Proximate Factors Regulating Maternal Options in the Eggplant Lace Bug, Gargaphia solani (Heteroptera: Tingidae)

The eggplant lace bug, Gargaphia solani, was used to investigate the proximate factors regulating maternal care and a noncaring, condition dependent strategy called egg dumping. We hypothesize that the act of delaying oviposition while searching for a dumping opportunity suppresses oogenesis and triggers guarding behavior. We examined several predictions of this hypothesis by measuring: (1) whether females do delay oviposition in the absence of dumping stimuli, (2) whether females in transition between egg dumping and egg guarding are capable of expressing either reproductive option, (3) the effect of nymphal interactions and antennal ablation on the duration of maternal care, and (4) oogenesis in guarding and dumping females. We found that females without a dumping opportunity wait, on average, 30 h longer to oviposit than females exposed to a dump mass. Females that had initiated their own egg mass could resume eggdumping if they had laid less than half of their eggs but were unlikely to abandon their eggs when most had been laid. Maternal care in G. solani can be prolonged if interactions with nymphs are artificially prolonged. Females require antennae to maintain maternalcare. Presumably antennae transduce cues from eggs and nymphs. Dissections of dumping and guarding females 72 h after their first oviposition demonstrated that dumpers continue to produce primary oocytes after a dumping event but guarders terminate oogenesis whilecaring for their first brood. We interpret all of these results within the context of the hypothesis that juvenile hormone titers regulate the expression of both egg dumping and egg guarding.     

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.     

Host discrimination modulates brood guarding behaviour and the adaptive superparasitism in the parasitoid wasp Trissolcus semistriatus (Hymenoptera: Scelionidae)

Because hosts utilized by parasitoids are vulnerable to further oviposition by conspecifics, host guarding benefits female wasps. The present study aims to test whether female adults regulate brood guarding behaviour by host discrimination in a solitary parasitoid Trissolcus semistriatus by presenting an intact or parasitized host egg mass to a female adult. Virgin females without oviposition experience have host discrimination ability, which enables them to adjust the number of eggs laid in the hosts. Mating experience increases superparasitism by female adults, whereas mated females achieve a higher discrimination ability as a result of oviposition experience and show a lower superparasitism rate. As expected, females exhibit brood guard after parasitizing an intact host egg mass, whereas those females visiting a previously parasitized host egg mass, do not. Because the survival of eggs in superparasitized hosts is relatively low, regulating brood guarding behaviour by host discrimination is adaptive for female wasps.     

Costs and benefits of maternal behaviour in the green lynx spider (Oxyopidae,Peucetia viridans)

I examined the adaptiveness of maternal behaviour in the green lynx spider Peucetia viridans (Hentz), by measuring the costs and benefits to the female of egg-sac tending. P. viridans guards her egg sac for 6–8 weeks until the young have emerged and dispersed. I removed females from egg sacs in the field either immediately after oviposition or after they had guarded the egg sacs for 2 weeks, and compared the fate of these sacs with that of control sacs which had females present throughout. Survival rates of egg sacs were significantly improved by the presence of a female. The major sources of mortality were ant predation and sac dislodgement, both of which were significantly reduced by a guarding female, and mantispid brood parasitism, which was not affected by the female. To measure the spiders' lifetime reproductive output, I allowed caged females to construct egg sacs. Half had their first egg sac removed shortly after oviposition, while half were allowed to guard. Significantly more non-guarding than guarding females produced second egg sacs. Thus for the green lynx spider, egg sac guarding decreases lifetime egg production, but increases offspring survival. Under north Florida conditions, the benefits of guarding outweight the costs, and the females have a higher lifetime reproductive success than if they did not guard.     

