Maternal provisioning and possible joint breeding in the burrower bug Adomerus triguttulus (Heteroptera: Cydnidae)

Subsocial burrower bugs (Heteroptera: Cydnidae) provide unique opportunities to investigate evolutionary ecological questions regarding parental provisioning and family dynamics. Observations and marked nutlet‐setting experiments in the field showed that Adomerus triguttulus females progressively delivered mint nutlets into nests harbouring nymphs under the litter. More than one female often attended nymphs, but not eggs, in a nest in the field. The number of nymphs aggregating in a nest with a single female was usually smaller than that in a nest with two females, suggesting the joining of different families and facultative joint parental care. There was a positive correlation between the number of nutlets delivered and the number of nymphs in a nest. The number of attendant females also affected the amount of provisioning; more nutlets were found for second‐instar broods with more females. The effect of brood size on provisioning was confirmed for families under laboratory rearing. Maternal provisioning also varied with the developmental stage of offspring; second‐instar broods received more nutlets than first‐instar broods, with a temporal decrease in provisioning during the moulting of nymphs. Considering the growing evidence of food solicitation signals of young in subsocial insects, the observed finely tuned supply of food by the female could be induced by begging signals from the nymphs.     

Maternal effects across life stages: larvae experiencing predation risk increase offspring provisioning

1. Maternal provisioning can reduce offspring vulnerability to predators by promoting offspring growth and eliciting of antipredator behaviours. Mothers perceiving predation risk may improve offspring survival if producing larger, higher‐quality offspring. However, empirical evidence suggests that offspring quality is often reduced, probably reflecting predator‐induced physiological costs, or a selfish maternal strategy aimed at producing more offspring by sacrificing their quality. While perception and impact of predators can vary across the prey's life stage, a majority of studies have focused on understanding how reproductive allocation decisions are influenced by the risk of predation during adulthood. 2. In this study, Leptinotarsa decemlineata beetles were used to examine if the risk of predation during the larval stage: (i) impacts the mother's physiological condition, including body mass and metabolic rate; and (ii) alters maternal allocation of reproductive resources to offspring quantity versus quality. 3. Results revealed that L. decemlineata mothers responded to perceived predation risk by producing clutches with fewer but larger eggs, thus increasing offspring provisioning. Surprisingly, while females that had faced predation risk as larva emerged with a similar body mass to control females, they exhibited lower metabolic rates. 4. Although predation risk in L. decemlineata larvae is known to impair their ability to acquire and maintain energy resources, adult females appeared to ameliorate such costs by improving their metabolic efficiency and by allocating more of their limited reproductive resources to produce fewer but better‐quality offspring.     

Protective function of maternal care against egg predators in the shield bug Sastragala esakii (Hemiptera: Acanthosomatidae)

Females of many acanthosomatid shield bugs attend eggs and young nymphs by covering them with their bodies. Although such form of maternal care has been suggested to have evolved independently in four genera, Elasmucha, Sastragala, Acanthosoma and Sinopla, previous studies exploring its adaptive function have solely focused on species of Elasmucha. This study pioneered an experimental examination of maternal care in the Japanese species Sastragala esakii. Field experiments demonstrated that unattended egg masses suffered intense predation, whereas egg masses attended by their mothers were rarely preyed upon. The ant Crematogaster sp. was the most common egg predator, while two other insect species were also observed to prey on eggs. The exclusion of ant workers and other walking predators from accessing egg masses through the utilization of a sticky trap resulted in a remarkably improved egg survival rate in orphaned egg masses, suggesting that, at least in our study site, maternal care of S. esakii primarily functions to protect eggs from walking predators. Laboratory observations revealed that egg protection against predators was achieved through at least two distinctive defensive behaviors shown by egg-tending females: (i) tilting their bodies when approached by predators; and (ii) fanning their wings when in contact with predators. However, females displayed a limited response to predators approaching from the posterior direction, suggesting that visual cues play a significant role in predator recognition. These results indicate a similarity in the maternal care functions between Elasmucha and Sastragala, and suggest the parallel evolution of female defensive behaviors.     

