Egg Dumping Research Articles

Breeding Biology of the White-Browed Tit-Warbler (Leptopoecile Sophiae) in Alpine Shrubs, Southern Tibet

Abstract. We studied the breeding ecology of the White-browed Tit-Warbler (Leptopoecile sophiae) in the alpine zone of southern Tibet, elevations 4110–4780 m. The earliest breeder among local passerines, the species initiated egg-laying in early April and ended by late July. We located its domed nests in 13 species of shrubs at an average height of 0.9 m (range 0.2–2.5 m) above the ground. Clutch size averaged 4.7 (range 4–6) eggs, declining through the season. Brood size was 4.3 (range 2–6) at hatching and 3.8 (range 1–5) at fledging. Incubation lasted 20.5 (range 16–23) days, and nestlings fledged at 17.5 (range 14–21) days of age, when they were 4% above the adult weight. Of the nests we observed 66% fledged at least one young. Most pairs were monogamous, and both sexes shared all nesting duties. We noted two females attending a single nest, with a brood of normal size, and egg dumping by an additional female. At the population level, the sex ratio of offspring, determined by sexual differences in plum...

Group structure, mating system and extra‐group paternity in the co‐operatively breeding White‐breasted ThrasherRamphocinclus brachyurus

Avian co‐operative breeders show a diverse range of social and reproductive systems. Here we combined microsatellite genotyping with field observations over three consecutive breeding seasons to investigate the social structure and mating system of the White‐breasted ThrasherRamphocinclus brachyurus. Co‐operative breeding was facultative in this species, with approximately one‐third of nests having helpers. Breeding groups comprised a pair of breeding adults and up to three helpers of either sex. Within co‐operative groups, age was associated with parentage in both sexes. Helpers were retained offspring and either rarely or never achieved parentage in current broods. There was no evidence of egg dumping and a relatively small percentage of chicks (7.5%,n = 67) were fathered by males from outside the co‐operative group. The White‐breasted Thrasher is a globally Endangered species with a very small and declining population. A captive breeding programme for this species is not recommended at present, but may become necessary in the future, and any such programme should take account of our findings regarding breeding behaviour and group structure.

Extra-pair paternity in the socially monogamous Sedge Warbler Actocephalus schoenobaenus as revealed by multilocus DNA fingerprinting

A marked population of Sedge Warblers Acrocephalus schoenobaenus was studied from 1993-95, the reproductive success of pairs was monitored and blood samples collected for DNA profiling. Measures of male physical and behavioural traits were also made, including song repertoire size, the amount of time spent singing and song-flighting, as well as territory quality. Although the Sedge Warbler is generally considered to be monogamous, social polygyny occurred in both 1994 and 1995, giving an overall rate for the two years of 7.7% of males. The overall rate of extra-pair paternity in the population was found to be 8.4% of offspring, with 34.4% of broods containing one or more extra-pair young. No incidences of intra-specific brood parasitism (egg dumping) were identified. Cuckolded males were compared with other males, but no significant differences were identified for any of the physical, behavioural or territorial measures. In particular, there was no evidence that previously detected mate choice cues are related d to patterns of paternity within the population, and hence no evidence that such cues indicate genetic benefits to females. It therefore seems likely that repertoire size reflects male phenotypic quality in this species.

Mixed mating strategies in cooperatively breeding apostlebirds Struthidea cinerea

In cooperatively breeding apostlebirds Struthidea cinerea philopatry by both sexes is coupled with low levels of dispersal, resulting in large groups (up to 17 birds) that may include multiple males and females of varying age and relatedness. We investigated mating patterns within 18 groups of apostlebirds using a set of six polymorphic microsatellite loci. Apostlebirds appear to adopt flexible and context‐dependent mating strategies, with both monogamy and polygamy detected in separate groups. Most groups (11/18, 17/26 group‐seasons) were putatively monogamous, with a single pair matching all typed offspring. Despite the potential for inbreeding, members of breeding pairs were less closely related than other potential within‐group mating combinations. Polygamy was inferred in four groups (four group‐seasons) from the presence of more than four alleles at one locus among offspring within a brood. Pairwise relatedness was lower among adults in polygamous groups than those in groups that included a monogamous breeding pair. There was no conclusive evidence of extra‐group paternity or egg dumping.

