morph, which the use of discriminant function analysis and other investigations confirmed. There was also good evidence of two size morphs in adult males.
3. Both body size and ovariole number had significant effects on m.p.f. but these effects were different in the two morphs. While the food plants had significant influences on size and fecundity, these were relatively small. The m.p.f. of the small morph was 125.8±11.1 eggs, and of the large morph 318.7±8.3 (95% cl).
4. Laboratory experiments on the flight performance of newly-emerged midges indicated that the small morph was not dispersive and that about 80% of the large morph was dispersive. No differences in size, wing length or fecundity could be found between the two behavioural types of the large morph.
5. The nature of the polymorphism is unusual, particularly since the dispersive individuals potentially have similar (or greater) fecundity than the non-dispersive ones.
6. Estimates of achieved fecundity (eggs actually laid, rather than m.p.f.) were 67.8±9.6 (field, minimum) and 138.9±30.4 (laboratory), the latter estimate being based on large-morph females.     

Host selection and egg allocation behaviour by Aphytis melinus and A.lingnanensis: comparison of two facultatively gregarious parasitoids

Abstract. 1. The ovipositional and egg allocation behaviour of individual females of Aphytis melinus DeBach and A.lingnunensis Compere were compared.
2. Both Aphytis species exhibit the same behavioural sequence during oviposition.
3. Aphytis melinus laid most of its female eggs on the dorsum of a scale-insect beneath its cover, and most of its male eggs under the scale-insect's body. Aphytis lingnanensis also oviposited both dorsally and ventrally on scale-insect hosts, but female and male progeny arose with equal frequency from eggs laid in both locations.
4. Both A.melinus and A. lingnanensis are facultatively gregarious parasitoids. The degree of gregariousness depends on host size, i.e. the larger the host, the more the Iikelihood that several eggs will be deposited at each visit by the parasitoid.
5. When two eggs were laid during the same host visit, both A.melinus and A.lingnanensis laid one female and one male egg more often than would be expected under an assumption of random allocation of sexes.
6. Because A.melinus successfulIy utilize smaller hosts than A.lingnanensis to produce progeny, these parasitoids should not be considered ecological homo-logues, as suggested by DeBach & Sundby (1963).     

Effect of Food and Density on Development of Typical and Cannibalistic Salamander Larvae in Ambystoma tigrinum nebulosum

Two subspecies of the tiger salamander, Ambystoma tigrinum,have a distinctive polymorphic life history that can includefour adult morphs as well as typical and cannibalistic larvalmorphs. We evaluated the effect of environment on developmentof larval morphology in two laboratory experiments. In ExperimentI, 180 larvae were raised in individual 3-liter containers andfed one of three food levels. Larvae in Experiment II receivedone of two levels of food, and were raised at three densities:one larva per 3 liters of water (50 containers), three larvaeper 22 liters of water (18 containers), or seven larvae per22 liters of water (18 containers). Cannibalistic morphs developedonly in nine containers at the highest density, and their occurrencewas independent of the two food levels. Our results suggestthe typical and cannibalistic larvae which occur in some populationsof Ambystoma tigrinum nebulosum is an environmentally induceddevelopmental polymorphism that results from some individualsresponding to the environment differently than others. Thisdifference in response may or may not be associated with geneticdifferences between these morphs. Based on our results we cannotdiscriminate between two models that differ in their assumptionsabout the genetic background of individual larvae.     

