Oviposition date and pattern of embryogenesis in the grasshopper Chorthippus brunneus (Orthoptera, Acrididae)

The morphological development of eggs of the grasshopper Chorthippus brunneus , laid during successive periods through a single season, was closely monitored in the field at a site in NW England. Eggs within the same clutch developed synchronously. However, there were significant differences between clutches laid at the same time, reflecting spatial variation in microclimate (and possibly maternal effects and/or genetic differences). Eggs laid early in the season entered diapause and hatched relatively synchronously in the following spring. Eggs laid later failed to reach the diapause-stage before winter, but hatched, albeit late and with greater variance. Diapause was averted in these eggs. Plasticity in the nature of diapause has been documented from laboratory studies of many insect species (including C. brunneus ,) but this study directly relates the incidence of such developmental plasticity to oviposition date and seasonal temperature cycles in the field. The adaptive significance of diapause aversion and the evolutionary consequences for female reproductive behaviour are considered.     

中国石龙子雌体繁殖特征和卵孵化的地理变异

浙江丽水和广东韶关中国石龙子均年产单窝卵,窝卵数,窝卵重和卵重均与雌体SVL呈正相关,雌体头部形态,繁殖特征,产卵起始时间和孵孵化的热依赖性等有显著的地理变异;韶关石龙子产卵起始时间为5月中旬,比丽水经子约早两周,韶关石龙子窝卵数较大,卵较小,窝卵重与丽水石龙子无显著差异。韶关石龙子特定SVL的窝卵数比丽水石龙子多2．8枚卵,中国经子卵数量和大小之间有种群间权衡,无种数内权衡,同一种群内卵数量与卵大小无关,孵化温度影响石龙子孵出幼体的一些特征,24℃孵出细幼体比32℃孵出幼体大,躯干发育好,剩余卵黄少,韶关24℃孵出幼体的体重,躯干干重小于丽水幼体,韶关32℃孵出幼体的SVL小于丽水幼体,剩余卵黄大于丽水幼体,表明适宜卵孵化温度范围有地理变异。丽水石龙子卵对极端高温和低温的耐受性较强,适宜卵孵化温度范围较宽。     

Maternal effects on progeny body size and color in the desert locust, Schistocerca gregaria: examination of a current view

Hatchling body color and size of the desert locust, Schistocerca gregaria, are determined by the population density of the mothers during their reproductive period. Smaller green hatchlings are produced by adults at low population density (solitarious conditions) and larger dark hatchlings at high population density (gregarious conditions). One claim states that a pheromonal factor secreted by gregarious mothers into foam plugs of egg pods induces darkening in hatchlings. Previous research suggests that the foam factor can be removed by separating eggs individually within 1h of deposition, causing presumptive gregarious eggs to hatch without darkening. The present study re-examined this claim and possible factors that have been proposed which could account for the difference between our results and those reported earlier. Early separation was performed on eggs with a low mortality rate. The results showed that the egg separation did not increase the incidence of green hatchlings. Once chorionated in the ovary, eggs remained unchanged in size until the second day after oviposition in either isolated or crowded locusts. This and other results suggest that the phase-dependent differences in body size and color of hatchlings are established in the ovary and that modifications by the accessory gland factor either in the oviduct or after deposition are unlikely.     

Changes in egg colour, egg weight and oviposition rate with the number of eggs laid by wild females of the small heath butterfly, Coenonympha pamphilus

ABSTRACT. 1. The colour of eggs laid by individual Coenonympha pamphilus (L.) (Lep., Satyridae) females changed over their lifespan. The first laid eggs were green, but after having laid about 100 eggs, females laid only yellow eggs.
2. By following females in the field, or by capturing wild females, and noting the colour of their eggs, it was established that younger females showed higher oviposition rates and laid heavier eggs than older females.
3. Hence, wild C.pamphilus females are unable to sustain a constant egg production by extracting amino acids or any other nutrients from nectar. We hypothesize that this is a general phenomenon in nectar feeding butterflies.
4. The egg colour did not seem to match the colour of the substrate on which the female chose to deposit the egg.     