Why do host‐deprived seed beetles ‘dump’ their eggs?

Abstract If host seeds are absent, females of the seed beetle Callosobruchus maculatus sometimes ‘dump’ eggs on unsuitable substrates, which causes complete larval mortality and decreases female lifespan. To understand the possible function of this behaviour, the present study examines genetic variation in dumping behaviour between and within populations. When deprived of hosts, females from an African population are much more prone to dump eggs than Asian females, most of which dump no eggs over their lifetimes. Egg dumping therefore cannot be explained as a simple, species‐wide constraint imposed by the accumulation of mature oocytes. A transfer experiment tested the hypothesis that dumping eggs improves a female’s ability to exploit the subsequent availability of seeds, perhaps by preventing a disruption of oocyte maturation. Contrary to prediction, the number of eggs laid after females are transferred to seeds is inversely related to the number dumped during deprivation. Two bidirectional selection experiments revealed heritable variation in egg‐dumping behaviour within the African population. Deprived females from the dumper line dump more than twice as many eggs as do females from the nondumper line. Pre‐adult development time is significantly longer in the nondumper line, which suggests that trade‐offs with other fitness components could maintain variation in egg‐laying behaviour within populations. The divergent responses to host availability by African and Asian females may represent a pleiotropic effect of similarly divergent responses to host quality.

Great reed warbler Acrocephalus arundinaceus and reed warbler Acrocephalus scirpaceus respond differently to cuckoo dummy at the nest

A cuckoo Cuculus canorus dummy was exposed at 24 nests of great reed warbler Acrocephalus arundinaceus (GRW) and 34 nests of reed warbler Acrocephalus scirpaceus (RW) during the egg-laying stage. The eight GRW pairs attacked the cuckoo directly, striking the dummy, but such a behaviour was not recorded in RWs. Also, other behavioural measures (closest distance from the model, duration of distress calls and number of excitement calls) indicated a lower level of defence by RWs compared to GRWs. In the study area, the parasitism rate was much lower in GRWs (1.7% of nests) than in RWs (11.3%). We suggest that one of the reasons for the lower level of cuckoo parasitism on GRWs is its stronger nest defence and hence higher risk of injury or even death for the cuckoo during egg dumping.

Fitness conflicts and the costs of sociality in communal egg layers: a theoretical model and empirical tests

Individuals within complex social groups often experience reduced reproduction owing to coercive or suppressive actions of other group members. However, the nature of social and ecological environments that favour individual acceptance of such costs of sociality is not well understood. Taxa with short periods of direct social interaction, such as some communal egg layers, are interesting models for study of the cost of social interaction because opportunities to control reproduction of others are limited to brief periods of reproduction. To understand the conditions under which communal egg layers are in fitness conflict and thus likely to influence each other's reproduction, we develop an optimality model involving a brood guarding 'host' and a nonguarding disperser, or 'egg dumper'. The model shows that when, where intermediate-sized broods have highest survival, lifetime inclusive fitnesses of hosts and dumpers are often optimized with different numbers of dumped eggs. We hypothesize that resolution of this conflict may involve attempts by one party to manipulate the other's reproduction. To test model predictions we used a lace bug (Heteroptera: Tingidae) that shows both hosts and egg dumpers as well as increased offspring survival in response to communal egg laying. We found that egg-dumping lace bugs oviposit a number of eggs that very closely matches predicted fitness optimum for hosts rather than predicted optimum of dumpers. This result suggests that dumpers pay a social cost for communal egg laying, a cost that may occur through host suppression of dumper reproduction. Although dumper allocation of eggs is thus sub-optimal for dumpers, previous models show that the decision to egg dump is nevertheless evolutionarily stable, possibly because hosts permit just enough dumper oviposition to encourage commitment to the behaviour.

