Maternal behavior and clutch manipulation in the ring-legged earwig (Dermaptera: Carcinophoridae)

The expression and maintenance of maternal behavior in the earwig,Euborellia annulipes, was examined through manipulation of clutch size, age, and species and through observations of interactions between brooding females. Females underwent discrete gonadotrophic cycles culminating in oviposition of first clutches that were highly variable in size. Neither the head capsule width nor the age of the mother was correlated with clutch size. Maternal care extended through embryogenesis and for the week following hatching. Clutch removal significantly shortened the interclutch interval, indicating that the presence of brood inhibited the onset of the second gonadotrophic cycle. Brooding females readily accepted replacement clutches of the same age. Thus, mothers did not appear to distinguish their own eggs from those of other females. Experimental doubling of clutch size did not significantly reduce the proportion hatching or fledging. In contrast, reducing clutch size diminished the percentage successfully fledging. Manipulation of clutch age resulted in reduced hatching/fledging success. Placing two females, each with newly laid clutches, in the same cage usually resulted in egg transfer from the nest of one female to that of the other within 12 h. Nests of females with larger forceps were significantly more likely to contain both clutches. When mothers with first clutches were paired with mothers with third clutches, eggs were more likely to be transferred to the nest of the older female.E. annulipes females with newly laid clutches appeared to accept as replacement clutches eggs of the earwigDoru taeniatum. Alien clutches were maintained for the typical duration of embryogenesis; however, noD. taeniatum hatchlings were observed.     

A comparison of the reproductive strategies of three species of Dysdercus from Africa (Hemiptera,Pyrrhocoridae)

Summary Reproductive strategies have been investigated for three species of Dysdercus from Africa, D. intermedius, D. cardinalis and D. nigrofasciatus. Reproductive parameters were measured for each species on individuals cultured in the laboratory. D. cardinalis is associated more with herbaceous than with arboreal host plants and is intermediate in body size. It tended to have the least age at first reproduction, reproduced most frequently, and layed the smallest clutches on average, spreading relatively high amounts of parental investment in more clutches than the other two species. It also laid the greatest number of clutches and eggs in a lifetime. D. intermedius specializes on arboreal host plants, and is the largest of the three species. It had the greatest age at first reproduction, reproduced less frequently than D. cardinalis, and layed the largest clutches. Its parental investment over time declined earlier than that of D. cardinalis. D. intermedius layed the least number of clutches in a lifetime, but its lifetime gross reproduction was intermediate in magnitude. D. nigrofasciatus has the widest diet breadth, and is the smallest in body size. Its age at first reproduction was similar to that of D. cardinalis, but D. nigrofasciatus reproduced least frequently, laying relatively large clutches. It was similar to D. intermedius in its pattern of parental investment over time, and essentially layed the same number of clutches in a lifetime as the latter species, although its lifetime gross reproduction was the smallest.It is suggested that the reproductive strategies of the three species might have evolved as adaptations to selection events in the niches of the immature stages.     

Intra-specific variations of egg size,clutch size and larval survival relatedto maternal size in amphidromous <Emphasis Type="BoldItalic">Rhinogobius</Emphasis> goby

Synopsis Newly hatched amphidromous Rhinogobius sp. CB (cross band type) larvae drift downstream to the sea to grow and develop before returning upstream as juveniles. Since larger and older individuals usually inhabit the upper reaches of rivers, larvae from larger females are more likely to suffer higher risks of starvation or predation during their longer migration to the sea. We examined the relationship between reproductive parameters (egg volume and clutch size) and maternal length. We collected adult Rhinogobius sp. CB along the course of the Aizu River (Wakayama Prefecture, Japan) and spawned them under laboratory conditions. We measured egg volumes and clutch sizes, as well as larval starvation tolerance. Both egg volume and clutch size increased with standard length or age of the maternal fish, while egg density in these clusters did not correlate with standard length. Gonad-somatic index (GSI) also tended to increase with maternal standard length. There was a significant positive correlation between egg size and the 72-h survival rate of unfed hatchlings. Intra-specific variation of egg volumes and clutch sizes in this species seems to be an adaptation for enhancing offspring survivorship during migration to the sea. Some females spawned a second time. Second spawned egg sizes were smaller than first spawned egg sizes, although there was no difference in clutch size between the two. Egg size variation between subsequent spawns may be an adjustment to the changes in seasonal environmental conditions.     