Changes in the behavior of the female short-tailed cricket,Anurogryllus muticus (De Geer) (Orthoptera: Gryllidae) following mating

InAnurogryllus muticus females, mating stimulates burrow construction, burrow provisioning, feeding, egg production, and egg-laying. Since mating often occurs before the ovaries are fully developed, the time span between mating and oviposition is used for increased food intake and the accumulation of nutrient reserves in the fatbody. Oviposition triggers maternal care of eggs and emerging hatchlings, and blocks egg consumption. Hatchling behavior is investigated. When hatchlings eat eggs, they prefer newly laid over older, embryonic eggs.     

Seasonal declines in incubation periods of Brünnich's Guillemots Una lomvia:testing proximate causes

In many species of bird, eggs laid late in the laying period hatch after a shorter incubation period than do eggs laid early. These seasonal declines in incubation period are generally thought to confer evolutionary advantages, but the proximate mechanisms that underlie them are poorly understood. Seasonal declines in incubation period are usually attributed to: (1) seasonal increases in ambient air temperatures; (2) seasonal changes in the behaviour of incubating birds; and/or (3) seasonal declines in egg size. In a previous study, Common Guillemot Una aalge incubation periods declined with laying date at a low-Arctic colony. As there was no support for hypotheses 1 or 2, it was suggested that this occurred because egg size declined with laying date, but eggs were not measured in that study. We recorded similar seasonal declines in the incubation periods of the single eggs laid by Brünnich's Guillemots Una lomvia at two low-Arctic colonies in four years. Neither seasonal variation in ambient air temperatures, nor in the behaviour of incubating adults, appeared to cause the declines. As predicted for Common Guillemots, incubation period increased with egg size among Briinnich's Guillemots, in one of two years. However, incubation period declined with laying date in the absence of corresponding declines in egg size. We conclude that none of the three commonly proposed proximate mechanisms adequately explains the seasonal variation in guillemot incubation periods. Several testable, alternative mechanisms are explored.     

Ecological correlates and heritability of reproductive variation in two co-occurring grasshopper species

ABSTRACT. 1. Relationships are examined between mean egg size and possible selection pressures acting on two grasshopper species, Chorthippus brunneus (Thunb.) and Myrmeleotettix maculatus (Thunb.), co-occurring at three sites in a sand dunes area.
2. Heavy eggs were laid where the mean crowding of hatchlings was high in relation to the amount of grasses with thin-edged leaves. To a lesser extent, large eggs were also found in more desiccating conditions. Mean egg dry weights did not correlate with measures of crowding or measures of crowding in relation to total grass abundance.
3. Significant differences in egg size and clutch sizes were observed between the stocks of second generation, laboratory-reared M.maculatus from three study sites. The significant difference in egg size (site 2 stock > site 1 stock) ran counter to the difference observed in the field but the difference in clutch size (site 3< site 2) was consistent with that observed between field populations.
4. We argue that egg size is primarily determined by selection pressures on the size of hatchlings but that predictably poor feeding conditions for juveniles and adults at site 2 results in the production of smaller adults which, in M.maculatus , then lay smaller eggs. We further argue that M. maculatus from site 2 have been selected to lay larger eggs for their body size and body condition in order to compensate partially for this environmental constraint.     

Performance of a spring-feeding moth in relation to time of oviposition and bud-burst phenology of different host species