Egg dumping in insects.

Females that place eggs under the care of conspecifics have been labeled egg dumpers. Egg dumping is an effective reproductive alternative that lowers risks for, and has the potential to increase fecundity in, its practitioners. Although insect egg dumpers can be social parasites of the maternal behavior of egg recipients, dumping is more likely to be a viable reproductive alternative when the costs to egg recipients are low and thus the defense by potential hosts against egg dumping intrusions is minimal. These conditions are met in insects that guard only eggs or in insects whose eggs hatch into self-supporting precocial young that need little beyond defense from parents. When this is the case, egg dumping is favored by natural and/or kin selection as a mechanism by which dumpers can avoid parental risks and increase fecundity, and egg recipients can enhance offspring survival by diluting predation.

The Emergence of Conservation Behavior

The Emergence of Conservation Behavior

Egg dumping and life history strategy of Callosobruchus maculatus

Abstract. Previous studies have demonstrated that female Callosobruchus maculatus adjust oviposition rates to cope with changes in host availability. A female lays fewer eggs when host availability is low, and hence decreases larval competition. However, females will also dump eggs on unsuitable substrates under conditions of host deprivation. Because the female does not feed as an adult, egg dumping possibly wastes energy and may thus be a maladaptive behaviour. In this study, the effect of mating and age on the egg‐dumping behaviour and the life history strategy of the female are explored. Under host‐deprived conditions, mating is seen to trigger egg‐dumping behaviour. Also, females mated at 6 days dump significantly fewer eggs and live longer than females mated at 0 or 3 days. Thus, a trade‐off between fecundity and longevity is seen among females subjected to different manipulations. In addition, 6‐day‐old virgin females contain more mature eggs than females mated at 6 days can produce when deprived of hosts. This finding indicates that the female reallocates internal energy resources by oosorption in a resource‐limited environment. To test the maladaptive hypothesis, mated females were deprived of a host for 6 days and then given sufficient hosts each day. The results show that the more eggs dumped by a female in the first 6 days, the more eggs are laid later on beans. Egg dumping is thus not maladaptive.

No evidence of extra‐pair paternity in a colonial seabird, the common tern (Sterna hirundo)

The incidence of extra‐pair paternity and egg dumping was investigated in a colony of common terns (Sterna hirundo), a colonial seabird, in the Venetian lagoon. Ten families were sampled and multilocus DNA fingerprinting analysis was performed. No indication of extra‐pair paternity or egg dumping was found in any of the families. The results are discussed in the light of life‐history strategies, the benefits of coloniality and the evolution of adoption behaviour in the species.

A field test of the intraspecific brood parasitism hypothesis in the golden egg bug ( Phyllomorpha laciniata)

In natural populations of golden egg bugs ( Phyllomorpha laciniata), females lay eggs on plants where they develop unattended, or on conspecifics, where they remain firmly glued until the nymphs hatch and start an independent life. Mortality rates among eggs laid on plants are higher than among eggs carried by adults. Because females cannot lay eggs on themselves, in order to improve offspring survival, they have to lay eggs on other individuals. Two hypotheses have been proposed to explain egg carrying: (1) the mating pair intraspecific brood parasitism hypothesis suggests that females dump eggs on copulating pairs, and (2) the paternal care hypothesis suggests that the system is driven mainly by males accepting eggs to improve the survival rates of their own offspring. Our data from the field show that 77% of the eggs are carried by males, because more males than females carry eggs, and because males carry a greater number of eggs. In addition, we show that mating males carry more recently laid eggs than single males. These results support the view that egg carrying is performed predominantly by males and that eggs are laid on males by their current mating partner, probably between repeated copulations. Males are likely to accept eggs, despite intermediate levels of paternity, because they cannot discriminate in favour of their own eggs, because rejected eggs will face 97% mortality rates on plants, and because they do not suffer mating costs when they carry eggs. However, females carry 23% of the eggs, but no differences in egg carrying have been found between mating and single females, suggesting that this is not the result of egg dumping while females are copulating. Egg carrying by females could reflect low levels of intraspecific parasitism, which is likely to reflect the low rate of successful attempts by egg-laying females who try to oviposit on other conspecifics rather indiscriminately, in an effort to improve the survival of their offspring.