MATERNAL INHERITANCE OF CONDITION AND CLUTCH SIZE IN THE COLLARED FLYCATCHER

Maternal effects may strongly influence evolutionary response to natural selection but they have been little studied in the wild. We use a novel combination of experimental and statistical methods to estimate maternal effects on condition and clutch size in the collared flycatcher, where we define “condition” to be the nongenetic component of clutch size. We found evidence of two maternal effects. The first (m) was the negative effect of mother's clutch size on daughter's condition, when mother's condition was held constant. The second (M) was the positive effect of mother's condition on daughter's condition, when mother clutch size was held constant. These two effects oppose one another because mothers in good condition also lay many eggs. The maternal effects were large: Experimentally adding an egg to a mother's nest reduced clutch sizes of her daughters by 1/4 egg (i.e., m = -0.25). Measured degree of resemblance between mother and daughter clutch sizes yielded M = 0.43. The results weakly support the presence of heritable genetic variation in clutch size: additive genetic variance/total phenotypic variance = 0.33. This estimate was highly variable probably because, as we show, mother-daughter resemblance may depend hardly at all on the amount of genetic variance when maternal effects are present. Daughter-mother regression (a standard method for estimating heritability) is consequently a poor guide to the amount of genetic variance in clutch size. Our results emphasize the value of combining field experiments with observations for studying inheritance.     

Clutch size in a leaf-mining fly (Pegomya nigritarsis: Anthomyiidae)

ABSTRACT.

• 1 Pegomya nigritarsis Zetterstedt is a gregarious leaf-mining fly attacking Rumex sp. The larvae from one clutch of eggs all inhabit the same mine.
• 2 Egg-batch size varies between host plants and is correlated with leaf size.
• 3 On Rumex obtusifolius L. larval survival as a function of clutch size was measured in the field. This measure was incorporated in an optimal clutch size model and a clutch size prediction obtained.
• 4 The predicted optimal clutch size, three, is that most frequently found in the field.

     

A comparative study of oocyte development in cotton stainers (Dysdercus spp., pyrrhocoridae) and the factors that control egg production

The relative importance of three factors-mating, nutrition, and crowding-in regulating the reproductive biology of Dysdercus fasciatus Signoret, D. nigrofasciatus Stål and D. cardinalis Gerst. was studied under standard laboratory conditions. Observations on mated and virgin females of these three species of Dysdercus showed that there were no significant differences in pre-oviposition period and inter-oviposition period. In D. fasciatus and D. nigrofasciatus there was, further, no significant difference in the rate of oocyte development; but D. cardinalis differed from these two species in that mating had a profound effect on the rate of oocyte development and consequently on the first oviposition. However, in all three species of Dysdercus mating had a significant effect on the number of eggs produced in each batch, the egg batches of mated females being significantly larger than those produced by virgin females. Further, in both D. fasciatus and D. nigrofasciatus virgin females lived longer than mated females but in D. nigrofasciatus mating had no significant effect on adult longevity. Mating activity was drastically reduced in starved adult bugs. But where it occurred under these conditions, mating did not have any effect on the development of the oocytes, even in D. cardinalis. Under starvation conditions, growth proceeded up to the stage where yolk deposition would normally occur, then stopped; the oocytes showed no signs of degeneration or oosorption. Crowding of female adults led to the acceleration of oocyte development and the onset of oviposition in all three species. These findings are discussed in relation to the known field biology of the three Dysdercus species.