Effects of trophic-eggs on offspring performance and rivalry in a sub-social bug

Trophic eggs, which are inviable and usually function as a food supply for offspring, have been regarded as extended parental investment or the outcome of parent–offspring conflict in sibling oophagy. Adomerus triguttulus (Heteroptera: Cydnidae) is a sub-social bug showing a complex pattern of maternal care, including progressive provisioning of host seeds and trophic-egg production. To investigate the functions of trophic eggs, we removed trophic eggs from clutches under different resource conditions. The longevity of nymphs was greatly extended by feeding upon trophic eggs when seeds were excluded. When seeds were provided, trophic-egg feeding by nymphs enhanced their development, but there were no significant effects on brood survival. Some viable eggs were also fed upon by sibling nymphs. However, there was no difference in the proportion of viable eggs consumed between clutches with and without trophic eggs. Females lay viable eggs within the first oviposition day. The synchronous hatching resulting from this oviposition mode seems to prevent sib-cannibalism. The body size of females affected their relative investment in trophic eggs; larger females produced more viable eggs with relatively fewer trophic eggs. The functions and adaptive allocation of trophic eggs are discussed in light of the two hypotheses mentioned above.     

Intraclutch egg‐size variation in acanthosomatid bugs: adaptive allocation of maternal investment?

If there are differences in predation risk among the offspring within a clutch, parents may allocate less resources to the offspring facing higher risk. I examined parental investment in terms of egg size within clutches in five species of stink bugs (Heteroptera, Acanthosomatidae). In subsocial Elasmucha and Sastragala species, the female guards her eggs and first-instar nymphs against invertebrate predators by covering her clutch with her body. Large differences in survival from predation between offspring at the centre and offspring at the periphery of the clutch have been reported in such subsocial insects. I found that Elasmucha and Sastragala females laid significantly smaller eggs in the peripheral (and thus more vulnerable) part of the clutch. Phenotypic trade-offs between egg size and clutch size were detected in these subsocial species. Egg size was positively correlated with hatched first-instar nymph size: smaller nymphs hatched from smaller peripheral eggs. In asocial Elasmostethus humeralis , however, no significant difference in size was detected between the eggs at the centre of and those at the periphery of the clutch. Thus, in subsocial acanthosomatid bugs, females seem to allocate their resources according to the different predation risks faced by the offspring within the clutch.     

Differences in the effects of parental age on offspring life history between tropical and temperate populations of milkweed bugs (Oncopeltus spp.)

Summary We compare the effects of parental age on several offspring life history traits in two milkweed bug populations: a typically univoltine population ofOncopeltus fasciatus from the Sacramento Valley of California and a typically multivoltine population ofO. cingulifer from Monteverde, Costa Rica. Reared under identical conditions (27°C, 12 h light: 12 h dark photoperiod), these bugs exhibit significant differences in the effects of parental age on offspring life history. As they age,O. fasciatus females from Sacramento lay clutches of eggs of decreasing average weight and a decreasing proportion of their eggs produce offspring that survive to adulthood. Those offspring which do survive have a significantly faster developmental rate and females have a larger body size at adult eclosion. AsO. cingulifer from Monteverde age, they also produce lighter eggs, but there is no significant change in the offspring developmental rate, survival or female adult body size. We suggest that these results are largely explicable as the consequence of different parental investment strategies associated with the predictable relationship between parental age and time of season inO. fasciatus but not inO. cingulifer. AsO. fasciatus from Sacramento age, they may be increasing their investment per developing offspring so as to increase the probability that nymphs hatching late in the season will reach a prereproductive adult diapause before the first killing frost.     

Prolonged maternal care inElasmucha dorsalis (Heteroptera: Acanthosomatidae)

Female parents ofElasmucha dorsalis attend their offspring and show specific sequential defensive behavior when disturbed. Intense responses of brooding females such as wing fanning were usually triggered by exposure to crushed nymphs. Maternal care continued for a long time, sometimes into the fourth or last nymphal instar. Female removal experiments showed that maternal defense is highly effective under field conditions. High egg mortality observed in the field was probably due to the predation by an ant,Myrmica ruginodis, which occurred at high densities on the host plant,Aruncus dioicus. Most of the egg masses were found on the leaves subtending axillary inflorescence of the female plants, and nymphs usually aggregated and fed on fruits in the inflorescence. Because the food of nymphs is temporally limited, the female can produce only a single brood. Thus, temporality of food resources as well as a heavy predation pressure may have led to prolonged maternal care.     