Individual Variation in Trophic Egg Production: Evidence for Maternal Manipulation in Response to Resource and Competition Levels

Females of the subsocial shield bug Parastrachia japonensis (Heteroptera: Parastrachiidae) incorporate trophic eggs (nutritive eggs) into their egg mass. Considerable variation occurs among females in trophic egg number and the proportion of an egg mass that is composed of trophic eggs. Because trophic eggs are essential to the development and survival of young, this variation could significantly impact female fitness. We tested the hypothesis that trophic egg abundance is induced by maternal phenotype (weight, body size) and resource exposure. We predicted that resource limitations would cause females to produce fewer fertile eggs and more trophic eggs and that larger and heavier females would produce more of each egg type. Females ovipositing early in the season are exposed to different resource conditions than those that oviposit late. Thus, we compared egg production patterns between these two groups and several other factors related to nesting. No correlation was seen between body size and trophic egg abundance, or, indeed, egg production, overall; however, heavier females produced heavier egg masses. Counter to our prediction, late females, which had greater access to food, produced significantly more total eggs, fewer fertile eggs, and more trophic eggs than early females. A binomial generalized linear model analysis indicated that the factors most correlated with the percentage of an egg mass destined to become trophic eggs were resource abundance, resulting from early or late oviposition, and distance of the nest from the host tree, with closer females producing more trophic eggs. The findings support our hypothesis that resource availability and, to a lesser extent, maternal phenotype affect trophic egg abundance.     

A reproductive tradeoff in an herbivorous lady beetle: egg resorption and female survival

A reproductive tradeoff between current egg production and subsequent survival in a lady beetle, Epilachna niponica, a specialist herbivore on a thistle, Cirsium kagamontanum, was investigated at the two study sites, A and F. Survival of reproductive females decreased consistently from early May until mid-June, but apparently increased thereafter. In contrast, males showed a consistent decrease in survival throughout the reproductive season, without any sign of recovery. Dissection of ovaries of sampled females revealed that egg resorption increased late in the reproductive season, coincident with increased female survival. Reproductive females stopped oviposition immediately after a large flood in 1979 at site F. Two weeks after the habitat perturbation, females resumed oviposition in response to a flush of new leaves on damaged plants. Female survival sharply increased during the nonoviposition period, and declined when egg-laying resumed. Approximately 40% of long-lived reproductive females at site F survived up to the following reproductive season in the next year. Also, some of these long-lived females were observed ovipositing in the following reproductive season. The long-lived reproductive females which had previously invested in reproduction survived equally well as newly emerged females which had not reproduced in summer. These results suggest that there is a reproductive tradeoff between current egg production and subsequent survival. Egg resorption may be an adaptive ovipositional response to habitat perturbation such as flooding, which considerably reduces offspring fitness due to absolute shortage of food. Also, increased female survival accompanied by egg resorption enhances the likelihood of the future oviposition in the second reproductive season, thereby increasing a female's lifetime reproductive success.     

Subsociality and female reproductive success in a mycophagous thrips: An observational and experimental analysis

Oviparous females of the haplodiploid, facultatively viviparous thrips Elaphrothrips tuberculatus(Thysanoptera: Phlaeothripidae) guard their eggs against female conspecifics and other egg predators. The intensity of maternal defense increases with clutch size. Field and laboratory observations indicate that cannibalism by females is an important selective pressure favoring maternal care. Experimental removals of guarding females showed that egg guarding substantially increases egg survivorship and that the survivorship of undefended eggs is higher in the absence of nonguarding female conspecifics than in their presence. The fecundity of viviparous females increases with the number of eggs cannibalized. The reproductive success of oviparous females increases with body size and local food density and decreases with local density of breeding females. Social behavior may not have advanced beyond maternal care in Elaphrothrips tuberculatusbecause, relative to Hymenoptera, capabilities for helping relatives are few or nonexistent, and the causes of variation in female reproductive success are not influenced easily by cooperation among females.     

Factors influencing egg parasitism in sub‐social insects: insights from the treehopper Alchisme grossa (Hemiptera,Auchenorrhyncha, Membracidae)

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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.     