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Résumé La biologie de la reproduction de 3 espèces de Dysdercus du Kenya a été étudiée au laboratoire. Des 3 espèces, D. fasciatus Signoret présente les périodes les plus courtes de pré-copulation et de pré-oviposition. Puisque cette espèce est inféodée à un relativement petit nombre de plantes-hôtes, qui ont une courte saison de fructification procurant alors une ample source de nourriture, elle tire un avantage maximum de la brièveté de cette période favorable, grâce à la réduction des phases de pré-copulation et de pré-oviposition.Par contre D. nigrofasciatus Stål est adaptée à un très grand nombre de plantes hôtes et pour elle il n'est pas indispensable d'avoir une très grande capacité de reproduction. Cela s'applique encore mieux à D. cardinalis Gerst. qui a une plus longue période de pré-copulation et de pré-oviposition. Cette espèce préfère les plantes qui sont dispersées sur une grande superficie et cela exige une longue période de vol exploratoire avant de s'alimenter et de se reproduire.Chacune de ces 3 espèces s'accouple de façon presque continue, mais les observations faites au laboratoire montrent que l'accouplement n'a aucun effet notable sur la longévité des adultes, sur la durée des phases de pré-oviposition ou d'inter-oviposition et sur le nombre total d'ufs pondus chez D. fasciatus et D. nigrofasciatus. Chez D. cardinalis, au contraire, l'accouplement a un effet important sur la vitesse de croissance des ovocytes, mais non sur les autres aspects de la reproduction évoqués ci-dessus.Le rassemblement en masse des femelles adultes accélère le développement des ovocytes et déclenche la ponte dans chacune des 3 espèces de Dysdercus.L'alimentation a un effet sur le développement des ovocytes. De jeunes femelles adultes doivent être alimentées pendant au moins 3–4 jours avant de déposer leur première ponte. Après chaque ponte une alimentation plus abondante est encore nécessaire pour assurer le maintien de la production d'ufs. Les femelles de chacune des 3 espèces peuvent survivre avec de l'eau pure mais leurs ovocytes arrêtent leur développement au stade où normalement se dépose le vitellus, mais ils ne sont pas résorbés.

     

Responses of Primula vulgaris to light quality in the maternal and germination environments

• In the model species Arabidopsis thaliana phytochromes mediate dormancy and germination responses to seasonal cues experienced during seed maturation on the maternal plants. However, the effect of the maternal light environment on seed germination in native wild species has not been well studied. This is particularly important given its practical application in the context of environmental restoration, when there can be marked changes in the canopy.
• Plants of Primula vulgaris were grown in the field over two vegetative seasons under four shading treatments from low to high ratio of red to far‐red light (R:FR). Leaf and seed traits were assessed in response to the light treatments. The germination of seeds from these four maternal environments (pre‐dispersal) was investigated at seven light and five temperature treatments (post‐dispersal).
• Thinner leaves, larger leaf area and greater chlorophyll content were found in plants growing in reduced R:FR. Shading in the maternal environment led to increased seed size and yield, although the conditions experienced by the maternal plants had no effect on seed germination. Seeds responded strongly to the cues experienced in their immediate germination environment. Germination was always enhanced under higher R:FR conditions.
• The observed phenotypic trait variation plays a major role in the ability of P. vulgaris to grow in a wide range of light conditions. However, the increased germination capacity in response to a higher R:FR for all maternal environments suggests potential for seedling establishment under vegetative shade only in the presence of canopy gaps.

     

Crowding‐induced changes in growth,reproduction and morphology of Daphnia

• 1 Daphnia may reach high population densities seasonally, or in patches, in lakes. To test the effects of chemicals released by high daphniid densities on their life‐history traits, nine species of Daphnia, D. magna, D. pulicaria, D. pulex, D. hyalina, D. galeata, D. laevis, D. lumholtzi, D. ambigua and D. cucullata, were grown in water from crowded Daphnia cultures in a flow‐through system in the presence of abundant food.
• 2 Water from Daphnia at 85 L^‐1 depressed growth rate, and lowered body size and clutch at first reproduction of six species of small‐bodied Daphnia (adult body length < 1.8 mm), but had no significant effects on larger species. Two clones of D. pulex differed in their growth rate in response to crowding, indicating that response patterns may vary within species.
• 3 Chemicals released by crowded D. magna reduced tail spine length in D. lumholtzi and D. cucullata by 37% and 11%, respectively, and induced changes in carapace morphology in D. lumholtzi and D. ambigua.
• 4 Chemicals released by crowded conspecifics may provide an additional, density‐dependent mechanism of population regulation; when large species of Daphnia coexist at a high population density with small species, these chemicals may reinforce the competitive advantage of large species.