Abstract. 1. As a spring-feeding moth committed to immature foliage, the autumnal moth Epirrita autumnata (Lepidoptera, Geometridae) must have egg hatch synchronised with the bud-burst of its host plants. Due to large individual variation in the length of the pupal period, however, E. autumnata populations exhibit a prolonged period of flight and oviposition. Because the timing of oviposition in autumn is associated with the timing of egg hatch in the following spring, the time window for egg hatch expands and more potential hosts may become attainable. This suggestion was evaluated under field conditions by rearing E. autumnata eggs and larvae on four different hosts.
2. The performance of E. autumnata was measured by using estimates for fecundity (pupal mass) as well as survivorship of eggs and larvae. Based on the availability of foliage and phenological synchrony between larval and leaf development, early-laid eggs and the larvae originating from them were predicted to perform better on the hosts that have early-flushing leaves. On the late-flushing hosts, the larvae that hatched later were predicted to perform better than the larvae that hatched earlier. Half of the trials were exposed to predators and parasitoids, while the rest were conducted inside mesh-bags preventing larval dispersal and mortality due to natural enemies.
3. The results of the experiment did not support the simple predictions. In particular, host-plant quality and natural enemies appeared to operate discordantly between early- and late-laid eggs. Larvae from the late-laid eggs had rapid development during the larval stages and pupated at the same time and with the same pupal mass as the larvae hatched from the early-laid eggs.
4. The results indicate an occurrence of several, unknown selective forces in E. autumnata populations maintaining variation in the length of the pupal period, timing of oviposition, and timing of egg hatch.     

Variation in trichome density and resistance against a specialist insect herbivore in natural populations of Arabidopsis thaliana

Abstract.  1. The annual herb Arabidopsis thaliana is a prime model organism of plant molecular genetics, and is currently used to explore the molecular basis of resistance to herbivores. However, both the magnitude and the causes of variation in resistance among natural populations of A. thaliana are poorly known. The hypotheses (a) that resistance to a specialist herbivore, the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), is positively related to the density of leaf trichomes, and (b) that oviposition preference by female moths is positively correlated to larval performance on different populations and maternal lines of the host plant were examined.
2. Variation in leaf trichome density and resistance to P. xylostella within and among six natural populations of A. thaliana in Sweden was quantified . Resistance was quantified by examining the number of eggs laid on plants exposed to ovipositing female moths and by monitoring larval development on plants of different origin.
3. Trichome density varied significantly among populations; for 4-week-old plants (exposed to ovipositing moths), it also varied significantly among maternal families within populations. The rate of oviposition varied significantly both among populations and among families within populations. This variation could partly be explained by a negative relationship between trichome density and egg number, and a positive relationship between plant size and egg number. Time to pupation, pupal mass, and time to adult eclosion did not vary among populations or maternal lines of the host plant, and offspring performance was not related to P. xylostella oviposition preference . The results indicate that A. thaliana populations may respond to selection for increased resistance to P. xylostella , and suggest that trichome production contributes to resistance against this specialist herbivore.     

CLUTCH SIZE, EGG SIZE, HATCH WEIGHT AND LAYING DATE IN RELATION TO EARLY MORTALITY IN RED GROUSE LAGOPUS LAGOPUS SCOTICUS CHICKS

Clutches of Red Grouse eggs were collected from the wild and subsequent hatching and rearing done in standard conditions in captivity. Variations in chick survival from one clutch to another in the same year were related to differences in hatch weight. Hatch weight was determined only partly by egg size. Weight loss between laying and hatching was related to survival independently of egg size. Variation in this weight loss obscured any simple relationship between egg size and survival, except in eggs laid by captive hens. Intrinsic differences amongst hens caused some variations in laying date, egg size, hatch weight and chick survival. Variations in egg size and hatch weight accounted for less than half the variation in survival; other unmeasured intrinsic factors were also important. Big clutches hatched earlier than small ones. The commonest clutches were of seven and eight eggs, with six and nine frequent. Very big clutches of ten or more eggs were infrequent and chicks from them sometimes survived worse than from smaller clutches. As in other species, the commonest clutch sizes were not the most productive. There was no simple relationship between egg size and clutch size.     

