Contribution of males to brood care can compensate for their food consumption from a shared resource

The sharing of the same food source among parents and offspring can be a driver of the evolution of family life and parental care. However, if all family members desire the same meal, competitive situations can arise, especially if resource depletion is likely. When food is shared for reproduction and the raising of offspring, parents have to decide whether they should invest in self‐maintenance or in their offspring and it is not entirely clear how these two strategies are balanced. In the burying beetle Nicrophorus vespilloides, parents care for their offspring either bi‐ or uniparentally at a vertebrate carcass as the sole food source. The question of whether biparental care in this species offers the offspring a better environment for development compared with uniparental care has been the subject of some debate. We tested the hypothesis that male contribution to biparental brood care has a beneficial effect on offspring fitness but that this effect can be masked because the male also feeds from the shared resource. We show that a mouse carcass prepared by two Nicrophorus beetles is lighter compared with a carcass prepared by a single female beetle at the start of larval hatching and provisioning. This difference in carcass mass can influence offspring fitness when food availability is limited, supporting our hypothesis. Our results provide new insights into the possible evolutionary pathway of biparental care in this species of burying beetles.     

Effects of diapause duration on future reproduction in the cabbage beetle,Colaphellus bowringi: positive or negative?

Abstract.  Cabbage beetles, Colaphellus bowringi , undergoing an imaginal summer and winter diapause in the soil, show a great difference in diapause duration (from several months to more than 3 years) under natural conditions. The effects of diapause duration on future reproduction in the beetle are investigated at 25 °C with an LD 14 : 10 h photoperiod and under natural conditions. The fecundity of postdiapause adults with a short diapause of 5 months and nondiapause adults is similar, showing that a short diapause has no affect on reproduction, whereas the longevity of postdiapause adults with a short diapause of 5 months is significantly shorter than nondiapause adults, showing that a short diapause has a negative affect on longevity. The mean total egg production per female and longevities of postdiapause adults with long diapause periods of 16, 22, 29 and 34 months are similar to nondiapause adults, but the mean daily egg production per female is significantly higher than nondiapause adults, showing that extended diapause has a positive effect on postdiapause reproduction. The offspring of postdiapause parents require a relatively shorter time for egg development compared with the offspring of nondiapause parents, showing that diapause has a positive effect on their offspring's performance. However, there are no significant differences among offspring performance in terms of survival, adult longevity, mean egg production per female and mean daily egg production per female.     

Does resource availability affect offspring begging and parental provisioning in a partially begging species?

In species where parents repeatedly provide their offspring with food, the offspring often communicate their need to the parents. Burying beetles, which breed on a wide size range of carcasses of small vertebrates, are interesting model systems to test theories on begging, because the larvae show partial begging, that is, they obtain food through both signalling to their parents (begging) and feeding directly from the carcass. We manipulated resource availability inNicrophorus vespilloides by providing parents with mouse carcasses spanning a wide size range, and allowing them to rear the larvae that hatched, so that both the amount of resources and the number of siblings varied. Time spent begging by each larva was strongly influenced by the time parents spent near the larvae. Brood size had a nonlinear effect on larval begging, with begging increasing with brood size for relatively smaller broods and decreasing again for larger broods. Carcass size and number of parents present had no effect on begging. Time spent provisioning the larvae by the parents was strongly associated with the time spent begging by each larva. Parents spent more time provisioning under biparental care than under uniparental care, while brood size and carcass size had no significant effect. These findings suggest that the larvae adjust their begging to the behaviour of their parents and the number of siblings, but not to the amount of resources. Furthermore, parents adjust the time spent provisioning to the average amount of begging by each larva in the brood, and not to the availability of resources.     

