Effect Of Brood Size Research Articles

Experimental evidence for a predation cost of begging using active nests and real chicks

The existence of begging costs has recently been the focus of evolutionary studies on parent–offspring conflict. Indirect findings seem to support the existence of a nest predation cost of begging. However, empirical direct evidence of nestling begging attracting the attention of nest predators has been criticized because of methodological problems, mainly the use of artificial nests and playbacks. In order to avoid this problem, we used natural and active nests and real chicks. We utilized Common Blackbird (Turdus merula) nests to experimentally create high and control begging nests by introducing a Great Spotted Cuckoo (Clamator glandarius) or Blackbird chick respectively. We also explored the effect of brood size comparing between Blackbird broods augmented and reduced by one chick. Experimentally parasitized nests were significantly more predated than simply augmented or reduced broods: nest predation for Cuckoo introduced (high begging) nests was more than double that predation for nests of the other two groups. However, we did not find any effect of the brood size. The presence of a Great Spotted Cuckoo chick in a nest also favors an earlier detection of the nest by predators. Parental activity at the nests seems not to be a factor explaining these results as we did not detect significant differences between Cuckoo and Blackbird introduced nests in visit rates. Therefore, our findings support the hypothesis that nestling begging has a severe cost in terms of predation and, hence, this cost may impose an important restriction to the evolution of begging behavior.

An experimental study of fledging success in American Coots (Fulica americana): Effects of brood size, food availability, and hatching asynchrony

ABSTRACT. Brood size and hatching patterns in birds are widely believed to represent adaptations to food availability during the brood-rearing period. Experimental manipulations of brood size and hatching patterns have been widely employed to determine whether parents can raise larger or more synchronous broods, but concurrent manipulations of food abundance to test the supposed causal mechanism have been rare. I studied survival to fledging of 1,060 American Coot (Fulica americana) chicks from 99 broods for which brood size, hatching asynchrony, and food availability had been experimentally manipulated via inter-nest transfers of newly hatched young and provisioning of supplemental food. Survival of color-marked young was measured until 45 days posthatch using Cormack-Jolly-Seber mark—resighting models. Survival of offspring from unsupplemented broods declined linearly with experimental increases in brood size, and this decline was large enough to ameliorate any benefits to parents from larger broods. Ho...

The effects of brood size on growth and steroid hormone concentrations in nestling eastern bluebirds (Sialia sialis)

Birds in multi-nestling broods often experience reduced growth and elevated nutritional and social stress as a result of competition for parental resources. While responses are often species-specific, experimentally increasing brood size has been shown to decrease growth and increase production of both testosterone and corticosterone in nestling passerines. To investigate the relationship between brood size, growth, and steroid hormone production in eastern bluebirds, we cross-fostered nestlings to small, medium, and large broods. Body mass, skeletal size, serum testosterone, and baseline serum corticosterone concentration were measured prior to fledging. Bluebird nestlings raised in large broods weighed less than nestlings in small and medium-sized broods and secreted elevated concentrations of testosterone. Nestling corticosterone concentrations did not vary in response to brood size or body condition, but concentrations were higher in males compared to females. Our results suggest that nestling bluebirds experiencing nutritional and social stress increase testosterone production. Elevated concentrations may enhance begging and competitive abilities in smaller young. Higher concentrations of corticosterone in males may be related to sex-biased provisioning. Further experimental work is needed to ascertain the effects of elevated testosterone concentrations in nestling bluebirds, as well as the causes and consequences of elevated corticosterone for male nestlings.

Sudden weather deterioration but not brood size affects baseline corticosterone levels in nestling Alpine swifts

While evidence is accumulating that stress-induced glucocorticoid responses help organisms to quickly adjust their physiology and behaviour to life-threatening environmental perturbations, the function and the ecological factors inducing variation in baseline glucocorticoid levels remain poorly understood. In this study we investigated the effects of brood size by experimentally manipulating the number of nestlings per brood and the effect of weather condition on baseline corticosterone levels of nestling Alpine swifts ( Apus melba). We also examined the potential negative consequences of an elevation of baseline corticosterone on nestling immunity by correlating corticosterone levels with ectoparasite intensity and the antibody production towards a vaccine. Although nestlings reared in enlarged broods were in poorer condition than nestlings reared in reduced broods, they showed similar baseline corticosterone levels. In contrast, nestling baseline corticosterone levels were higher immediately after cold and rainy episodes with strong winds. Neither nestling infestation rate by ectoparastic flies nor nestling antibody production against a vaccine was correlated with baseline corticosterone levels. Thus, our results suggest that altricial Alpine swift nestlings can quickly modulate baseline corticosterone levels in response to unpredictable variations in meteorological perturbation but not to brood size which may be associated with the degree of sibling competition. Apparently, short-term elevations of baseline corticosterone have no negative effects on nestling immunocompetence.