     

Transgenerational plasticity in Silene vulgaris in response to three types of stress

• The environment experienced by plants can influence the phenotype of their offspring. Such transgenerational plasticity can be adaptive when it results in higher fitness of the offspring under conditions correlated with those experienced by the mother plant. However, it has rarely been tested if such anticipatory parental effects may be induced with different environments.
• We grew clonal replicates of Silene vulgaris under control conditions and three types of stress (nutrient deficiency, copper addition and drought), which are known from natural populations of the species. We then subjected offspring from differently treated mother plants to each of the different stress treatments to analyse the influence of maternal and offspring environment on performance and several functional traits.
• Current stress treatments strongly influenced biomass and functional traits of the plants, mostly in line with responses predicted by the theory of functional equilibrium. Plant performance was also influenced by maternal stress treatments, and some effects independent of initial size differences remained until harvest. In particular, stressed mothers produced offspring of higher fitness than control plants. However, there was no evidence for treatment‐specific adaptive transgenerational plasticity, as offspring from a mother plant that had grown in a specific environment did not grow better in that environment than other plants.
• Our results indicate that the maternal environment may affect offspring traits and performance, but also that this transgenerational plasticity is not necessarily adaptive.

     

Direct and transgenerational effects of an experimental heatwave on early life stages in a freshwater snail

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Chemical egg defence in the large milkweed bug,Oncopeltus fasciatus,derives from maternal but not paternal diet

Many herbivorous insects sequester defensive compounds from their host‐plants and incorporate them into their eggs to protect them against predation. Here, we investigate whether transmission of cardenolides from the host‐diet to the eggs is maternal, paternal, or biparental in the large milkweed bug, Oncopeltus fasciatus (Dallas) (Hemiptera: Lygaeidae). We reared individual bugs on either milkweed seeds [MW; Asclepias syriaca L. (Apocynaceae)] that contain cardenolides, or on sunflower seeds [SF; Helianthus annuus L. (Asteraceae)] that do not contain cardenolides. We mated females and males so that all four maternal/paternal diet combinations were represented: MW/MW, MW/SF, SF/MW, and SF/SF. Using larvae of the common green lacewing, Chrysoperla (Chrysopa) carnea (Stevens) (Neuroptera: Chrysopidae), we conducted two‐choice predation trials to assess whether maternal, paternal, or biparental transmission of cardenolides into the eggs of O. fasciatus increased protection against predation. Furthermore, we used high performance liquid chromatography (HPLC) to assess putative cardenolide content of eggs from the various parental diet treatment groups. The predation trials suggested that regardless of male diet, eggs were afforded better protection when females had been raised on milkweed. However, many eggs were at least partially consumed. This suggests that although chemical defence of eggs does not guarantee protection to eggs on an individual basis, they may increase the probability that some eggs in a clutch are left intact thereby potentially conferring a fitness advantage to more offspring than if eggs are left unprotected. Based on HPLC analysis we found that maternal contribution of cardenolides was significantly greater than paternal contribution of cardenolides to the eggs, supporting the results of our predation trials that a maternal diet of milkweed makes eggs more distasteful than a paternal diet of milkweed.     

Population characteristics and life cycle of the Lake Baikal invader <Emphasis Type="Italic">Gmelinoides fasciatus</Emphasis> (Stebbing, 1899) (Crustacea: Amphipoda in Lake Ladoga

Dynamics of the size-age structure and sexual structure of the population and characteristics of reproduction and life cycle have been studied in the Baikal invader Gmelinoiudes fasciatus (Stebbing, 1899) in Lake Ladoga. The studied characteristics display both similar features with and differences from populations of this species in other waterbodies. It is found that the size and age structure of the G. fasciatus population changes in the circannian aspect under the influence of complex of environmental factors. The sexual structure of the population is dynamic, but a sex ratio of approximately 1: 1 is maintained in the circannian cycle. Six principal periods of hatching are recognized. The main factor determining the timing of the start and termination of the reproduction period in G. fasciatus is temperature. The weight of eggs oviposited by the female over the whole life cycle (about 21.5 cal in the energy equivalent) amounts to about 60% of the female body weight.     