Flexible parents: joint effects of handicapping and brood size manipulation on female parental care in Nicrophorus vespilloides

Parental care is highly variable, reflecting that parents make flexible decisions in response to variation in the cost of care to themselves and the benefit to their offspring. Much of the evidence that parents respond to such variation derives from handicapping and brood size manipulations, the separate effects of which are well understood. However, little is known about their joint effects. Here, we fill this gap by conducting a joint handicapping and brood size manipulation in the burying beetle Nicrophorus vespilloides. We handicapped half of the females by attaching a lead weight to their pronotum, leaving the remaining females as controls. We also manipulated brood size by providing each female with 5, 20 or 40 larvae. In contrast to what we predicted, handicapped females spent more time provisioning food than controls. We also found that handicapped females spent more time consuming carrion. Furthermore, handicapped females spent a similar amount of time consuming carrion regardless of brood size, whereas controls spent more time consuming carrion as brood increased. Females spent more time provisioning food towards larger broods, and females were more likely to engage in carrion consumption when caring for larger broods. We conclude that females respond to both handicapping and brood size manipulations, but these responses are largely independent of each other. Overall, our results suggest that handicapping might lead to a higher investment into current reproduction and that it might be associated with compensatory responses that negate the detrimental impact of higher cost of care in handicapped parents.     

Residency Time as an Indicator of Reproductive Restraint in Male Burying Beetles

The cost of reproduction theory posits that there are trade-offs between current and future reproduction because resources that are allocated to current offspring cannot be used for future reproductive opportunities. Two adaptive reproductive strategies have been hypothesized to offset the costs of reproduction and maximize lifetime fitness. The terminal investment hypothesis predicts that as individuals age they will allocate more resources to current reproduction as a response to decreasing residual reproductive value. The reproductive restraint hypotheses predicts that as individuals age they will allocate fewer resources to current reproduction to increase the chance of surviving for an additional reproductive opportunity. In this study, we test for adaptive responses to advancing age in male burying beetles, Nicrophorus orbicollis. Burying beetles use facultative biparental care, but the male typically abandons the brood before the female. Previous work in male burying beetles has suggested several factors to explain variation in male residency time, but no study has observed male behavior throughout their entire reproductive lifetimes to determine whether males change residency time in an adaptive way with age. We compared residency time of males that reproduced biparentally, uniparentally, and on different-sized carcasses to determine if they used an adaptive reproductive strategy. Males did not increase residency time as they aged when reproducing biparentally, but decreased residency time with age when reproducing uniparentally. A decrease in parental care with age is consistent with a reproductive restraint strategy. When female age increased over time, males did not increase their residency time to compensate for deteriorating female condition. To our knowledge, this is the first test of adaptive reproductive allocation strategies in male burying beetles.     

Effect of Population Density and Female Body Size on Number and Size of Offspring in a Species with Size‐Dependent Contests over Resources

When size‐dependent contests over resources influence reproductive success, the trade‐off between number and size of offspring depends on the frequency of contests. Under these circumstances, clutch size should decrease and offspring size should increase as contests become more frequent. We tested these predictions with the burying beetle Nicrophorus pustulatus through manipulation of rearing densities. Burying beetles reproduce on small vertebrate carcasses, a rare but high quality food source for the larvae. Large beetles are more likely to win contests over carcasses and gain exclusive access to a carcass. The winner of a contest kills eggs and larvae already present on a carcass. As a result of the rarity of carcasses, burying beetles are unlikely to breed more than once. As predicted, brood size of N. pustulatus decreased with increasing rearing density. Despite a negative correlation between brood size and larval mass, larval mass did not increase with increasing rearing density. This may be due to the special biology of N. pustulatus which can use snake eggs for reproduction. Potentially larger supply of resources and generally small population densities of N. pustulatus may weaken selection on body size and thus the correlation between brood size and larval mass. As size‐dependent constraints can limit reproductive phenotypes, we examined whether female size influenced reproductive phenotype. Small females produced larger broods with smaller, but more variable, offspring than large females. Mechanical constraints of egg size seem an unlikely explanation for the differences because burying beetles can compensate for small egg size through parental care. Energetic constraints may impact small females because body mass and brood size of small females decreased with increasing density. Yet, at all density levels small females produced larger, not smaller, broods than large females. The larger and more variable broods of small females seem to be in agreement with a bet‐hedging strategy.     