Effects of brood size on multiple-paternity rates: a case for ‘paternity share’ as an offspring-based estimate

Effects of brood size on multiple-paternity rates : a case for ‘paternity share’ as an offspring-based estimate

Adjustment of brood size and androgen levels in a teleost species with exclusive male parental care

In maximizing reproductive success, individuals face a trade-off between parental care for their current offspring and investing in androgen-dependent sexual traits to produce further offspring. It has thus been proposed that parental effort would suppress androgen levels. Here, we studied parental effort by manipulating brood size in the peacock blenny, Salaria pavo, a littoral fish species in which males show long periods of paternal care. We focused on the effects of brood size on female spawning rate (measured as number of eggs received in the nest by focal males) and androgen levels. We found a positive linear relationship between brood size and the number of eggs received subsequently. Accordingly, spawning rate increased for males with experimentally enlarged broods while it decreased for males in which brood size was reduced. However, over a longer time interval, brood sizes of both treatments returned to the nonmanipulated brood size, suggesting an effect of additional factors such as male quality. Brood size did not show the expected negative relationship with levels of testosterone and 11-ketotestosterone. On the contrary, testosterone levels were positively correlated with brood size. However, in agreement with the prediction, changes in the level of parental care by manipulation of brood size showed an inverse relationship with testosterone levels. As with brood size, given time, testosterone levels tended to return to nonmanipulated levels. Such changes suggest that males adjust their brood size to an individual value through androgen modulation of courtship or other traits influencing female spawning rate.

Paternal aggression towards a brood predator during parental care in wild smallmouth bass is not correlated with circulating testosterone and cortisol concentrations

Male smallmouth bass ( Micropterus dolomieu) provide sole parental care including frequent aggressive actions towards conspecifics and potential brood predators. Failure to defend the brood through continual vigilance results in predation reducing the number of offspring and promoting abandonment by the nesting male. However, little is known about how biochemical and endocrine factors and brood size collectively influence paternal aggression. Behavioral assays were conducted during the egg stage of offspring development by placing a brood predator in a jar on the nest to quantify aggression (number of attacks on the potential brood predator in a minute). To determine the correlates of parental aggression, we temporarily removed fish from their nests and measured circulating levels of testosterone and indicators of the primary (plasma cortisol) and secondary stress response (plasma glucose, Cl −, Na +, K +) from non-lethal blood samples. While the male was removed from the nest, a snorkeler quantified the size of the brood. Brood size was positively correlated with male aggression. The only biochemical correlate of parental aggression was plasma glucose, which also had a positive relationship with brood size. When the effect of brood size was removed, no biochemical or endocrine factors were predictive of male aggression. Hence, brood value appeared to influence parental aggression independent of biochemical or endocrine status. While several-fold individual differences in aggression towards brood predators were noted, the role of androgens and glucocorticoids in mediating these behaviors is currently not well understood.

Sex‐specific effects of altered competition on nestling growth and survival: an experimental manipulation of brood size and sex ratio

1. An increase of competition among adults or nestlings usually negatively affects breeding output. Yet little is known about the differential effects that competition has on the offspring sexes. This could be important because it may influence parental reproductive decisions. 2. In sexual size dimorphic species, two main contradictory mechanisms are proposed regarding sex-specific effects of competition on nestling performance assuming that parents do not feed their chicks differentially: (i) the larger sex requires more resources to grow and is more sensitive to a deterioration of the rearing conditions ('costly sex hypothesis'); (ii) the larger sex has a competitive advantage in intra-brood competition and performs better under adverse conditions ('competitive advantage hypothesis'). 3. In the present study, we manipulated the level of sex-specific sibling competition in a great tit population (Parus major) by altering simultaneously the brood size and the brood sex ratio on two levels: the nest (competition for food among nestlings) and the woodlot where the parents breed (competition for food among adults). We investigated whether altered competition during the nestling phase affected nestling growth traits and survival in the nest and whether the effects differed between males, the larger sex, and females. 4. We found a strong negative and sex-specific effect of experimental brood size on all the nestling traits. In enlarged broods, sexual size dimorphism was smaller which may have resulted from biased mortality towards the less competitive individuals i.e. females of low condition. No effect of brood sex ratio on nestling growth traits was found. 5. Negative brood size effects on nestling traits were stronger in natural high-density areas but we could not confirm this experimentally. 6. Our results did not support the 'costly sex hypothesis' because males did not suffer from higher mortality under harsh conditions. The 'competitive advantage hypothesis' was also not fully supported because females did not suffer more in male-biased broods. 7. We conclude that male nestlings are not likely to be more expensive to raise, yet they have a size-related competitive advantage in large broods, leading to higher mortality of their on average lighter female nest mates.