Ineffective Maternal Care of a Subsocial Bug against a Nymphal Parasitoid: a Possible Consequence of Specialization to Predators

In the subsocial bug Elasmucha putoni, females lay egg masses on leaves of fruit-bearing wild mulberry trees. Parent females invariably remained straddling their offspring until the offspring moulted to the second instar on natal leaves. Thereafter, the females usually settled on the stem of shoots which harboured second-or-later-instar nymphs feeding on fruit or aggregating on leaves, and faced toward the base of the shoot. To evaluate the effectiveness of maternal attendance on parasitism of nymphs by a braconid wasp, I divided the boughs of each mulberry tree into two experimental groups: a single bough which was isolated from the others by Tanglefoot treatments and on which guarding females were removed from broods before the second instar, and other boughs on which broods were left intact as controls. The experiments showed that nymphal parasitism was not affected by the presence of females. The position and orientation of females attending second-or-later-instar nymphs is probably effective for detecting predators approaching the nymphs by walking along the stem; however, this posture may prevent the females from detecting parasitoids, which fly and land directly on plant parts close to the nymphs. The ineffectiveness of the females in providing defence against the parasitoid is possibly associated with a specialization of the attending posture to pedestrian predators: a parental defensive behaviour specific to these predators may make the offspring more vulnerable to the parasitoid.     

Differential efficacy of toxic pederin in deterring potential arthropod predators of Paederus (Coleoptera: Staphylinidae) offspring

This study investigates the effects of pederin, a hemolymph toxin that is accumulated in the eggs of most Paederus females, on potential arthropod predators of the offspring of P. fuscipes and P. riparius. Insects generally do not respond to pederin present in the prey. Paederus larvae are sufficiently agile to escape from these predators by running away, and the eggs are hidden by the females. Unlike insects, (wolf) spiders are deterred by prey with pederin. They turn away from larvae they have already captured and exhibit cleansing behavior. Larvae containing pederin survive the attacks of spiders without damage, whereas larvae descended from females that do not transfer pederin into their eggs are often killed and eaten. In the case of sudden attacks by spiders, the larvae have no chance of escape. Their survival thus depends on chemical defense. These investigations show for the first time why pederin might be of considerable importance for Paederus in the field.     

Unequal maternal investment in offspring quality in relation to predation risk

Equal investment within broods does not always maximize parental reproductive value if the reproductive value of some of the young is low. We examined maternal investment in terms of offspring size in relation to the prospects of survival from predation within broods of the shield bug Elasmucha ferrugata Fabr. (Heteroptera; Acanthosomatidae). Shield bug females guard eggs and first instar nymphs against invertebrate predators by covering the clutch with their body and by behaving aggressively towards their enemies. Survival of eggs was not possible without maternal care. When females were allowed to guard their brood, eggs at the periphery were more vulnerable to predators than eggs at the centre. We found that females laid significantly larger eggs in the safest, central part of the clutch. There seems to be an advantage of large nymph size, since when nymphs were reared separately with low food resources, the larger ones were more likely to survive. Larger nymphs were also more likely to push themselves to the safest, central part of the clutch. Females seem to allocate their resources more to the offspring with the highest probability of avoiding predation. Thus our study supports unequal maternal investment within broods of E.ferrugata.     

Maternal behaviour of a webspinner (Order Embiidina): mother-nymph associations

ABSTRACT.

• 1 The life cycle of Antipaluria urichi (Saussure) (Embiidina: Clothodidae) was aseasonal and without overlap of generations within fifteen closely monitored colonies observed over a 9½ month field season. Adult females remained with their nymphs after hatch but disappeared prior to maturation of their offspring.
• 2 Time budgets of nymphs and their mothers differed significantly. Adult females spent significantly more time spinning silk than their offspring. After the nymphs hatched, the increase in area of the silk covering approximated an exponential curve, reflecting the increase in silk spinning behaviour exhibited by the adult female. These results indicate that silk may be a major contribution of the mother.
• 3 Observations suggest that silk protects embiids from some predators and from rainfall, which is often torrential in the rainforest.
• 4 In experimental field colonies, nymphs with their mothers developed significantly faster than those alone, indicating that maternal behaviours promote the development of their nymphs.