Risks for larvae mediated by plasticity in hatching

Risks associated with benthic and planktonic development can be mediated by plasticity in hatching. The ability to delay hatching while awaiting favorable planktonic conditions, combined with the ability to accelerate hatching when encapsulated offspring are at risk, should be advantageous. We tested this predicted association of hatching plasticities with a barnacle. In the winter, broods of barnacles (Balanus glandula) reached hatching‐capable stages at widely varying times, but these broods hatched in the spring within about 2 weeks, consistent with a synchronizing environmental stimulus for hatching. In contrast, the same adults held subsequent broods (during later spring and summer) briefly. Either an environmental stimulus for hatching was not needed later in the season, or it was more frequently present. Dissections of brood lamellae that scattered smaller clumps of the encapsulated nauplii induced hatching. Crabs eating brooding adults had a similar effect: crabs broke the barnacles' tests, and many nauplii hatched. In contrast, when whelks ate barnacles, they left the barnacles' wall plates and opercula in place, and few nauplii were released. In some cases, numerous hatched nauplii were trapped within the test of the killed mother. At a field site with abundant whelks, many dead barnacles had opercular plates in place. Plasticity in hatching of broods adjusted risks for planktonic larvae against risks of death of the parent before release of embryos, but escape or death of brooded offspring depended on the kind of damage to the brooding mother and thus on the kind of predator. Although both predators killed brooding parents, subtle snails imposed a greater risk than crushing crabs.     

The phenology of Onitis alexis (Coleoptera: Scarabaeidae) in the Araluen Valley: Survival in a marginal environment

The introduced African dung beetle, Onitis alexis Klug, has become established in the warmer regions of Australia. The south-eastern limit of its current distribution is Moruya, NSW, and the Araluen Valley 50 km inland. At Araluen newly emerged beetles are present in dung in late spring, summer and autumn. Egg-laying starts 1-2 weeks after emergence and continues throughout the summer and autumn, as indicated by the presence of parous females in the population and of broods under experimental pads. Eggs laid in December/January produce adults in late summer and autumn, those laid from February to April produce adults in the following spring and summer. In the laboratory, mortality of larvae is high in cold (0–16°C), wet conditions and their development is delayed in warm (25°C and 27°C), dry conditions. This delay was confirmed in the field during the summer drought of 1982-83 when predicted times of emergence (based on day-degree summation in the soil) always preceded the observed emergence time of the local population, as well as preceding the emergence of beetles developing from eggs laid at known times. Follicle resorption in adult females was related directly to increasing age and to rainfall. Dung collected from hayed-off pasture did not affect fecundity, but caused larval mortality. Adults survived the winters at Araluen in some years, and immatures survived best during dry winters, being facilitated in this by a cold-induced larval diapause. Onitis alexis larvae can survive wet or dry summers, and cold dry winters (down to about 0°C) but not wet winters. This seems to be the major factor limiting the southern distribution of the species.     

Abundance and reproduction of Nilsson's pipefish on tidal flats

Nilsson's pipefish Syngnathus rostellatus were found on tidal flats in the Dollard (Ems estuary, Wadden Sea) in high numbers in May and June (20 to 150 1000 m^-2) and August and September (150 to 300 1000m^-2). Low numbers were found in April, July, November and December. Nilsson's pipefish with eggs and larvae in their brood pouch were most numerous in May and August and September. Both the fraction of Nilsson's pipefish with a brood pouch and the fraction of brood pouches with eggs and larvae increased with fish length. Analysis of length-frequency distributions suggests that two cohorts live in the Dollard (spring cohort, April–August; summer cohort, July–December). Reproducing animals were found in both cohorts. It was assumed that the spring cohort was the previous year's summer cohort, which migrate to the tidal flats in spring, reproduced and then disappeared. The summer cohort was their offspring (0 group) which leave the tidal flats during November and December.     