Recognition and avoidance of previously laid egg masses by the oblique-banded leafroller (Lepidoptera: Tortricidae)

Investigations were conducted on oviposition by laboratory-reared oblique-banded leafrollers, Choristoneura rosaceana Harris. In paperbag oviposition chambers, females deposited their egg masses in a significantly nonrandom, uniform manner in 16 of 20 replicates. Inpetri dish oviposition chambers, gravid females were exposed to egg masses recently deposited by other females. As determined by the magnitude of the mean direction vector of the second egg mass in relation to the first, females detected and avoided egg masses laid by other females. In addition, they avoided waxed paper from which a recently oviposited egg mass had been removed. This phenomenon is most likely due to a pheromone laid down with each egg mass.     

Relationship between reproductive resource allocation and resource capacity in the matriphagous spider, <Emphasis Type="Italic">Chiracanthium japonicum </Emphasis>(Araneae: Clubionidae)

Females of the Japanese foliage spider, Chiracanthium japonicum, are eaten by their offspring at the end of the maternal care period. To examine the patterns of allocation of maternal investment to their offspring associated with female resource capacity, the amounts of female body reserves accumulated before oviposition, reproductive components at the egg-production phase and those at the matriphagy phase were measured using an artificial breeding nest. Regardless of size, female bodies were completely consumed by the offspring, and larger females, i.e. those having larger reserves, produced a larger number of offspring, but not larger offspring. Furthermore, the proportion of reserves allocated to egg production was not affected by the total amount of the reserves, which indicated that the females systematically divided the resources for individual offspring between egg yolk and food for the growth and survival of the offspring. These results suggest that C. japonicum females adjust egg production to their own resource capacity so that they can achieve an investment per individual offspring which is not dependent on resource capacity. Electronic Publication     

Oviposition site selectivity of some stream-dwelling caddisflies

Population dynamics depends upon the spatial distribution of individuals in heterogeneous environments. The various processes surrounding insect oviposition are central to understanding their population dynamics because the choice of oviposition site ultimately influences the survivorship and spatial distribution of their progeny. Aquatic insects are often assumed to have non-selective oviposition habits, but empirical data are scarce and selective oviposition may be quite common. We quantitatively sampled egg masses of stream-dwelling caddisflies (Trichoptera) that specialise in egg-laying on hard substrata underwater, in order to characterise oviposition site selectivity and test for communal oviposition. In a field survey of two Scottish streams, we sampled egg masses of three species, Polycentropus flavomaculatus, Hydropsyche siltalai, Rhyacophila dorsalis, with the aim of testing whether egg mass abundance varied with current (riffles vs. pools), location within the channel (margins vs. centre) and rock exposure (emergent vs. fully submerged). In one stream, we captured adults landing on emergent rocks and assessed whether females were modified morphologically for swimming. The egg masses of two species (P. flavomaculatus, H. siltalai) occurred primarily on submerged rocks in pool margins, and adult females had legs modified for swimming. In contrast, egg masses of R. dorsalis were most abundant on the underside of emergent rocks in riffles, and females were not modified for swimming. Communal oviposition was evident for all three species, with most egg masses aggregated on the minority of potential rocks. How females locate oviposition sites and the consequences of these highly specialised oviposition behaviours to the survival and spatial distribution of larvae now require investigation. The effects of these behaviours on population dynamics are likely to differ from terrestrial herbivores because oviposition sites are not food resources for these aquatic species.     

Costs and Benefits of Maternal Care in a Subsocial Spider,Coelotes terrestris

The maternal social spider Coelotes terrestris demonstrates extended care towards its progeny: the mother guards its egg sac for 3–4 weeks, then stays with its young from the time of their emergence until their dispersal about 1 month later. The present investigation evaluates the adaptiveness of these maternal behaviours by comparing the fitness of females performing them with that of females separated from the egg sac or the spiderlings. By protecting their egg sacs from predation and parasites, and by pursuing this task while supplying the young with food, mothers enhance the survival rate and the development of many of their spiderlings. The costs linked with these activities, estimated by the ability to produce another clutch, appear variable according to the stage in the reproductive cycle. In such terms, the egg sac guarding appears to have a low cost in relation to the care given to the spiderlings.     