Context-Dependent Plastic Response during Egg-Laying in a Widespread Newt Species

Previous research on predator-induced phenotypic plasticity mostly focused on responses in morphology, developmental time and/or behaviour during early life stages, but the potential significance of anticipatory parental responses has been investigated less often. In this study I examined behavioural and maternal responses of gravid female smooth newts, Lissotriton vulgaris, in the presence of chemical cues originating from invertebrate predators, Acilius sulcatus water beetles and Aeshna cyanea dragonfly larvae. More specifically, I tested the extent of oviposition preference, plasticity in egg-wrapping behaviour and plasticity in egg size when females had the possibility to lay eggs at oviposition sites with and without predator cues during overnight trials. I found that individuals did not avoid laying eggs in the environment with predator cues; however, individuals that deposited eggs into both environments adjusted the size of the laid eggs to the perceived environment. Females deposited larger eggs earlier in the season but egg size decreased with time in the absence of predator cues, whereas individuals laid eggs of average size throughout the investigated reproductive period when such cues were present. Also, egg size was found to be positively related to hatching success. Individuals did not adjust their wrapping behaviour to the presence of predator cues, but females differed in the extent of egg-wrapping between ponds. Females’ body mass and tail depth were also different between ponds, whereas their body size was positively associated with egg size. According to these results, female smooth newts have the potential to exhibit activational plasticity and invest differently into eggs depending on temporal and environmental factors. Such an anticipatory response may contribute to the success of this caudate species under a wide range of predator regimes at its natural breeding habitats.     

The influence of maternal thermal environments on reproductive traits and hatchling traits in a Lacertid lizard,Takydromus septentrionalis

The thermal environment can induce substantial variation in important life-history traits. Experimental manipulation of the thermal environment can help researchers determine the contribution of this factor to phenotypic variation in life-history traits. During the reproductive season, we kept female northern grass lizards, Takydromus septentrionalis (Lacertidae), in three temperature-controlled rooms (25, 28 and 32 °C) to measure the effect of the maternal thermal environment on reproductive traits. Maternal thermal environment remarkably affected reproductive frequency and thereby seasonal reproductive output, but had little effect on reproductive traits per clutch or hatchling traits. Females kept at 32 °C produced more clutches and thus had shorter clutch intervals than females from 28 to 25 °C. Clutch size, clutch mass, relative clutch mass, egg size and hatchling traits did not vary among the three treatments. The eggs produced by the females were incubated at 27 °C and the traits of hatchlings were measured. The result that egg (offspring) size was independent of maternal thermal environments is consistent with the prediction of the optimal egg size (offspring) theory. The eggs produced by low temperature females (28 and 25 °C) took longer time to complete their post-oviposition development than did eggs produced by high temperature females (32 °C). This suggests that the eggs from low temperatures might have been laid when the embryos were at relatively early stages. Therefore, maternal thermal environment prior to oviposition could affect post-oviposition development in T. septentrionalis.     

Trade-off in oviposition strategy: choosing poor quality host plants reduces mortality from natural enemies for a salt marsh planthopper

Abstract.  1. Both host plant nutrition and mortality from natural enemies have been predicted to significantly impact host plant selection and oviposition behaviour of phytophagous insects. It is unclear, however, if oviposition decisions maximise fitness.
2. This study examined whether the salt marsh planthopper Pissonotus quadripustulatus prefers higher quality host plants for oviposition, and if oviposition decisions are made so as to minimise mortality at the egg stage.
3. A controlled laboratory experiment and 4 years of field data were used to assess the rates of planthopper oviposition on higher quality 'green' and lower quality 'woody' stems of the host plant Borrichia frutescens . The numbers and percentages of healthy eggs and eggs that were killed by parasitoids or the host plant were recorded.
4. In all years, including the laboratory experiment, Pissonotus planthoppers laid more eggs on lower quality woody stems than on higher quality green stems. While host plant related egg mortality was higher in woody stems, the percentage of eggs parasitised was much greater in green stems. This resulted in a lower total mortality of eggs on woody stems.
5. The results of this study demonstrate that, although Pissonotus prefers lower quality host plants for oviposition, this actually increases fitness. These data seem to support the enemy free space hypothesis, and suggest that for phytophagous insects that experience the majority of mortality in the egg stage, oviposition choices may be made such that mortality is minimised.     

