Brood size and begging intensity in nestling birds

Theoretical models suggest that sibling competition should selectfor conspicuous begging signals. If so, begging intensity mightbe expected to increase with the number of competitiors. Thepurpose of our study was to examine the relationship betweenbegging intensity and brood size using nestling tree swallows(Tachycineta bicolor) as our model. Over 2 years, we videotapedbegging behavior in unmanipulated broods of different sizes.We found that begging intensity increased with brood size. Theaverage weight of nestlings in each brood did not vary withbrood size, but feeding rate per nestling decreased with broodsize, suggesting that nestlings in larger broods begged moreintensively, possibly because they were hungrier. We also conductedan experiment to examine the effect of nest mates on beggingin different-sized broods. We found that nestlings with similarweights, previous competitive environments, and food deprivationbegged more intensively in large broods than in small broods.Overall, our study indicates that begging intensity increaseswith brood size in tree swallows. This relationship may resultfrom interactions among brood mates rather than from lower feeding rates to individual nestlings in larger broods.     

Genetic evidence for prevalence of alloparental care in a socially monogamous biparental cichlid fish,Perissodus microlepis,from Lake Tanganyika supports the “selfish shepherd effect” hypothesis

Alloparental care – care for unrelated young – is rare in animals, and its ecological or evolutionary advantages or, alternative maladaptive nature, remain unclear. We investigate alloparental care in the socially monogamous cichlid fish Perissodus microlepis from Lake Tanganyika that exhibits bi‐parental care. In a genetic parentage analysis, we discovered a surprisingly high percentage of alloparental care represented by brood mixing, extra‐pair paternity and extra‐pair maternity in all broods that we investigated. The percentage of nondescendant juveniles of other parents, i.e., brood mixing, ranged from 5% to 57% (mean = 28%). The distribution of genetic parentage also suggests that this socially monogamous species has, in fact, polygamous mating system. The prevalence of genetically mixed broods can be best explained by two, not mutually exclusive hypotheses on farming‐out and fostering behaviors. In the majority of broods, the sizes of the parents’ own (descendant) offspring were significantly larger than those of the adopted (nondescendant) juveniles, supporting the ‘selfish shepherd effect’ hypothesis, i.e., that foster parents preferentially accept unrelated “smaller or not larger” young since this would tend to lower the predation risks for their own larger offspring. There was also a tendency for larger parents particularly mothers, more so than smaller parents, to care predominantly for their own offspring. Larger parents might be better at defending against cuckoldry and having foreign young dumped into their broods through farming‐out behavior. This result might argue for maladaptive effects of allopatric care for the foster parents that only larger and possibly more experienced pairs can guard against. It needs to be determined why, apparently, the ability to recognize one's own young has not evolved in this species.     

Sexual conflict among polygynous pied flycatchers feeding young

In the polygynous pied flycatcher, Ficedula hypoleuca, reproductivesuccess of females is constrained by male food provisioningduring the nestling period. Hence, there will be conflictinginterests among the male and each of his mates as to how malefeeding effort should be shared among broods. This paper describesthree experiments designed to examine the parental behaviorof the members of a bigynous trio, i.e., the male and his twomates, in light of these conflicts. In all experiments, primaryand secondary broods were manipulated to hatch on the same dayto reduce the difference in brood-reproductive value due toage. Males divided their effort equally when the two broodswere the same size. However, males did not allocate their investmentin proportion to brood size when brood sizes differed, but investedmore heavily per young in the larger broods. This finding suggeststhat males tried to optimize the joint effort of their two mates.Males and females showed similar responses to experimental reductionin brood demands, which indicates no difference in their willingnessto invest in offspring. When one of the male’s mates wasremoved temporarily, the male increased his total feeding rateand provided proportionately more food to the "motherless" brood.Through flexible allocation of parental investment, males seemable to optimize their reproductive interests in the two broods.The only way a polygynously mated female might successfullyincrease the amount of male assistance at her nest is to makeher own brood more valuable for the male, relative to the otherbroods he might have. We discuss some ways this might be achieved.[Behav Ecol 1991;2:106–115]     

Insemination Capacity and Dispersal in Relation to Sex Allocation Decisions in Goniozus legneri (Hymenoptera: Bethylidae): Why Are There More Males in Larger Broods?