Life-history consequences of predation for a subantarctic beetle: evaluating the contribution of direct and indirect effects

1. Recently, a small predatory beetle, Trechisibus antarcticus (Carabidae), was accidentally introduced onto the island of South Georgia, sub-Antarctic.
2. From the presumed site of introduction the beetle is invading the coastal lowland area, building up high densities locally in the tussock-forming grass Parodiochloa flabellata .
3. In the coastal area the endemic detritivorous/herbivorous beetle Hydromedion sparsutum (Perimylopidae) is common, especially in and beneath the tussocks.
4. The first three, out of six, larval instars of H. sparsutum are easily taken prey by the carabid.
5. In sites colonized by the carabid, total abundance and the ratio between larval and adult numbers of H. sparsutum are far lower, and its adult body size clearly larger, than in comparable sites where the carabid is absent.
6. Two hypotheses are proposed for explaining the increase in adult body size of H. sparsutum : (i) the increase is a direct effect of predation: selection by the predator favours large hatchlings and/or larvae with a high growth rate; and (ii) the increase is an indirect effect of predation: by lowering the density of H. sparsutum , predation has increased its per capita food supply, enabling a higher growth rate and a larger adult body size.
7. A food addition experiment in a carabid-free site showed availability of high quality food to be insufficient for sustaining the initial larval population.
8. In the laboratory, females from the predator-infested sites produced larger eggs and hatchlings than females from the carabid-free sites, but mass specific growth rates of the larvae were not higher.
9. Field and laboratory data give stronger support to the food hypothesis than to the size selectivity hypothesis.     

Age‐specific maternal effects interact with larval food supply to modulate life history in Coleomegilla maculata

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豚草条纹叶的生态学特性

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Superparasitism in Cotesia glomerata: response of hosts and consequences for parasitoids

Abstract.  1. Superparasitism occurs in Cotesia glomerata (Hymenoptera: Braconidae), a gregarious endoparasitoid of Pieris spp. (Lepidoptera: Pieridae). The response of P. brassicae larvae to superparasitism and the consequences for the parasitoid were examined in order to elucidate the ecological significance of this behaviour.
2. Field surveys of a Swiss population revealed that C. glomerata brood sizes from P. brassicae larvae ranged from three to 158, and both the female ratio and the body weight of emergent wasps correlated negatively with brood size. In the laboratory, single oviposition on P. brassicae larvae did not produce any brood size larger than 62, but brood size increased with superparasitism.
3. Laboratory experiments demonstrated that both naive and experienced female wasps were willing to attack hosts that had been newly parasitised by themselves or conspecifics. Superparasitism reduced survivorship but increased food consumption and weight growth in P. brassicae larvae. Superparasitism lengthened parasitoid development and prolonged the feeding period of host larvae.
4. Despite a trade-off between maximising brood size and optimising the fitness of individual offspring, two or three ovipositions on P. brassicae larvae resulted in a greater dry female mass than did a single oviposition on the host. Thus, superparasitism might be of adaptive significance under certain circumstances, especially when host density is low and unparasitised hosts are rare in a habitat.     

A comparative study of larval cannibalism in three species of ladybird

Abstract. 1. Cannibalism was studied under laboratory conditions in three species of Coccinellidae (Coleoptera): Cycloneda sanguinea , Olla v-nigrum , and Harmonia axyridis . Larval cannibalism varied among species, C. sanguinea  > H. axyridis  ≥  O. v-nigrum .
2. Larvae of all species cannibalised more in response to reduced food availability (14 h starvation daily) than in response to reduced food quality (dry Ephestia eggs).
3. Larvae of H. axyridis cannibalised siblings at lower rates than non-siblings, but larvae of C. sanguinea and O. v-nigrum did not. Rates of cannibalism increased significantly with increasing size disparity among larvae of all three species.
4. Cannibalism in C. sanguinea and H. axyridis , but not in O. v-nigrum , increased with larval density, suggesting that not all attacks on conspecifics are driven by hunger.
5. Costs, not benefits, were the predominant effects of exclusively and partially cannibalistic diets. A diet of conspecific eggs yielded survival equivalent to the Ephestia egg diet for H. axyridis and O. v-nigrum , but developmental time was extended and adults were smaller. Larvae of C. sanguinea had superior survival and faster development on conspecific eggs than on the Ephestia egg diet. Only H. axyridis larvae survived as well on diets comprised exclusively or partially of conspecific larvae as on Ephestia eggs, although they developed more slowly.     