Evolution of Egg Dumping in a Subsocial Insect

Evolution of Egg Dumping in a Subsocial Insect

Evolution of Egg Dumping in a Subsocial Insect

Egg dumping, or abandonment of eggs and young to the care of other conspecifics, frees individuals from costs of maternal care while potentially imposing energetic and ecological costs on egg recipients. It is not clear, however, that egg dumping necessarily represents selfish manipulation of egg recipients, and in some ecological contexts, recipients may benefit from enlarged broods. Thus, egg dumping may either be mutually beneficial for dumpers and recipients or entail costs for dumpers that are compensated by other means, such as improving reproduction of genetically related egg recipients. Here I use field experiments to test the relative importance of manipulation (i.e., "parasitism"), mutualism, and kin selection in the evolution of egg dumping in the tingid lace bug Gargaphia solani. In support of mutualism and kin selection, I found that reproduction of egg recipient G. solani benefits from brood enlargement, most likely because eggs and gregarious nymphs find safety in greater numbers. But contrary to both parasitism and mutualism, egg dumper reproduction was not improved by offspring abandonment. Indeed, dumpers laid smaller clutches than recipients, and dumpers did not convert a survival advantage into greater future reproduction. Genetic analyses of a natural G. solani population revealed, however, that dumpers are related to their egg recipients. Moreover, Hamilton's rule showed that egg-dumping G. solani earn sufficient indirect genetic benefits for kin selection to favor the behavior. Thus, egg dumping in some species may be kin-selected cooperation rather than parasitism or mutualism.

Extra-pair paternity and egg dumping in birds: life history, parental care and the risk of retaliation.

Molecular techniques have revealed striking variation among bird species in the rates of extra-pair paternity (EPP) and intraspecific brood parasitism (IBP). In terms of the proportion of broods affected, rates of EPP and IBP vary across species from 0-95% and 0-50%, respectively. Despite a plethora of hypotheses and several careful comparative analyses, few robust correlates of this interspecific variation have been identified. One explanation for this shortfall is that most comparative studies have tended to focus on contemporary ecological factors and ignored fundamental differences in reproductive biology that evolved millions of years ago. We show that, for both EPP and IBP, over 50% of interspecific variation is due to differences among taxonomic families and orders. Therefore, we test hypotheses that predict interspecific variation in the rate of alternative reproductive strategies should be associated with differences in life history and the form of parental care. Our analyses largely support these predictions, with high rates of reproductive cheating being associated with 'fast' life histories. High EPP rates are associated with high rates of adult mortality and reduced paternal care. High IBP rates are associated with high-fecundity rates. These patterns remain intact whether we use species as independent data points or evolutionary contrasts based on either molecular or morphological phylogenies. These results are interpreted as supporting the idea that alternative reproductive strategies are most common in taxa in which the risks of retaliation are low. We suggest a hierarchical explanation for interspecific variation in the incidence of alternative reproductive strategies. Variation between major avian lineages in the EPP and IBP rates are determined by fundamental differences in life history and parental care that evolved many millions of years ago. Variation between populations or individuals of the same species, however, are more likely to be determined by differences in contemporary ecological and genetic factors.