Ovary structure in a presocial insect,Elasmucha grisea (Heteroptera,Acanthosomatidae)

First generation egg clusters of Elasmucha grisea are more closely guarded than second generation clusters. The ovaries of this species are structured to enhance this behavior. The population of E. grisea from S-W Poland breeds in the spring (May–June) and late summer (July–August). The second generation clutches contain fewer eggs and are destroyed 3–4 days after oviposition by predators and parasitoids.The ovary structure in the studied species differs from that found in other Heteroptera. The average number of ovarioles per ovary is 24 while in the other investigated species the number of ovarioles per ovary is 6–7. Lateral oviducts are elongated and the ovarioles are arranged in a pennate pattern. Each ovariole contains only one growing ovarian follicle. Differentiation of the ovarioles and ovarian follicles is synchronised thus enabling simultaneous oviposition. A comparative analysis of the ovary structure during the life cycle, particularly the presence of atresive ovarian follicles in the ovarioles of egg- and nymph guarding females, as well as the shape and structure of the apical part of the tropharium all support the hypothesis of cooperation between females in E. grisea. A similar ovary structure has been observed in the Coccoidea (Hemiptera, Homoptera) which indicates presocial behavior.     

Does it Pay to Care?: Exploring the Costs and Benefits of Parental Care in the Hibiscus Harlequin Bug Tectocoris diophthalmus (Heteroptera: Scutelleridae)

Knowledge of the selective pressures favouring parental care can inform our understanding of the evolutionary origins and transitions of sociality in insects. Here, we report upon investigations designed to estimate the costs and benefits of parental care in the egg‐guarding hibiscus harlequin bug Tectocoris diophthalmus (Heteroptera: Scutelleridae). We found that the presence of a guarding female had no effect on hatching success under benign laboratory conditions. In the wild, however, guarding decreased the likelihood of total clutch failure, and produced a fourfold greater egg‐hatching rate relative to unguarded clutches. Females guarded against generalist invertebrate egg predators, including conspecific nymphs, but were ineffective against hymenopteran egg parasitoids. Females continued to feed during the guarding period and exhibited no signs of weight loss or increased mortality due to this behaviour. We did not observe the production of subsequent clutches in any experimental females; therefore, the lifetime fecundity costs of providing parental care in T. diophthalmus remain indeterminate. Overall, maternal egg guarding appears to function as a relatively low cost–low benefit strategy that increases hatching success by protecting against predation – but not parasitism.     

Female Reproductive Cycle and Sexual Conflict over Precopulatory Mate-guarding in Thermosphaeroma (Crustacea, Isopoda)

In species with time-limited opportunities for insemination, precopulatory mate-guarding is expected to coevolve with the duration of female reproductive cycles. Despite this adaptation to female characteristics, it may also be advantageous for males to adjust the duration of guarding with respect to sex ratio because the benefits of guarding are dependent on the availability of females. If female fitness is reduced because of guarding, male guarding behavior leads to intersexual conflict. We studied these aspects of male mate-guarding behavior in two closely related, thermal-spring isopods (Thermosphaeroma). First, guarding duration showed species specificity which was related to the duration of reproductive cycle; cycle length for females and duration of guarding by males in T. milleri were twice as long as in T. thermophilum. Second, males in both species adjusted their guarding duration with sex ratio, guarding longer when a competing male was present. Third, in T. thermophilum, ovarian development began immediately after the birth of the previous brood and continued through guarding, sexual molt and post-molt periods until oviposition, whereas in T. milleri, ovarian development was largely postponed until the post-molt period. Because guarding during ovary provisioning periods may be costly for females, we tested the existence of intersexual conflict over guarding duration in T. thermophilum. We compared the duration of guarding of control pairs with those of pairs in which either male guarding ability or female ability to resist guarding was reduced experimentally. Guarding durations for manipulated and control males were equal, but manipulated females were guarded longer, suggesting that conflict exists and that females can effectively shorten guarding duration by their behavior. Moreover, we suggest that selection in the context of intersexual conflict may play an important role in the evolution of delayed oviposition and sperm-storage organs in mate-guarding crustaceans.     