Food competition among Scotophugu stercoraria Iarvae with emphasis on its effects on reproductive success

Abstract. 1. Laboratory experiments with yellow dung fly larvae, Scatophaga sfercoraria L. were carried out in the hope of clarifying the causal relationship underlying the extreme variation in body size in wild populations.
2. First, parents were collected in the field and newly hatched larvae exposed to different density levels under two temperature regimes (15°C and 20°C). Second, heritability experiments were done.
3. The results showed that density during the larval stage has a very significant effect on adult size. Male size is more affected than female size and the male larvae spend more time feeding than the female larvae with the result that the males are on average larger. Also, the additive genetic component is of little or no significance, supporting the notion that body size is positively related to fitness.
4. Finally, the results are discussed in relation to field data on the effects of size on both male and female reproductive success.     

Relationships between body size,reproductive traits,and food resource in three species of tropical flesh flies

Intra- and inter-specific comparisons in progeny size and clutch size were used to identify possible selection pressures acting on three sympatric species of ovoviviporous flesh flies. Both progeny and clutch sizes increased with body weight of the female parent in all species. There was no evidence for a simple trade-off between the two traits. In one species, the larger the female adult the smaller the relative biomass of larvae per clutch (total weight of larvae/body weight of female parent). Female flies produced similar progeny and clutch sizes over successive batches. Variation in progeny size did not influence either larval survival or duration of larval and pupal development. However, larger first instar larvae tended to form larger pupae in two of the species. The experiments testing the effect of progeny size on pupal weight under limited food conditions suggest a selective advantage for larger progeny size.     

Effects of suppressed oviposition activity and flight muscle histolysis on food consumption and ovarian development in a wing-dimorphic cricket: an explanation for sporadic conclusions related to physiological trade-offs

The effects of deprivation of oviposition substrate on food consumption and egg production were compared between the long-winged (LW) and the short-winged (SW) morph of a cricket, Modicogryllus confirmatus, to determine how suppressed oviposition activity would influence these traits in each wing morph. Food consumption was greatly suppressed in females deprived of oviposition substrate (-OS) compared to those given access to it (+OS) during the 2-week feeding trial in the SW morph but not in the LW morph. Some LW females shed their hindwings and histolyzed the flight muscles. Such de-alated LW (DLW) morphs tended to consume more food than intact LW (ILW) morphs. In all morphs, ovarian weight was heavier under -OS conditions than under +OS conditions during the second week of adulthood, although the differences were greater in SW morphs than in ILW morphs. In DLW morphs in which flight muscle histolysis was induced by artificial de-alation at adult emergence, the temporal changes in ovarian weight were similar to those of SW morphs.In SW morphs, food consumption was also significantly reduced when ovipositing females were deprived of oviposition substrate for 2 days compared to those allowed to oviposit continuously, but food consumption was not reduced in ILW or DLW morphs. SW females from which one ovary was extirpated at adult emergence, SW (-o), also showed a significant difference in food consumption when treated as above, indicating that food consumption was not determined simply by the number of ovarian eggs. The crop content was positively correlated to food consumption and smaller under -OS conditions than under +OS conditions. The 2-day deprivation of oviposition substrate caused no significant difference in the total number of deposited and ovarian eggs in any group, but the ovarian mass of developing oocytes tended to be smaller under -OS than under +OS conditions, particularly in SW morphs.These results indicate the possibility that some inconsistent results and conclusions discussed in recent studies, concerning the physiological trade-offs between flight capability and reproduction, were caused by the suppressed oviposition activity and failure to recognize the occurrence of flight muscle growth and histolysis in the test crickets.     