Conspecific Brood Parasitism and Population Dynamics

Conspecific brood parasitism (CBP), defined as parasitic laying of eggs in a conspecific nest without providing parental care, occurs in insects, fishes, amphibians, and many birds. Numerous factors have been proposed to influence the evolution of CBP, including nest site limitation; effects of brood size, laying order, or parasitic status on offspring survival; randomness of parasitic egg distribution; adult life-history trade-offs; and variation in parental female quality or risk of nest predation. However, few theoretical studies consider multiple possible types of parasitism or the interplay between evolution of parasitism and population dynamics. We review existing theory of CBP and develop a synthetic modeling approach to ask how best-of-a-bad job parasitism, separate-strategies parasitism (in which females either nest or parasitize), and joint-strategies parasitism (in which females can both nest and parasitize) differ in their evolutionary conditions and impacts on population dynamics using an adaptive dynamics framework including multivariate traits. CBP can either stabilize or destabilize population dynamics in different scenarios, and the role of comparable parameters on evolutionarily stable strategy parasitism rate, equilibrium population size, and population stability can differ for the different modes of parasitism.

Corticosterone metabolites in blue tit and pied flycatcher droppings: Effects of brood size, ectoparasites and temperature

The hypothalamic-pituitary-adrenal (HPA) axis of birds induces the secretion of corticosterone (CORT) as a response to different ecological variables. In this study we tested experimentally if manipulations of brood size or ectoparasitism led to subsequent differences in the concentration of excreted CORT metabolites of adult and nestling blue tits ( Cyanistes caeruleus). No significant effect of the manipulation of brood size was detected in adults or nestlings. No significant effect of ectoparasitism was detected in males or nestlings, although females from uninfested nests showed lower concentrations of excreted CORT metabolites. In addition, we analysed if weather conditions had an influence on the concentration of excreted CORT metabolites of blue tits and pied flycatchers ( Ficedula hypoleuca) breeding in the same forest. We detected no effect of weather conditions on adults, but nestlings of both species showed a negative correlation between their excreted CORT metabolites and the average mean temperatures they were subjected to during their growth. This effect was not found in blue tits in a colder year, suggesting that the sensitivity of the HPA axis to ambient temperature may be subjected to interannual variation. Moreover, we found a positive effect of the maximum temperature on the day of sampling on the concentration of CORT metabolites of blue tit nestlings in one of the years. These results suggest that weather conditions may act as environmental stressors to which the HPA axis of blue tit and pied flycatcher nestlings may be sensitive.

Shared or Unshared Parental Care in Overwintering Brent Geese (Branta bernicla hrota)

AbstractWe examine brood size effects on the behaviour of wintering parent and juvenile brent geese (Branta bernicla hrota) to test predictions of shared and unshared parental care models. The behaviour of both parents and offspring appear to be influenced by declining food availability over the winter. Parental vigilance increased with brood size and may be explained by vigilance having functions in addition to antipredator behaviour where the benefits are shared among the brood. There was no increase in parental aggression with brood size and this does not fit the prediction of shared care. Nevertheless, large families are able to monopolize better feeding areas compared with smaller families and large families static feed more but walk feed less than do small families, the former apparently being the preferred mode. The presence of additional young, rather than increasing the amount of parental aggression, seems to enhance the family's competitive ability. Because parents with large broods benefit from enhanced access to resources there is likely to be no additional significant cost in the parental care of larger broods (sensuTrivers 1972).