The seasonal timing of oviposition in sand lizards (Lacerta agilis): why early clutches are better

We studied a population of sand lizards (Lacerta agilis) near the northern edge of the species' range in coastal Sweden. We captured, marked, released and recaptured 98 adult female lizards over 5 years. Hatchlings from 146 laboratory-incubated clutches (1279 eggs) from field-caught gravid females were measured, weighed, marked and released at the study site. Female sand lizards usually laid only a single clutch of 4 to 15 eggs each year, but varied considerably in the time of year at which they laid their eggs. Oviposition dates shifted between years depending on weather (basking opportunities), but the relative timing of oviposition was consistent within a given female from year-to-year. The first females to oviposit each year were large animals in good physical condition, that had grown rapidly in previous years. “Early” clutches were larger than “later” clutches, had higher hatching success, and tended to have higher post-hatching survival rates. Offspring from early clutches were larger than “later” hatchlings, and differed in body proportions (probably because seasonal changes in maternal temperatures directly modified offspring phenotypes). Overall, our study documents several strong correlates of the timing of oviposition, and suggests that variation in this trait among females has strong fitness consequences, perhaps related to maternal “quality”. The correlations we observed between oviposition date and other traits that have been invoked as determinants of hatchling survival in reptiles (e.g., hatchling size, body shape, opportunities for multiple mating by the mother) suggest that hypotheses advocating simple causal connections between these traits and hatchling success should be viewed with caution.     

Effects of time-dependent intraspecific competition on offspring survival in the butterfly, Anthocharis scolymus (L.) (Lepidoptera: Pieridae)

Effects of mothers' eclosion and oviposition timing on the survival of their offspring in the pierid butterfly Anthocharis scolymus (L.) was examined. I recorded the performance of individual eggs and larvae that differed in their mother's eclosion and oviposition timing in a natural population, where A. scolymus feeds on Turritis glabra (L.) Bernh. Eggs laid early in the season, and larvae emerging from these eggs, had higher survival than eggs laid later, and larvae emerging from eggs that were laid later in the season, the causal factor being egg cannibalism by larvae on the same host plant. Logistic regression showed that females eclosing early in the season had higher offspring survival than females eclosing later. I conclude that optimal timing of adult eclosion in A. scolymus is a trade-off between eclosing early with associated higher offspring survival but lower egg-laying rate, and eclosing later with associated lower offspring survival but higher egg-laying rate. Received: 4 August 1997 / Accepted: 19 October 1997     

Use of statistical models based on radiographic measurements to predict oviposition date and clutch size in rock iguanas (Cyclura nubila)

The ability to noninvasively estimate clutch size and predict oviposition date in reptiles can be useful not only to veterinary clinicians but also to managers of captive collections and field researchers. Measurements of egg size and shape, as well as position of the clutch within the coelomic cavity, were taken from diagnostic radiographs of 20 female Cuban rock iguanas, Cyclura nubila, 81 to 18 days prior to laying. Combined with data on maternal body size, these variables were entered into multiple regression models to predict clutch size and timing of egg laying. The model for clutch size was accurate to 0.53 ± 0.08 eggs, while the model for oviposition date was accurate to 6.22 ± 0.81 days. Equations were generated that should be applicable to this and other large Cyclura species. © 1995 Wiley-Liss, Inc.     

Is juvenile hormone involved in the maternal regulation of egg size and progeny characteristics in the desert locust?

The role of juvenile hormone (JH) in the maternal regulation of progeny characteristics was examined in the desert locust, Schistocerca gregaria. Female adults of this species are known to produce smaller but more eggs when reared in isolation than do those reared in a group. Eggs laid by isolated females develop green hatchlings typical of solitarious forms, whereas those laid by the latter produce black hatchlings typical of gregarious forms. Topical application of a juvenile hormone analog (JHA), fenoxycarb, or implantation of corpora allata (CA) taken from the migratory locust, Locusta migratoria, caused crowded S. gregaria females to deposit smaller eggs, but did not have a significant effect on the number of eggs per egg pod except at high doses of JHA. The production of smaller eggs by treated and untreated crowded females was closely associated with earlier deposition of the egg pods and shorter oviposition intervals. However, neither JHA application nor CA implantation influenced the progeny characteristics in actively reproducing aged females under crowded conditions, while untreated control females started producing smaller and more eggs upon transfer to isolated conditions. These results may suggest that JH is not directly involved in the maternal regulation of phase-dependent progeny characteristics.     