     

Maternal care in Gargaphia solani (Hemiptera: Tingidae)

We investigated the extent to which maternal behaviour exists in the lacebug Gargaphia solani as well as the adaptive relevance of maternal behaviour to predation, interspecific competition, feeding facilitation, and environmental heterogeneity. G. solani females aggressively defend their progeny against predators until maturity but provide no assistance in feeding, aggregating, or locating resources. In the absence of predators, nymphs suffer no ill effects if raised without their mother, but when nymphs are orphaned in normal field conditions only 3% survive to maturity. Such strong selection has given rise to maternal defensive behaviour that enables Gargaphia females to defend their progeny against predators.     

Factors influencing offspring traits in the oviparous multi-clutched lizard,<Emphasis Type="Italic">Calotes versicolor</Emphasis> (Agamidae)

The determinants of offspring size and number in the tropical oviparous multi-clutched lizard,Calotes versicolor, were examined using both univariate and multivariate (path) analyses. InC. versicolor maternal snout-vent length (SVL) and body condition influence clutch mass and clutch size but have no significant influence on offspring size. The positive effect of maternal SVL and body condition on offspring number is counterbalanced by a negative effect of breeding time on egg mass. In fact, breeding time directly influences the offspring body mass and condition through variation in the egg mass. There is a trade-off between offspring mass and condition with offspring number, and breeding time influences both. Offspring hatched from the eggs of early (May–June) or mid (July–August) breeding periods invariably show lower mass and condition than those hatched from the eggs of late breeding season (September–October). Yet, there is no variation in offspring SVL among early, mid and late clutches. Thus, inC. versicolor offspring SVL is optimized while body mass and condition are not optimized.     

Offering offspring as food to cannibals: oviposition strategies of Amazonian poison frogs (Dendrobates ventrimaculatus)

Species utilizing distinct resources for offspring production often show plasticity in reproductive strategies as a function of resource quality. For species using ephemeral pools, strategies are mainly shaped by a time constraint related to pool stability, resource availability and the colonizing community. We studied reproductive strategies in Amazonian poison frogs (Dendrobates ventrimaculatus) that are characterized by oviposition in distinct, small and resource-limited water bodies in leaf axils of plants and the transport of newly hatched tadpoles on the back of males to similar water bodies. Cannibalism of eggs by tadpoles was found to be the main cause of egg mortality. Typically, at the end of the rainy season new clutches of eggs were deposited in water bodies already containing a tadpole. Manipulation of the available number of water bodies showed that this observation did not result from resource limitation. We conclude that D. ventrimaculatus has a plastic reproductive strategy that includes provisioning its tadpoles with fertilized eggs as a function of desiccation risk of water bodies housing its offspring. Provisioning behavior is expected to increase developmental rate and, therefore, chances of metamorphosis for tadpoles that hatched towards the end of the rainy season. The plastic food provisioning strategy may be an important evolutionary link to bi-parental and female care with development of obligate unfertilized egg provisioning in the genus Dendrobates.     

Maternal androgens in eggs of communally breeding guira cuckoos (Guira guira)

Variation of maternal androgens in avian eggs may be a mechanism of maternal influence on offspring development, growth, and/or behavior. We studied yolk androgen concentrations in eggs of guira cuckoos (Guira guira) to understand how females might enhance the success of offspring in a complex communal breeding system. We measured concentrations of androstenedione, 5alpha-dihydrotestosterone, and testosterone in yolks and identified eggs and clutches of individuals in joint nests by yolk protein electrophoresis. Androstenedione had the highest yolk concentration, at least 10 times higher than that of testosterone and 5alpha-dihydrotestosterone. The first eggs of individual females that laid two or three eggs in a joint nest had lower androstenedione concentrations than their second and third eggs, the latter having a lower probability of being ejected from the nest. This implies that guira cuckoo females may influence offspring survival and competitiveness in communal nests by means of differential allocation of androstenedione and laying tactics. There was significant variation in yolk androstenedione among females, but the order in which females entered laying in the communal clutch had no effect on the concentrations. Androstenedione yolk concentrations increased with communal clutch size, which may indicate that higher levels of competition in larger groups lead to higher yolk androgen concentrations. Finally, androstenedione concentrations were higher in clutches in the later wetter periods of the rainy season than during the earlier drier period. This may be explained by the high frequency of large clutches in the later periods, with more females contributing to a joint clutch.     