Hormonal control of egg dumping and guarding in the lace bug, Gargaphia solani (Hemiptera: Tingidae)

This study examines the effect of juvenile hormone (JH) or a similarly acting juvenoid on the expression of maternal egg guarding and its life history alternative, egg dumping, in the lace bug, Gargaphia solani Heidemann. JH manipulations were indirect: methoprene, a synthetic JH analog, and precocene II, an allatocidal phytochemical commonly used to reduce JH synthesis, were applied exogenously to test the hypothesis that high JH titers promote egg production and egg dumping behavior, while low titers terminate egg production and initiate maternal care. As predicted, egg dumpers treated with precocene II ignored dumping opportunities and became egg guarders. Similarly, egg guarders that were treated with methoprene became gravid within 2 days and abandoned their eggs to become egg dumpers. These manipulations suggest that hormones can trigger the expression of both egg dumping and egg guarding in G. solani even when environmental conditions are inappropriate.

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.

Alloparental care between catfishes in Lake Tanganyika

A bagrid catfish Auchenoglanis occidentalis cared for its brood, deposited within an accumulation of shells and gravel in the centre of a large saucer‐like depression, for up to 2 weeks in Lake Tanganyika. Adults of another catfish Dinotopterus cunningtoni (Clariidae) persistently came to A. occidentalis nests near days of host spawning. Eggs of D. cunningtoni were found in the host nests on the day and within 2 days of host spawning. During and for a few days after the end of host brooding, associate species' young of much older ages than the host brood were often found in the nests together with those of a similar age to the host young. Associate young were rarely found in nests before spawning and in unused nests. These findings suggested that this brood‐mixing has two origins: egg dumping by associate adults and voluntary intrusion of large associate young into host nests. Possible benefits to the associate species are to take advantage of nest preparation and parental behaviour of the host species and to feed on the host brood.

Egg-dumping lace bugs preferentially oviposit with kin

Egg dumping, or abandoning eggs and young to the care of other conspecifics, results in an extreme form of alloparental care. It is unclear, however, if egg dumpers discriminate among kin and nonkin egg recipients. In the lace bugGargaphia solani (Heteroptera: Tingidae), some females with eggs (guards) also accept and defend eggs of conspecifics. Other females (egg dumpers) abandon their offspring after oviposition, leaving a single guard as the caregiver. We asked if egg dumpers preferentially dump their eggs among unguarded eggs of kin or nonkin. When given a choice between dumping among eggs of full siblings and eggs of nonsiblings, most eggs (67%) were dumped with full siblings’ eggs. Furthermore, egg dumpers were just as likely to oviposit among eggs of kin with whom they had interacted on a shared host plant during juvenile development as they were to oviposit with kin reared on different host plants. Thus, egg dumpers discriminate kin by using cues associated with eggs, and such cues are not likely to be acquired through interaction on a common host plant environment.

Multi-locus DNA fingerprinting supports genetic monogamy in Florida scrub-jays

Extensive behavioural and pedigree data on a colour-marked population of Florida scrub-jays (Aphelocoma coerulescens) suggested that this cooperatively breeding species is monogamous, with extremely rare exceptions in which males have two mates. We used multi-locus DNA fingerprinting to test these observations by determining genetic parentage. Despite restricted dispersal and high relatedness between behavioural parents and non-breeding members of the group, DNA fingerprints provided sufficient variability to determine parentage unambiguously in almost all cases. We found no evidence of extra-pair fertilisation of females or egg dumping, and confirmed a suspected case of polygyny in which a mother and daughter laid and incubated in the same nest. Our results confirm that detailed behavioural data allow accurate assignment of genetic parentage in this species. In Florida scrub-jays, large territory size may limit opportunities for cuckoldry, and persistent intense competition for limited breeding space may lead to low variance in the quality of established male breeders. These factors would reduce both the opportunity for, and benefits of engaging in extra-pair fertilisations. Delayed dispersal and cooperative breeding in this species have not evolved as avenues for direct reproduction by unpaired individuals.