Maternal Effects of Photoperiod and Food Level on Life History Characteristics of the Cladoceran Daphnia Pulicaria Forbes

The response of various life-history characteristics of Daphnia pulicaria to photoperiod and food concentration was measured in 16 combinations of maternal and offspring environments (long vs. short day, high vs. low food) in flow-through experiments. Response variables in offspring were time and survival to release of first offspring, clutch size and neonate mass in the first brood, mass of adult females after 30days and somatic growth rate during the course of the experiment. Most of these parameters were directly controlled by food concentration in the offspring environment, but maternal effects frequently modified the response. A long day length in the maternal environment resulted in a prolongation of the time to first clutch release in offspring similar to the direct effect of low food. Likewise, survival to maturation and female mass were affected by maternal photoperiod. Somatic growth rate and clutch size responded to combined effects of maternal food conditions and photoperiod. The laboratory results were used to predict the seasonal change of fecundity of Daphnia in the field. When data on clutch size are ordered in a sequence as the different combinations of maternal and offspring environment occur during the seasonal succession in a temperate lake, they show a bimodal distribution with a high peak in spring and a lower peak in fall. This pattern is consistent with field observations. We conclude that photoperiod and maternal effects are important factors influencing life history and population dynamics of Daphnia.     

Divergent phenotypic responses to predators and cyanobacteria in Daphnia lumholtzi

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An energetic analysis of host plant selection by the large milkweed bug,Oncopeltus fasciatus

Summary The large milkweed bug, Oncopeltus fasciatus, is a specialized seed feeder that has been observed completing nymphal development in the field on only a small proportion of its potential host species within the genus Asclepias. In central Missouri only two of the six milkweed species studied, A. syriaca and A. verticillata, commonly supported nymphal O. fasciatus growth in the field. The seed of all six species, however, was equally suitable food for bugs reared in the laboratory. In laboratory preference tests, adult bugs chose to feed on the largest seeds, A. hirtella, but such a preference could not explain the observed field feeding patterns.One explanation to account for the observed host plant selection is based upon an energetic analysis. Only A. syriaca provided enough seed biomass for a clutch of O. fasciatus nymphs to develop on a single plant, and only A. verticillata grew in high enough density that a clutch could find sufficient food within the limited range of nymphal movement. These results illustrate a corollary of the resource concentration hypothesis: within a plant group whose members share similar secondary plant chemistries, the only species that will be viable hosts for a specialized herbivore are those that provide the minimal resource density necessary for the completion of nymphal development.In central Missouri, O. fasciatus has specialized on a critical resource density, not traits of individual Asclepias species. The appearance of host selection within the potential host plant spectrum is the result of a characteristic growth form, seed output, and dispersion pattern for each milkweed species that makes some species much more likely than others to produce sufficient seed resources.     

An investigation of allelopathic effects of Daphnia

• 1 The hypothesis is tested that large daphnids are able to suppress their own and other species' feeding and reproduction by means of excreting an inhibitory chemical (or chemicals).
• 2 In laboratory experiments with an Australian species, Daphnia carinata, water preconditioned with 3–67 daphnids 1-^?1 for 30 h had the effect of reducing feeding rates of D. carinata and D. lumholtzi provided with Selenastrum capricornutum.
• 3 For the two Daphnia species, there were highly significant negative correlations between feeding rate and the preconditioning density of D. carinata.
• 4 Water preconditioned with 20–30 daphnids 1-^?1 for 1–2 weeks reduced the grazing rates of Daphnia, Moina, and Diaphanosoma 2–3-fold.
• 5 Moina kept in such water for 2 days stopped feeding. Conditioned water kept for 3 days without animals still inhibited grazing by Moina. Hearing to 100°C removed the inhibitory effect.
• 6 Given excess food, and in non-renewed water, a gradient of D. carinata densities developed a strong negative correlation between clutch size and daphnid density after a 6-day time lag. This result may help explain the direct density-dependent regulation of cladoceran reproduction observed earlier in a subtropical lake.