Models considering sex ratio optima under single foundress strict local mate competition predict that female bias will be reduced by stochasticity in sex allocation, developmental mortality of males and limited insemination capacity of males. In all three cases the number of males per brood is expected to increase with brood size. Sex ratio optima may also be less female biased when several mothers contribute offspring to local mating groups or if non‐local mating occurs between members of different broods; again more males are expected in larger broods. In the parasitoid wasp Goniozus legneri (Hymenoptera: Bethylidae), sex allocation has only a small stochastic component, developmental mortality is low and non‐siblings are unlikely to develop in the same brood. However, the number of males per brood increases with the size of the brood (produced by a single mother). We investigated the further possibilities of limited insemination capacity and non‐local mating using a naturalistic experimental protocol. We found that limited insemination capacity is an unlikely general explanation for the increase in number of males with brood size. All males and females dispersed from both mixed and single sex broods. Although most females in mixed sex broods mated prior to dispersal, these data suggest that non‐local mating is possible, for instance via male immigration to broods containing virgin females. This may influence sex ratio optima and account for the trend in male number.     

Local Mating,Dispersal and Sex Ratio in a Gregarious Parasitoid Wasp

Parasitoid sex ratios can be greatly influenced by mating and dispersal behaviour. Many sex ratio models assume that mating is strictly local (only mated females disperse from the natal patch) and that a single male is sufficient to inseminate all females in a brood. Bethylids (aculeate parasitoids) have been used to test predictions of these models, but less attention has been paid to testing their underlying assumptions. We investigated the timing of eclosion, mating and dispersal in mixed-sex and single-sex broods of the bethylid wasp Goniozus nephantidis. In mixed-sex broods, almost all females mate before dispersal and a single male is sufficient to inseminate virtually all females, even when brood sizes are large. Males disperse from both mixed-sex and all-male broods, but males in all-male broods disperse more slowly. Virgin females disperse from all-female broods, which are common. Virgin females can produce a brood, mate with their own sons and subsequently produce mixed-sex broods, but their success rate is very low. Virgin females could potentially circumvent sex allocation constraints by superparasitizing mixed-sex broods, but when presented with hosts bearing mixed-sex broods they destroy all members of the initial brood before ovipositing. Because of the high prevalence of single-sex broods and dispersal of both sexes, the mating structure of G. nephantidis is unlikely to conform to the assumption of strict local mating.     

Evidence for Strategic Sex Allocation in the Guppy (Poecilia reticulata): Brood Sex Ratio and Size Predict Subsequent Adult Male Mating Success

Sex allocation theory predicts that females should produce more sons when the reproductive success of sons is expected to be high, whereas they should produce more daughters, not daughters when the reproductive success of sons is expected to be low. The guppy (Poecilia reticulata) is a live‐bearing fish, and female guppies are known to produce broods with biased sex ratios. In this study, we examined the relationship between brood sex ratio and reproductive success of sons and daughters, to determine whether female guppies benefit from producing broods with biased sex ratios. We found that sons in male‐biased broods had greater mating success at maturity than sons in female‐biased broods when brood sizes were larger. On the other hand, the reproductive output of daughters was not significantly affected by brood sizes and sex ratios. Our results suggest that female guppies benefit from producing large, male‐biased brood when the reproductive success of sons is expected to be high.     

Nestling sex predicts susceptibility to parasitism and influences parasite population size within avian broods

Vertebrate hosts differ in their level of parasite susceptibility and infestation. In avian broods, variation in susceptibility of nestlings to ectoparasites may be associated with non‐uniform distributions of parasites among brood mates, with parasites concentrating feeding on the most vulnerable hosts. The presence of a highly susceptible nestling in a brood can benefit the remaining young by reducing the parasite pressure they experience; however, from a parasite’s perspective, broods with fewer susceptible hosts may provide effectively fewer resources than broods of the same size containing a greater abundance of susceptible hosts, and this could limit the number of parasites that a host brood can sustain. To test whether variation in number of susceptible hosts affects the number of parasites in bird nests, we first examined the role of host sex and induced immunity (via methionine supplementation) on susceptibility of mountain bluebirds Sialia currucoides to parasitism by blow flies Protocalliphora spp. We then assessed the effect of variation in number of susceptible hosts on the number of parasites inhabiting the nest. Only females showed a benefit of methionine supplementation, gaining mass more rapidly following supplementation compared to males. This suggests that females are more susceptible to parasites in this system; this was further supported by parasite feeding trials, in which parasites extracted larger blood meals from female than male hosts. Finally, the abundance of parasites in nests was predicted by brood sex ratio: broods containing more female young harboured more parasites. Hence, within‐brood variation in host susceptibility to parasites can not only influence the costs of parasitism for individual nestlings, but may also have consequences for the size of parasite populations within nests. If patterns of maternal investment affect the abundance of nest‐dwelling parasites, these interactions may be important for understanding fitness consequences of maternal resource allocation in many vertebrate hosts.     