Ability of ovipositing seed beetles to discriminate between seeds with differing egg loads

Abstract. 1. Previous work has shown that ovipositing females of Callosobruchus maculatus (F.) avoid seeds already bearing eggs, and thereby reduce competition among their larval progeny within seeds.
2. This study demonstrates that females also detect small differences in egg density, and prefer to oviposit on seeds with a lower-than-average number of eggs. A nearly uniform dispersion of eggs is thus maintained even after all seeds bear several eggs.
3. In addition, variation in egg load influences oviposition rate. Transfer of females from seeds with few eggs to seeds with many eggs inhibits oviposition; the reverse transfer stimulates it.
4. The upper surface of the egg chorion or egg 'cover' remains intact on the seed surface after the larva has entered the seed and continues to deter egg-laying for at least as long as the period required for larval development. 14-day-old egg covers provide as much deterrence as freshly laid eggs.     

Ladybird egg cluster size: relationships between species, oviposition substrate and cannibalism

The success or not of ladybirds as biological control agents is dependent on both their foraging behaviour and their individual survival rates. The former is a function of the habitats they utilise; the latter, a consequence of their reproductive strategy. Egg clustering was investigated in two ladybird species, Aphidecta obliterata, a conifer specialist, and Adalia bipunctata, an arboreal woodland generalist. The effect of oviposition substrate (filter papers vs. spruce needles) on clutch size and oviposition preference was also tested. Adalia bipunctata laid significantly more eggs than A. obliterata. The size of egg clusters laid by the two coccinellids varied between species and substrate types. Adalia bipunctata laid larger egg clusters than A. obliterata, with batches reaching a maximum size of 32 eggs on spruce and 41 eggs on paper, while those of A. obliterata contained a maximum of 5 eggs on spruce and 9 eggs on paper. Of the clusters laid by A. obliterata, 18.6% of those on paper and 21.4% of those on spruce contained only a single egg, whereas a minimum of two eggs per cluster were laid by A. bipunctata. Smaller clusters were laid on the spruce cuttings by both species when compared with those laid on the filter paper, but A. obliterata laid significantly more eggs on spruce than on the filter paper (77% vs. 23%), whilst A. bipunctata laid significantly more eggs on the filter paper (91%). It is suggested that coccinellid eggs are more likely to be washed off spruce needles than broad leaves and that, by laying smaller egg clusters on spruce, A. obliterata reduces this risk. Adalia bipunctata usually lays its eggs on the underside of broad leaved trees and thus does not face this risk and thus can lay larger egg clusters. No differences in cannibalism rates were found between the two species. These findings have implications for the use of ladybirds as biological control agents in spruce forests.     

Pseudosexual and dominance behaviour: their relationship to fecundity in the unisexual gecko, Lepidodactylus lugubris

We examined the hypothesis that pseudosexual behaviour facilitates reproduction in the unisexual gecko, Lepidodactylus luguhris. Thirty-eight hatchlings were housed in isolated or social conditions. Age at initial egg development, days between egg development and laying for the geckos'first three clutches of eggs, inter-clutch intervals and dominance behaviours were recorded. Lepidodactylus lugubris developed their first clutch of eggs at about 9·5 months of age after their endolymphatic sacs became externally visible. The geckos laid their eggs about 28–30 days later with smaller geckos requiring more time between egg development and laying than larger geckos. Grouped geckos formed dominance hierarchies. In the triad and quadrad groups, the subordinate geckos'growth rates were slowed and their fecundity was suppressed. Pseudosexual behaviour was not observed in the geckos and was unrelated to fecundity because isolated geckos developed and laid three clutches of eggs at the same rate as grouped geckos. Dominance behaviour, not pseudosexual behaviour, was related to fecundity in L. lugubris. Dominance behaviour may facilitate dispersal in this colonizing species because subordinate geckos could increase their fecundity by moving to less populated areas.