     

Life-history variation in a hybrid species complex ofDaphnia

Summary Life-history variation was examined among members of theDaphnia longispina group, which consists ofD. galeata, D. hyalina, D. cucullata, and hybrids. Factorial experiments were conducted at two temperatures (14° and 20° C) and two food concentrations (0.2 and 1.0 mg Cl⁻¹). Differences in life-history features (size at maturity, age at first reproduction, size of first clutch, offspring size in first clutch) under the different environmental conditions were assessed among eightDaphnia clones, which represented members of this species complex. Significant differences between parentals and hybrids for most life-history features were observed under various treatments; generally, hybrid clones showed intermediate life-history traits when compared with parentals. When comparisons were made among clones within a given species (i.e.D. galeata, D. galeata xcucullata, D. cucullata), clonal differences were also noted for certain life-history traits. The data are discussed with reference to the formation and maintenance of hybrid species complexes in nature. This paper is dedicated to the memory of Hans Georg Wolf, who died suddenly in May 1990 at the age of 39 years.     

Life history patterns of parental and hybrid Daphnia differ between lakes

1. We investigated whether Daphnia galeata × hyalina hybrids of Lake Constance and Lake Greifensee show the same pattern of life history parameters as previously reported for D. galeata × cucullata hybrids and whether such a pattern is consistent between Daphnia populations from those two lakes. 2. Hybrids in Lake Constance were intermediate in size compared with the parental species. Hybrids in Lake Greifensee were smaller than D. galeata. The intrinsic growth rate (r) of hybrids from Lake Constance was not significantly different from the faster growing parental taxon D. galeata. However, r of hybrids from Lake Greifensee was significantly lower than that of D. galeata. 3. The observed juvenile body length differences between the taxa varied with the clutch number. The first clutch juvenile lengths of the three taxa did not differ for Lake Constance. First clutch juveniles of Lake Greifensee D. galeata were smaller than hybrid first clutch juveniles. The third clutch juvenile length did not differ between taxa from Lake Greifensee, but D. galeata juveniles from Lake Constance were bigger than those of D. hyalina. 4. The life history pattern found in Lake Constance corresponds to previous findings from other studies. The hybrids in this lake combine the faster population growth of one parental species with a relatively small size. In the case of Lake Greifensee hybrids, the relatively large size of first clutch juveniles and the small size of the adults could be interpreted as dual adaptations to invertebrate and fish predation. We speculate that the lower population growth rate of the hybrids is a trade‐off for this twofold protection.     

Maternal consumption of non‐toxic Microcystis by Daphnia magna induces tolerance to toxic Microcystis in offspring

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Effects of crowding and different food levels on growth and reproductive investment of Daphnia

The effects of daphniid crowding on juvenile growth rate, length at first reproduction, clutch size and egg size of four species of Daphnia were compared with the effects of food level. Juvenile Daphnia were grown to primipary in a flow-through system in water conditioned by different densities of the same, or another, species. At high ambient food levels, water from Daphnia that had been crowded at densities 150 l^–1 depressed growth rate and lowered body size and clutch size of D. hyalina and D. galeata; effects on the same traits of D. magna and D. pulicaria were variable (stimulation, depression, or no effect). D. hyalina and D. galeata responded to gradients of increasing daphniid density (0–300 l^–1) by altering egg mass, somatic mass and clutch size to maintain a relatively constant reproductive investment; egg mass increased with crowding and then decreased in a pattern consistent with Glazier's (1992) hypothetical model of changes in offspring size in relation to food quantity and maternal demand. Effects of crowding by conspecifics were indistinguishable from those of other species. This study, which uncouples the effect of crowding per se from ambient resource depletion, shows that chemical substances released by high densities of Daphnia can cause changes in life-history traits comparable to those that occur in response to low food levels.