Not for Parents Only: Begging Calls Allow Nest‐Mate Discrimination in Juvenile Zebra Finches

The benefits of recognition of family members may range from inbreeding avoidance to cooperative and coordinated behaviors within the family group. In birds, recognition of family members has almost exclusively been studied between parents and offspring or within cooperatively breeding societies. Yet, recognition of nest‐mates could be of special importance in recently fledged birds of colonial species by helping in locating the nest, maintaining family group cohesion, or allowing detection of feeding opportunities by recognizing the begging calls nest‐mates produced on the return of a parent. Here we study nest‐mate discrimination based on begging calls in fledglings of domesticated zebra finches (Taeniopygia guttata), a gregarious songbird living in loose colonies in which juveniles may gather in crèches and are fed by parents up to 20 d after fledging. Using playback tests, we show that fledglings called more and spent more time near the loudspeaker in response to the begging calls of their nest‐mates than to the calls of other familiar individuals. Because each fledgling was exposed to the repeated association of the begging calls of its nest‐mates and the subsequent feeding of its parents, this preferential response to the nest‐mates' calls could be a conditioned response to the food reward. Whereas fledglings answered more to male fledgling calls than to female fledgling calls, response to playback was influenced neither by the sex of the subject nor by its brood size. Discriminant function analysis based on acoustic parameters showed that begging calls carried an individual signature as well as a brood signature which might account for such nest‐mate discrimination. Begging signals are major study systems of the evolution of communication in the face of conflicts of interest between signalers and receivers. Our results suggest that eavesdropping and communication networks may be other informative frameworks to understand the design of offspring solicitation signals.     

Offspring desertion and parental care in the Whiskered Tern Chlidonias hybrida

In species with biparental care, a conflict of interest can arise if one mate tries to maximize its own reproductive success at the expense of the other's. One of the mates can desert the brood to accrue a number of benefits to enhance its own fitness, leaving parental care to the remaining parent. This study is the first to describe the desertion pattern in a tern species (Sternidae). We investigated offspring desertion in the Whiskered Tern Chlidonias hybrida, a species with semi‐precocial chicks. Offspring desertion was recorded in 52% of nests prior to fledging (n = 131 nests). Females also deserted during the post‐fledging period. Of the deserters, 97% were females. Desertions started when chicks were 5 days old and no longer required intense brooding. Desertions before fledging did not affect fledging success. Provisioning rates between pair members differed, and females supplied much less food than males. Female provisioning rate affected the chances of nest desertion significantly: daily desertion rates were lower when females supplied more food. After females had deserted, males increased their provisioning rates but compensated for the loss of female care only partly in two‐ and three‐chick broods. Only in small (one‐chick) broods was compensation full. We conclude that male and female Whiskered Terns adopt different reproductive strategies in the population studied here. Females invest much less in parental care than males, providing less food and deserting more frequently. Given the ready availability of food and low predation pressure, benefits appear to accrue to females that desert; selection forces may therefore not be acting against female desertion.     

Should mothers provision their offspring equally? A manipulative field test

Within‐brood variation in offspring size is universal, but its causes are unclear. Theoretical explanations for within‐brood variation commonly invoke bet‐hedging, although alternatives consider the role of sibling competition. Despite abundant theory, empirical manipulations of within‐brood variation in offspring size are rare. Using a field experiment, we investigate the consequences of unequal maternal provisioning for both maternal and offspring fitness in a marine invertebrate. We create experimental broods of siblings with identical mean, but different variance, in offspring size, and different sibling densities. Overall, more‐variable broods had higher mean performance than less‐variable broods, suggesting benefits of unequal provisioning that arise independently of bet‐hedging. Complementarity effects drove these benefits, apparently because offspring‐size variation promotes resource partitioning. We suggest that when siblings compete for the same resources, and offspring size affects niche usage, the production of more‐variable broods can provide greater fitness returns given the same maternal investment; a process unanticipated by the current theory.     