Delayed behavioral effects of postnatal exposure to corticosterone in the zebra finch ( Taeniopygia guttata)

Early developmental conditions can significantly influence the growth and survival of many animal species. We studied the consequences of exposure to corticosterone (CORT), a stress hormone, during the nestling stage on two behavioral traits (neophobia, social dominance) measured when the birds had reached independence. Nestling zebra finches ( Taeniopygia guttata) were exposed twice daily to exogenous CORT via oral administration for a 12-day period up until fledging. Experimental CORT administration depressed nestling growth rates, confirming results previously obtained in this species. Our data on neophobic behavior revealed a significant interaction between sex and treatment, with CORT-dosed males showing reduced latencies to approach a novel object, while there was little effect of corticosterone treatment on female neophobia. There was no significant effect of age (30 or 50 days), however, there was a non-significant trend towards an interaction between treatment and age, with neophobia increasing with age in the CORT-dosed birds, but decreasing in controls. At 50 days of age previous exposure to corticosterone resulted in reduced success in competitions for a non-food-based resource (a perch) in both sexes. There were no effects of brood size on any behavioral traits measured here, but this may be due to the small range in brood size used. Our results show that elevated levels of stress hormones during postnatal development can have significant effects on important behavioral traits, i.e., neophobia and dominance. Moreover, they confirm the importance of rearing conditions in shaping adult phenotypes.

Effect of brood size and hatching sequence on prefledging mortality of Sandwich terns: why lay two eggs?

The mortality of Sandwich tern Sterna sandvicensis chicks held in enclosures was studied in colonies on Griend, in the Dutch Wadden Sea, from 1992 to 1999, and on Hirsholm, in the Danish Katt- egat, in 1997. Survival of chicks until fledging was 73% for chicks hatching from first-laid eggs or single- egg clutches and 59-64% for partially hatched two-egg clutches, whereas 6% of second hatchlings survived until fledging. Less than 2% of all two-chick broods actually fledged two chicks. Because 18% of the two-egg clutches only hatched one egg, 7% of fledglings of two-egg clutches originated from a sec- ond-laid egg. In nests where both eggs hatched, the number of chicks was usually reduced soon after hatching. Within five days of hatching more than 50% of the second hatchlings died of starvation or were preyed upon. It seems that overproduction commonly occurs in Sandwich terns and that invest- ment in a surplus egg mainly serves as an insurance mechanism. On Griend and Hirsholm, chick pro- ductivity of two-egg clutches was somewhat higher than for one-egg clutches. Undernourishment was an important cause of death, either directly by starva- tion or by selective predation of chicks in poor condition. This, in combination with earlier, studies suggests that Sandwich tern parents on Griend are exposed to severe food stress.

Parental care behavior in the monogamous, sexually dimorphic Madagascar paradise flycatcher: sex differences and the effect of brood size

AbstractI studied the parental care behavior of the Madagascar paradise flycatcher Terpsiphone mutata in northwestern Madagascar. I especially focused on feeding, brooding and vigilance behaviors. Feeding rate did not differ between males and females, but females spent more time at the nest than males. Females dedicated their time to brooding, while males perched on the nest and were vigilant. Both parents changed the feeding rate in relation to brood size, so the feeding rate per nestling was not different among nests of different brood size. Duration of brooding by females increased with decreasing brood size, suggesting that the Royama effect, the pattern of lower feeding rate per nestling in larger broods, did not apply in this study. Males spent more time on vigilance than females. Anti‐predator vigilance by males should be important for nestling survival given the high predation pressure typical of this population. In conclusion, males provide considerable parental care probably to minimize nestling starvation and to avoid nest predation. My results are not consistent with the general pattern of less parental effort by males in monogamous, sexually dimorphic species.

Time constraints and trade-offs among parental care behaviours: effects of brood size, sex and loss of mate

Animals that provide care to their offspring are likely to face time constraints and, consequently, need to trade-off allocations of time among different behavioural activities. Parental allocation of time is often influenced by intrafamilial conflicts including conflicts of interests between parent and offspring and between parents over optimal parental effort. We investigated effects of offspring demand (by manipulating brood size) and loss of mate (by experimental removal of mate) on allocation of time among parental and nonparental behaviours in the burying beetle Nicrophorus orbicollis. With increasing offspring demand, allocation of time to parental care occurred at the cost of nonparental behaviours. Time allocation among parental care behaviours changed with offspring demand. Time spent on care behaviours from which offspring benefit simultaneously did not change with increasing offspring demand. In contrast, time spent on care behaviours that offspring receive individually increased with increasing brood size. This suggests that costs for parents and benefits for offspring differ considerably among parental care behaviours. Removal of the mate affected males and females differently. Widowed males increased their effort, whereas widowed females showed no change in their effort. This result suggests that males and females negotiate their parental effort differently, and costs and benefits of parental care differ considerably between the sexes. In general, our study shows a plastic parental response to mate loss and simultaneous change in offspring demand, indicating that parents negotiate parental efforts while considering offspring demands.