Oviposition preference for novel versus normal food resources in laboratory populations ofDrosophila melanogaster

Female fecundity, oviposition preference and specificity on one normal and two novel food media were assayed on four laboratory populations ofDrosophila melanogaster, revealing considerable among- and within-population variation in oviposition preference. Overall, there was a significant tendency of females to prefer novel media to their normal banana food as an oviposition substrate. Specificity in the populations was fairly high, implying that a large proportion of females tended to lay the majority of their eggs on the preferred medium. The results showed that oviposition preference for a given food medium could be affected by the alternative provided, and that, consequently, oviposition preference for a given food medium versus another cannot be predicted based upon a knowledge of what the preference for each of the two media was versus a common third medium. Specificity, on the other hand, was not significantly affected by the type of alternative food media provided in a given trial. Moreover, comparison of results from fecundity and oviposition preference assays also showed that the egg laying behaviour ofDrosophila females in response to different food media may be different in choice versus no-choice situations. Thus, a substrate on which fecundity is higher than on another, when assayed in a no-choice situation, may not be preferred over the other substrate when a choice between the two is provided to the ovipositing females. The latter two results point to possible complexity in the responses of females to various oviposition substrates based upon the overall setting of the assay, including the alternative substrates present for egg laying.     

Mating behavior and the evolutionary significance of mate guarding in three species of crane flies (Diptera: Tipulidae)

Three species of crane flies-Dactylolabis montana, Limonia simulans,and Antocha saxicola-gather near streams to mate and oviposit. All species are polygamous and sex ratios at these sites are male-biased. After a short mating bout, males guard females by standing over them during oviposition. Sperm competition appears to be intense and to follow last-male advantage, based on the packing of sperm within the two elongate spermathecae. Males of A. saxicolasuccessfully defend against rivals over 85% of the time. In contrast, defending males of D. montanaand L. simulanslose the female over 65% of the time during interactions with rivals. Despite the high frequency of loss, defending males gain additional oviposition time by engaging rivals in combat while the female continues to oviposit. Thus, a guarding male does not have to retain the female for guarding to be adaptive. Legs and claws of all species are sexually dimorphic and play an important role in guarding and defending.     

Seasonal decline in plant defence is associated with relaxed offensive oviposition behaviour in the viburnum leaf beetle Pyrrhalta viburni

1. Plant defence of Viburnum shrubs against oviposition by its specialist herbivore, the viburnum leaf beetle [VLB Pyrrhalta viburni(Paykull)], involves an egg‐crushing wound response in twigs. Although the response is variable among Viburnum species, it can have a strong impact on egg survivorship. Beetles typically aggregate egg masses with conspecifics along infested twigs, forming clusters that can overwhelm the twig response. It was investigated whether twig responses and beetle oviposition behaviour vary seasonally. 2. In a field experiment, twig defences decreased towards the end of the VLB oviposition period: wound response of the North American Viburnum dentatum L. and the European V. opulus L. was reduced by 100% and 54%, respectively, in September compared with the July to August period. 3. Oviposition trials demonstrated a corresponding behavioural change: VLB females displayed aggregative oviposition in August, but not in September. 4. Further tests revealed that late‐season VLB females reverted to aggregative oviposition after being kept on uninfested twigs, whereas females kept on heavily infested twigs did not. This behavioural change suggests that relaxation of aggregative oviposition originates from cues associated with high densities of egg masses. 5. Relaxation of aggregative oviposition may be adaptive (and beneficial for invasion) on shrubs with low levels of defences by reducing intra‐specific competition.