Male burying beetles extend,not reduce,parental care duration when reproductive competition is high

Male parents spend less time caring than females in many species with biparental care. The traditional explanation for this pattern is that males have lower confidence of parentage, so they desert earlier in favour of pursuing other mating opportunities. However, one recent alternative hypothesis is that prolonged male parental care might also evolve if staying to care actively improves paternity. If this is the case, an increase in reproductive competition should be associated with increased paternal care. To test this prediction, we manipulated the level of reproductive competition experienced by burying beetles, Nicrophorus vespilloides (Herbst, 1783). We found that caregiving males stayed for longer and mated more frequently with their partner when reproductive competition was greater. Reproductive productivity did not increase when males extended care. Our findings provide support for the increased paternity hypothesis. Extended duration of parental care may be a male tactic both protecting investment (in the current brood) and maximizing paternity (in subsequent brood(s) via female stored sperm) even if this fails to maximize current reproductive productivity and creates conflict of interest with their mate via costs associated with increased mating frequency.     

Do maternal food deprivation and offspring predator cues interactively affect maternal effort in fish?

The state of the environment parents are exposed to during reproduction can either facilitate or impair their ability to take care of their young. Thus, the environmental conditions experienced by parents can have a transgenerational impact on offspring phenotype and survival. Parental energetic needs and the variance in offspring predation risk have both been recognized as important factors influencing the quality and amount of parental care, but surprisingly, they are rarely manipulated simultaneously to investigate how parents adjust care to these potentially conflicting demands. In the maternally mouthbrooding cichlid Simochromis pleurospilus, we manipulated female body condition before spawning and exposure to offspring predator cues during brood care in a two‐by‐two factorial experiment. Subsequently, we measured the duration of brood care and the number and size of the released young. Furthermore, we stimulated females to take up their young by staged predator attacks and recorded the time before the young were released again. We found that food‐deprived females produced smaller young and engaged less in brood care behaviour than well‐nourished females. Final brood size and, related to this, female protective behaviour were interactively determined by nutritional state and predator exposure: well‐nourished females without a predator encounter had smaller broods than all other females and at the same time were least likely to take up their young after a simulated predator attack. We discuss several mechanisms by which predator exposure and maternal nutrition might have influenced brood and offspring size. Our results highlight the importance to investigate the selective forces on parents and offspring in combination, if we aim to understand reproductive strategies.     

Adult feeding affects fecundity of the predator,Calosoma sycophanta (Col.: Carabidae)

Calosoma sycophanta L. adults were fed either gypsy moth (Lymantria dispar L.) larvae or split grapes for set periods of time while their reproduction was monitored. Few female beetles reproduced unless fed gypsy moth larvae during the first week after they ended hibernation. Even females initially fed grapes that were later fed larvae had reduced reproduction. The implications these results have for relationships between beetle and gypsy moth populations are discussed.     

Carcass maintenance and biparental brood care in burying beetles: are males redundant?

1. Burying beetles inter small vertebrate carcasses that ultimately serve as a food source for their developing young. The male remains with the female on the carcass after the brood has been produced, purportedly to aid in the feeding and protection of larvae. However, numerous laboratory experiments have failed to demonstrate a beneficial effect of the male on the growth and survival of offspring.
2. A potential difficulty with laboratory studies is that beetles are typically held under relatively benign conditions, protected from the biotic and environmental challenges that they normally encounter. In nature, males may enhance offspring survival by aiding the female in ridding the carcass of mould, and by helping to preserve the carcass through the secretion of antibiotic substances in the beetles' saliva. To examine more rigorously the potential benefits of male parental care, an experiment was conducted under field conditions in which the reproductive output of male–female pairs was compared to that of single females.
3. Beetles were induced to bury carcasses in soil inside rigid plastic tubes that had been inserted into the ground. The experiment was a paired design involving pairs of sisters reproducing in adjacent tubes; one sister reproduced alone, whereas the other reproduced with the assistance of a male. Soil cores were recovered about 1 month later, and examined for viable pupae.
4. There was no significant difference in the number of offspring produced by single females and those reproducing with the assistance of the male, nor was there any significant difference in total brood mass. These results suggest that any benefits of extended male residency on the carcass do not stem from male participation in carcass maintenance or provisioning young.     