Mothers influence offspring body size through post-oviposition maternal effects in the redbacked salamander, Plethodon cinereus

In the terrestrial salamander (Plethodon cinereus), previous work has shown that mothers body size is positively correlated to offspring size at the time of hatching even after controlling for the effects of egg size. This study was designed to determine whether maternal body size affects offspring size via pre-oviposition factors (e.g., yolk quality, jelly coat composition, or maternal genes) or post-oviposition factors (e.g., parental care behaviors, parental modification of environment). Gravid females were captured and induced to lay eggs in experimental chambers in which the environment was standardized. Fifteen clutches were exchanged, or cross-fostered, between female pairs differing in body size. Ten females whose eggs were taken away and then returned served as controls for the crossing treatment. Foster mothers did not significantly differ from control mothers in the time spent with eggs, body position, or number of egg movements during brooding. Average egg mass measured midway through development was not significantly correlated to the body size of either the genetic or foster mother, but was correlated to pre-oviposition oocyte size. At hatching, offspring body length was positively correlated to egg size and the foster mothers body size. This correlation suggests that in P. cinereus post-oviposition maternal effects have a greater impact on offspring size than other maternal factors incorporated into the egg prior to oviposition. While our study showed that larger mothers moved their eggs less often and tended to spend more time in contact with their eggs, further work needs to be done to identify the specific mechanisms through which larger mothers influence the body size of their offspring. This is the first experimental demonstration of post-oviposition maternal effects for any amphibian with parental care.     

Brood conspicuousness and clutch viability in male‐caring assassin bugs (Rhinocoris tristis)

Abstract 1. Conspicuousness to mates can bring benefits to both males (increased mating success) and females (reduced search costs), but also brings costs (e.g. increased predation and parasitism). Assassin bugs, Rhinocoris tristis, lay egg clutches either on exposed stems or hidden under leaves. Males guard eggs against parasitoids. Guarding males are attractive to females who add subsequent clutches to the brood. This is an excellent opportunity to study the effects of conspicuousness on the fitness of males and females. 2. Using viable eggs in a multi‐clutch brood as a correlate of fitness, the present study examined whether laying eggs on stems affected (1) female fitness, through exposure to parasitism and cannibalism, and (2) male fitness, through attracting further females. 3. Stem broods were more parasitised. However, males on stems accumulated more mates and more eggs, a net benefit even accounting for parasitism. The eggs gained from being on a stem were cannibalised. By contrast, higher mortality on stems suggests that females should gain by ovipositing on leaves. To the extent that egg viability represents fitness, male and female interests may therefore differ. This suggests a potential for sexual conflict that may affect other species with male care. 4. Despite higher costs, females actually initiated more broods, and subsequently added bigger clutches to broods, on stems than under leaves. This suggests either that viable eggs do not reflect fitness, or that females laid in unfavourable locations. The key is now to address lifetime fitness, since unmeasured factors may affect offspring viability post‐hatching, and to investigate who controls the location of oviposition in R. tristis.     

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.     

Larger clutches of chemically defended butterflies reduce egg mortality: evidence from Battus philenor

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Maternal Defence of a Leaf Beetle is not Effective against Parasitoids but is against Pedestrian Predators

Females of the subsocial leaf beetle Gonioctena sibirica attend their broods on the willow Salix bakko. Females with broods settle on the underside of the basal part of leaves and face towards the base of the shoots. If other arthropods approach such attendant females, the females show aggressive behaviour against the intruder. The effectiveness of maternal care on offspring survival against pedestrian predators and against a parasitoid wasp was evaluated in field experiments using three groups: broods from which parent females were experimentally removed, those from which females were removed and pedestrian predators were excluded by tanglefoot treatments, and control broods. These experiments showed that maternal care was highly effective against pedestrian predators, but parasitism was not affected by the presence of females. Offspring mortality by predation or by parasitism was not usually dependent on larval densities. Position and orientation of attendant females will be effective for them to detect intruders which approach their larvae by walking along the stem, while it can inhibit the females from detecting parasitoids which fly and land directly on the leaf close to the larvae. The results suggest a trade-off in the effectiveness of prey defensive behaviour against different enemy species: prey defensive behaviour specific to one type of the enemy may make the prey more vulnerable to the other.