Adaptive plasticity of egg size in response to competition in the cowpea weevil,Callosobruchus maculatus (Coleoptera: Bruchidae)

Life history theory predicts that larger propagules should be produced when the offspring are expected to experience intense competition. This study tested whether female cowpea weevils responded to high larval or adult density by producing larger eggs. In a splitbrood design I measured the effect of density experienced by females at their larval stage (1 vs. 4–6 larvae/cowpea) on the size of eggs produced just after emergence. The females were then kept either at low adult density (1 female+1 male per vial), or at high adult density (10 females+10 males) for 2 days, and tested for the effect of this adult density treatment on the size of eggs laid subsequently. I measured egg length and width, as well as the diameter of the entrance tunnel made by the larva, which can be regarded as a crude measure of larval size. Females that experienced high adult density subsequently laid slightly wider eggs than those kept at low density. This difference, albeit small (about 1–4% after correction for female weight and the effect of family, depending on the statistical model used), was statistically significant and robust to alterations of the statistical model. It may be a remnant of a larger plastic response of egg size to competition that has become eroded during many generations in high-density laboratory cultures. There was no difference in egg length or the diameter of the entrance tunnel. Eggs laid just after emergence by females reared at high larval density also tended to be wider than those produced by females that had no competitors. This effect was only marginally significant, however, and sensitive to the statistical model. Both egg length and width and the diameter of the entrance tunnel increased with female weight and decreased with female age. The tunnel diameter was positively correlated with both egg length and width, but the effect of width was larger.     

Dispersal by bagworm larvae, Metisa plana: effects of population density, larval sex, and host plant attributes

Abstract 1. The work reported here investigated the incidence of dispersal by bagworm larvae Metisa plana (Walker) (Lepidoptera: Psychidae), a species with apterous females and winged males.
2. A mark–recapture study conducted in a plantation of oil palm revealed that a significant proportion of larvae undertakes multiple episodes of ballooning, and suggested that ballooning represents a strategy for foraging both within and between hosts.
3. Experiments conducted in a controlled cage environment indicated that increasing population density and defoliation of oil palm both promote dispersal by larvae.
4. Neonatal larvae typically remained on the host where they emerged, indicating that pupation site of late-instar females influences the distribution of their progeny, and suggesting a high level of genetic relatedness in localised bagworm populations.
5. Density-dependent dispersal may generate relatively uniform distributions of M. plana in plantations of oil palm, by simultaneously stabilising populations on heavily infested palms and redistributing larvae on lightly infested palms.
6. The rate of ballooning was greater for female than for male larvae, possibly resulting from the sex-specific impact of population density on reproductive success or from late-instar females seeking suitable oviposition sites for their future progeny.     

Runaway social games,genetic cycles driven by alternative male and female strategies,and the origin of morphs

Analysis of evolutionarily stable strategies (ESS) and decade-long field studies indicate that two color morphs of female side-blotched lizards exhibit density- and frequency-dependent strategies. Orange females are r-strategists: they lay large clutches of small progeny that are favored at low density. Conversely, yellow females are K-strategists: they lay small clutches of large progeny that are favored when carrying capacity is exceeded and the population crashes to low density. Interactions among three male morphs resembles a rock-paper-scissors (RPS) game. Fertilization success of males depends on frequency of neighboring morphs. Orange males usurp territory from blue neighbors and thereby mate with many females. However, orange males are vulnerable to cuckoldry by sneaky yellow males that mimic females. The yellow strategy is thwarted in turn by the mate-guarding strategy of blue. Sinervo and Lively (1996) developed a simple asexual model of the RPS game. Here, we model the dynamics of male and female morphs with one- and two-locus genetic models. Male and female games were considered in isolation and modeled as games that were genetically coupled by the same locus. Parameters for payoff matrices, which describe the force of frequency-dependent selection in ESS games, were estimated from free-ranging animals. Period of cycles in nature was 5 years for males and 2 years for females. Only the one locus model with three alleles (o, b, y) was capable of driving rapid cycles in male and female games. Furthermore, the o allele must be dominant to the y allele in females. Finally, the amplitude of male cycles was only reproduced in genetic models which allowed for irreversible plasticity of by genotypes, which is consistent with hormonally-induced changes that transform some males with yellow to dark blue. We also critique experimental designs that are necessary to detect density- and frequency-dependent selection in nature. Finally, runaway ESS games are discussed in the context of self-reinforcing genetic correlations that build and promote the formation of morphotypic variation.