Juvenile Barn Swallows Hirundo rustica L. from late broods start autumn migration younger,fuel less effectively and show lower return rates than juveniles from early broods

Long‐distance migratory passerines may exhibit different post‐fledging rearing times between their early and late broods due to limited time and energy resources. This could affect the migratory fuelling behaviour, condition and fitness of juvenile birds. We analysed long‐term (13‐year) biometric data (body mass, fat score) on Barn Swallows ringed at the nest and later recaptured during their initial stages of autumn migration. The return rates of early‐ and late‐born juveniles after overwintering in Africa were further compared to estimate their fitness. We found that juveniles from late broods started migrating south on average 11 days younger compared with juveniles from early broods. Body masses of nestlings did not differ between early and late broods, but juveniles from late broods were in better condition after leaving the nest, possibly due to a longer period of parental care. Having started migration, juveniles from late broods were, however, less efficient in accumulating energy than juveniles from early broods. A younger departure age, together with less efficient fuelling of late brood juveniles, may partly explain the 39% lower return probability and hence lower fitness value of late‐brood juveniles. Our study is the first to show that juveniles from the early and late broods exhibit different fuelling behaviour and departure strategies (in terms of age and timing), which may be reflected in the fitness value of offspring.     

Male Isaza (Gymnogobius isaza, Gobiidae) prefer large mates: a counterstrategy against brood parasitism by conspecific females

Like many other gobies, male Isaza (Gymnogobius isaza) which are endemic to Lake Biwa, Japan, exclusively care for broods in nests. This goby may have an optimal range of brood size (i.e., an average clutch size of about 2000–3000 eggs) within which they may produce larger numbers of hatching young because much larger broods may be destroyed by fungal infection before hatching. This optimal brood size hypothesis (Takahashi et al. in J Ethol 22:153–159, 2004) predicts that (1) after spawning, both males and females will refuse additional spawning by other gravid females (second females) to keep brood sizes within optimal ranges, (2) larger fish will repel second females more successfully than will smaller fish, and thus, (3) both sexes prefer larger mates. To examine these predictions, we first observed Isaza’s aggressive behaviors in aquaria and investigated whether fish attacked and repelled second females that were introduced after spawning, and, if so, what were the sizes of fish that did so. Large fish, regardless of sex, aggressively prevented second females from entering the nest, but second females larger than the pairs displaced the pair females forcibly and spawned eggs into their clutches. Mate choice experiments showed that males preferred large females. Although females’ choice of large mates was not confirmed, many results may largely coincide with the predictions of the optimal brood size hypothesis. Thus, Isaza males’ choice of large mates will be advantageous for defending against brood parasitism by conspecific females and for achieving optimal clutch size.     

Female mate choice,male-male competition and parental care in the river bullhead,Cottus gobio L. (Pisces,Cottidae)

Male body size was tested for its influence on female mate choice, male-male competition and ability to defend broods in the river bullhead, Cottus gobio L., a polygynous fish with paternal care. Females presented with two potential mates of different sizes significantly preferred to spawn with the larger male. Males smaller than, or 1·5 times longer than, the female were rarely selected as mates. Larger males were more successful in defending their brood from conspecifics, which may explain female preference for them. Unmated large males displaced smaller guarding males from their nests and retained the acquired egg masses. Competition between males for nest sites with eggs can be accounted for by the preference of females for males already guarding eggs: by seizing a nest containing egg masses, a male will increase his chance of being chosen.     

Alloparental care in the sea: Brood parasitism and adoption within and between two species of coral reef Altrichthys damselfish?