Effects of parental body condition and size on reproductive success in a tenebrionid beetle with biparental care

Abstract. 1. The beetle Parastizopus armaticeps (Coleoptera: Tenebrionidae) inhabits the Kalahari desert of southern Africa, constructs breeding burrows after rainfall, and shows extensive biparental care. Previous work has shown that it is predominantly male size, not female size, that determines breeding success; however, in the field these beetles show size assortative mating. This might obscure or override effects of female size on reproduction. Moreover, the inaccessibility of the breeding burrows makes it impossible to test effects of female and male size on offspring development and survival before adulthood. 2. To disentangle the effects of male and female length, body mass, and body condition on reproductive success, males and females were paired randomly in small breeding cages in the laboratory (n = 887 breeding pairs). The construction of the breeding cages allowed a clear view of the brood chamber contents at each stage in offspring development. Larva, pupa, and imago numbers and development were monitored daily, and imago mass at hatching from the pupa (hatchlings), offspring mass, and offspring body length at complete exoskeleton melanisation (juveniles) were determined. 3. There was a weak positive correlation between body condition and body length for females only. Breeding chronology was related to male body condition: males in better condition were fast to start and finish a breeding bout. Males in better condition produced heavier hatchlings and juveniles, and larger‐sized males produced larger‐sized juveniles. In contrast, numbers of larvae and juveniles produced were determined mainly by female length and body condition: larger females in better condition hatched more larvae and produced more offspring. 4. The results suggest that male size and condition will be the most important determinant of reproductive success under relatively dry conditions, when burrow length is critical for reproductive success. Female size might be more important for the pair's reproductive success under wet breeding conditions, when burrow length is less critical for successful reproduction.     

Cooperation, conflict, and crèching behavior in goldeneye ducks

Crèching behavior, or brood amalgamation, results in offspring being reared by adults other than their genetic parents. Although a variety of hypotheses have been proposed to explain this behavior, most assume either that brood amalgamation is accidental (i.e., nonselected) or that adoption of young is selected for because of social benefits to the young and/or adopting parents. We propose, instead, that brood amalgamation is a function of two separate processes: brood desertion and brood adoption. To examine brood desertion, we develop a graphic model to predict when parents should abandon their young and we test this model experimentally for the Barrow's goldeneye (Bucephala islandica). As predicted, females deserted their offspring when the size of the brood was experimentally reduced. Brood adoption occurred when deserted ducklings joined other broods. However, the success of ducklings in doing so was strongly dependent on the availability of potential host broods and on the age of the recipient broods. Foreign ducklings were readily accepted into young broods (<10 d old) but invariably were rejected from old broods. We could detect no benefits or costs of brood adoption to the host females, contrary to the expectations of a social benefit hypothesis. Our experiments indicate that Crèching behavior is driven by selection on adults to abandon their brood when the benefits of continued investment are outweighed by the reduction in future reproduction and selection on deserted ducklings to join other broods to obtain parental care. Rather than a form of cooperative brood care, Crèching in goldeneyes is perhaps best considered as a form of reproductive parasitism, entailing parent-offspring conflict over brood desertion and intergenerational conflict over adoption of abandoned young.     

Habitat selection and offspring survival rate in three paracoprid dung beetles: the influence of soil type and soil moisture

Paracoprid dung beetles build brood chambers in the soil beneath a dung pat and provide them with dung Onthophagus species lay one egg into each chamber This paper deals with the influence of soil type and soil moisture on micro-habitat selection and survival of offspring m three middle-European Onthophagus-species ( O coenobita, O fracticonis and O vacca) Discrimination between sandy soils with three different loam contents (0%, 20%, 40%) and four different water contents (4%, 8%, 12%, 16%) was tested in the laboratory During the first 24 h of each replicate beetles which colonized one of the patches did not distinguish between different soil conditions Emigration rates, measured as time when 50% of all individuals had left the patch, and numbers of brood chambers proved to be species specific and depended on soil moisture and soil type Survival rates of the larvae in the brood chambers were influenced nearly exclusively by soil moisture The results are discussed in relation to the ecology of the three species and in context with optimal foraging theories     