The evolution of parental care opens the door for the evolution of brood parasitic strategies that allow individuals to gain the benefits of parental care without paying the costs. Here we provide the first documentation for alloparental care in coral reef fish and we discuss why these patterns may reflect conspecific and interspecific brood parasitism. Species‐specific barcodes revealed the existence of low levels (3.5% of all offspring) of mixed interspecific broods, mostly juvenile Amblyglyphidodon batunai and Pomacentrus smithi damselfish in Altrichthys broods. A separate analysis of conspecific parentage based on microsatellite markers revealed that mixed parentage broods are common in both species, and the genetic patterns are consistent with two different modes of conspecific brood parasitism, although further studies are required to determine the specific mechanisms responsible for these mixed parentage broods. While many broods had offspring from multiple parasites, in many cases a given brood contained only a single foreign offspring, perhaps a consequence of the movement of lone juveniles between nests. In other cases, broods contained large numbers of putative parasitic offspring from the same parents and we propose that these are more likely to be cases where parasitic adults laid a large number of eggs in the host nest than the result of movements of large numbers of offspring from a single brood after hatching. The evidence that these genetic patterns reflect adaptive brood parasitism, as well as possible costs and benefits of parasitism to hosts and parasites, are discussed.     

Avoidance of Filial Cannibalism in the Amphipod Gammarus pulex

Adult animals that cannibalise juvenile conspecifics may gain energy but also risk filial cannibalism, that is, consumption of their own offspring. However, individuals vary in the magnitude of the costs and benefits of cannibalism depending on factors such as their current energy reserves or the probability that they have offspring in the vicinity. They may therefore also vary in the extent to which they participate in cannibalism. This study investigated whether the sex or brooding status of adult amphipods (Gammarus pulex) influenced whether they participated in cannibalism of juveniles. For females carrying embryos within their brood pouch, we also investigated two hypotheses to explain the presence or absence of cannibalistic behaviour by determining whether cannibalism was correlated with factors that might reflect energy demands (body length, brood size), or that might reflect a temporal change in cannibalistic behaviour (corresponding to stage of brood development). All reproductive classes of adults participated in some level of juvenile cannibalism, but females carrying offspring at an advanced stage of development (close to emergence from the brood pouch) consumed significantly fewer juveniles than other groups. Females thus appear to significantly reduce cannibalism of juveniles concurrent with the time when their own eggs are hatching within the brood pouch, prior to the release of their offspring. Because the experiment tested female responses to unfamiliar juveniles, this reflects a temporal change in behaviour rather than a response to phenotypic recognition cues, although additional direct recognition cannot be ruled out. Brooding females with large brood sizes or large body lengths, which might have disproportionately greater energetic demands, were not more likely to cannibalise juveniles. We also noted that juveniles that survived in trials where cannibalism occurred were significantly more likely to be found at the water surface, suggesting a possible adaptation to escape cannibalistic adults. Overall, our results provide evidence that amphipods use indirect temporal cues to avoid filial cannibalism.     

Serial monogamy in the European long-snouted seahorse Hippocampus guttulatus

Seahorses (Hippocampus spp.) are non-sex-role-reversed members of the Syngnathidae family that provide extensive brood care. Previous studies of seahorses have revealed monogamy within a single brood, but their longer term mating system had not been comprehensively evaluated. The parental contribution to 29 wild-born broods of Hippocampus guttulatus, sampled from six Portuguese populations with differing seahorse densities and sex ratios, was assessed using microsatellite DNA markers. To assess the longer term genetic mating system of this species parentage was determined in eleven broods sampled from a captive population over two breeding seasons. Genetic data suggest that this socially polygamous seahorse is serially monogamous across breeding seasons, i.e. monogamous within a season but may switch mates between seasons, and that differing population densities and sex ratios do not affect the mating system.     

Effects of Passive Social Support in Juvenile Greylag Geese (Anser anser): A Study from Fledging to Adulthood

In general, support by social allies may reduce stress, increase an individual's status and facilitate access to resources. In Greylag geese (Anser anser), offspring stay with their parents for an entire year or even longer and thereby enjoy social support in encounters with other flock members. We investigated the influence of spatial distance to one's allies on the outcome of agonistic encounters in a natural flock situation for a total of 12 sibling groups after fledging. In addition, we tested two groups of hand‐raised juvenile geese over a time span of 11/2 yr. Passive (i.e. not interfering) human supporters of different familiarity were placed at a standard distance during food provisionings, which produced a tight flock situation. Success in agonistic interactions increased with decreasing distance to members of their social unit. The hand‐raised juveniles were more successful in agonistic interactions and showed increased feeding rates when accompanied by a familiar human than when alone or with a non‐familiar human. The effect of the presence of familiar humans on success in agonistic encounters significantly decreased with increasing age, while feeding rates remained elevated. The positive effects of social support were particularly evident in females. We conclude that social support has similar effects and functions in the highly social greylag geese as reported for social mammals.     

Effects of mating order and male size on embryo survival in a pipefish

In species that provide parental care, individuals should invest adaptively in their offspring in relation to the pre‐ and post‐zygotic care provided by their partners. In the broad‐nosed pipefish, Syngnathus typhle L., females transfer large, nutrient‐rich eggs into the male brood pouch during mating. The male broods and nourishes the embryos for several weeks before independent juveniles emerge at parturition. Given a choice, females clearly prefer large partners. Yet, females provide protein‐richer eggs when the same individual mates with a smaller than a larger male. In the present study, we allowed each female to mate with one small and one large male, in alternated order. We found a strong effect of female mating order, with larger clutches and higher embryo mortality in first‐ than second‐laid broods, which may suggest that eggs over‐ripen in the ovaries or reflect the negative effects of high embryo density in the brood pouch. In either case, this effect should put constraints on the possibility of a female being selective in mate choice. We also found that small and large males produced embryos of similar size and survival, consistent with the reproductive compensation hypothesis, suggesting that, in this species, larger males provide better nourishment to the embryos than smaller males. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 639–645.     

Sibling competition does not exacerbate inbreeding depression in the burying beetle Nicrophorus vespilloides

Inbreeding results from matings between relatives and can cause a reduction in offspring fitness, known as inbreeding depression. Previous work has shown that a wide range of environmental stresses, such as extreme temperatures, starvation and parasitism, can exacerbate inbreeding depression. It has recently been argued that stresses due to intraspecific competition should have a stronger effect on the severity of inbreeding depression than stresses due to harsh physical conditions. Here, we tested whether an increase in the intensity of sibling competition can exacerbate inbreeding depression in the burying beetle Nicrophorus vespilloides. We used a 2 × 3 factorial design with offspring inbreeding status (outbred or inbred) and brood size (5, 20, or 40 larvae) as the two factors. We found a main effect of inbreeding status, as inbred larvae had lower survival than outbred larvae, and a main effect of brood size, as larvae in large broods had lower survival and mass than larvae in medium‐sized broods. However, there was no effect of the interaction between inbreeding status and brood size, suggesting that sibling competition did not influence the severity of inbreeding depression. Since we focused on sibling competition within homogeneous broods of either inbred or outbred larvae, we cannot rule out possible effects of sibling competition on inbreeding depression in mixed paternity broods comprising of both inbred and outbred offspring. More information on whether and when sibling competition might influence inbreeding depression can help advance our understanding of the causes underlying variation in the severity of inbreeding depression.     

Azure‐winged Magpies Cyanopica cyanus trade off reproductive success and parental care by establishing a size hierarchy among nestlings

It is common in birds that the sizes of nestlings vary greatly when multiple young are produced in one nest. However, the methods used by parents to establish size hierarchy among nestlings and their effect on parental provisioning pattern may differ between species. In the Azure‐winged Magpie Cyanopica cyanus, we explored how and why parents controlled the sizes of nestlings. Asynchronous hatching was the main cause of size hierarchy within the brood, although the laying of larger eggs later in the laying sequence reduced this effect. Parents with asynchronous broods produced more eggs and fledged more nestlings than those with synchronous broods but their brood provisioning rates, food delivery per feeding bout and feeding efficiency did not differ. We performed a cross‐fostering experiment to synchronize some asynchronous broods. Provisioning rates of asynchronous broods were lower than those of synchronized broods, but the daily growth rates and fledging body mass of their nestlings were not different. Our findings indicate that parents of asynchronous broods can achieve higher reproductive success than those of synchronous broods based on the same parental care, and the same reproductive success as those of synchronized broods based on less parental care. It appears that parent birds can better trade off reproductive success and parental care by establishing a size hierarchy among nestlings.