Brood ball size but not egg size correlates with maternal size in a dung beetle,Onthophagus atripennis

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Maternal and paternal effects on offspring phenotype in the dung beetle Onthophagus taurus

Abstract.— Parents often have important influences on the development of traits in their offspring. One mechanism by which parents are able to influence offspring phenotype is through the level of care they provide. In onthophagine dung beetles, parents typically provision their offspring by packing dung fragments into a brood mass. Onthophagus taurus males can be separated into two discrete morphs: Large, "major" males have head horns, whereas "minor" males are hornless. Here we show that a switch in parental provisioning strategies adopted by males coincides with the switch in male morphology. Male provisioning results in the production of heavier brood masses than females will produce alone. However, unlike females in which the level of provisioning increases with body size in a continuous manner, the level of provisioning provided by males represents an "all-or-none" tactic with all major males providing a fixed level of provisioning irrespective of their body size. Offspring size is determined largely by the quantity of dung provided to the developing larvae so that paternal and maternal provisioning affects the body size and horn size of offspring produced. The levels of provisioning by individual parents are significantly repeatable, suggesting paternal and maternal effects as candidate indirect genetic effects in the evolution of horn size in the genus Onthophagus .     

Social and nonsocial stimuli and juvenile hormone titer in a male burying beetle, Nicrophorus orbicollis

We investigated the interaction of social and nonsocial stimuli on juvenile hormone (JH) titer in male burying beetles (Nicrophorus orbicollis). The initial JH response to discovery of a carcass was substantial (10-15-fold increase over controls) and rapid (<1h), and occurred whether or not a female was present. By 3h after discovery, JH titers were declining, the decline being more pronounced when a female was not present. We also tested the effect of larval stimulation on JH titer in care-giving males by removing a male's brood and replacing it with a brood of first or third instar larvae. Males initially providing care for begging first instar larvae continued to maintain high titers of JH when the replacement broods were first but not third instars. Males caring for third instar larvae (normally low JH titers) maintained low levels of JH regardless of the developmental stage of the replacement brood. This suggests that once males begin to care for nutritionally independent third instar larvae, JH titers remain low regardless of subsequent larval stimulation. Burying beetles are socially and hormonally complex organisms in which stimuli from a breeding resource, mating partners, rivals and young interact to alter the JH profile of breeding adults.     

Bimodal life cycle of the burying beetle Nicrophorus quadripunctatus in relation to its summer reproductive diapause

Abstract 1. Under natural conditions in Kyoto, Japan, the reproductive activities of Nicrophorus quadripunctatus Kraatz (Coleoptera: Silphidae) decreased in summer and the species showed a bimodal life cycle.
2. In the laboratory, most adult pairs raised at 20 °C under a LD 12:12 h regime reproduced when provided with a piece of chicken. In adults raised at 20 °C under a LD 16:8 h regime, however, both reproductive behaviour and ovarian development were reduced. It is concluded that these adults entered a reproductive summer diapause.
3. High temperature (25 °C) also suppressed the reproductive behaviour even under a favourable LD 12:12 h regime. In the field, therefore, adults reduce their reproductive activity in summer because of diapause induced by long-day photoperiods and direct inhibition of reproduction by high temperatures.
4. When the temperature was changed from 20 °C to 25 °C immediately after hatching of larvae, they reached the wandering stage in 95% of adult pairs. When the temperature was changed from 20 °C to 25 °C immediately after oviposition, however, no larvae hatched in 85% of pairs. Egg mortality was significantly higher at 25 °C than at 20 and 22.5 °C; no eggs hatched at 27.5 °C. The physiological mechanisms for reducing reproduction probably prevent the beetles from inefficient oviposition in summer.