Offspring sex ratio variation in relation to brood size and mortality in a promiscuous species: the Aquatic Warbler Acrocephalus paludicola

We analyse nestling sex ratio variation in the Aquatic Warbler Acrocephalus paludicola to test for predictions from sex allocation theory that the brood sex ratio is close to parity. We also tested Fiala's (1980) prediction that there is no difference in sex ratio between broods affected and not affected by mortality, and whether a shift in primary sex ratios or simple differential mortality by sex underlies that difference. Furthermore, we explore additional analytical possibilities for inferring proximate mechanisms through simulation modelling. In the Aquatic Warbler, which is promiscuous, the overall sex ratio determined by molecular sexing of nestlings at 8–11 days of age did not deviate significantly from parity (proportion of females 0.509), nor did we find any predictive effect of brood size, maternal body mass, fat condition, wing and bill length, laying date, mean daily temperature, and multiple‐male mating. However, extensive simulation suggested that the whole pattern of sex ratio variation is unlikely to arise purely by chance: (1) there is a diverging sex ratio between complete and partial broods, (2) large broods tended to be female‐biased and small broods male‐biased, and (3) low ambient temperature prior to the laying period seemed to increase the proportion of female offspring in complete broods. We conclude that most variation in nestling sex ratio is non‐adaptive in nature, and results from variation in female nestlings mortality dependent on brood size and sex ratio.

The effect of brood size and age on partial filial cannibalism in the scissortail sergeant

Within the size range found in the field for the scissortail sergeant Abudefduf sexfasciatus, there was no correlation between the number of cannibalized eggs and total brood size. Very small broods were fully cannibalized. In a manipulation experiment, males were provided with broods of one of three standardized sizes: at the two extremes and in the middle of the range of naturally occurring broods. Brood size had no effect on partial filial cannibalism, but parental effort increased with increasing brood size. Field correlates and the manipulation experiment showed that the cost of cannibalism in terms of current reproductive success decreased significantly with increasing brood size. Contrary to expectations, there was no relationship between male size and the incidence of cannibalism. No preference for younger eggs was found either from field correlates or from a manipulative experiment in which males were provided with an equal number of young and old eggs at the beginning of the parental phase.

Ectoparasites and reproductive trade-offs in the barn swallow (Hirundo rustica).

Parents are predicted to trade offspring number and quality against the costs of reproduction. In altricial birds, parasites can mediate these costs because intensity of parasitism may increase with parental effort. In addition, parasites may mediate a trade-off between offspring number and quality because nestlings in large broods may have reduced anti-parasite immune defence. In this study, we experimentally analysed the effect of brood size on infestation by an ectoparasitic mite in nests of barn swallows (Hirundo rustica). Nests with an enlarged brood had larger prevalence and intensity of infestation than those with a reduced brood. Importantly, each nestling in enlarged broods was exposed to a larger number of mites, even when measured on a per nestling basis, than in reduced broods. Nestlings in enlarged broods had smaller body mass and T-cell-mediated immune response compared to reduced broods. T-cell-mediated immune response and feather growth were negatively correlated with per nestling intensity of infestation in enlarged but not in reduced broods. The results suggest that nestlings in enlarged broods have depressed immunity leading to larger per nestling mite infestation. Hence, exposure to parasites of offspring and parents increases with brood size, and parasitism can thus mediate trade-offs between reproduction and number and quality of the progeny in the barn swallow.

The presence of eggs in the nest and female choice in common freshwater gobies ( Rhinogobius brunneus )

In many species of fishes with paternal care, females prefer to spawn with males who are already guarding eggs. We studied the effect of egg presence on female mate choice in common freshwater gobies, Rhinogobius brunneus sp. OR. In our tests, females did not prefer males with eggs, suggesting egg presence per se may not act as a cue to attract females. We also examined the effects of brood size on paternal care and offspring survival to look for possible benefits females could obtain when choosing males with eggs. Both fanning by egg-guarding males and egg survival increased with brood size. The presence of neighboring males did have a significantly negative effect on males' parental activity, which subsequently results in a lower level of egg survival than in solitary egg-guarding males. This result provides a partial explanation for the result of eggs not attracting females to mate.

Begging signals more than just short-term need: cryptic effects of brood size in the pied flycatcher ( Ficedula hypoleuca )

The begging of nestling birds is known to reliably signal short-term nutritional need, which is used by parents to adjust rates of food delivery and patterns of food distribution within broods. To test whether begging signals reflect more than just short-term feeding history, we experimentally created 18 "small" (4-nestling) and 18 "large" (8-nestling) broods in the pied flycatcher (Ficedula hypoleuca). Compared to small broods, large broods were provisioned by parents at a greater rate, but at a lower visit rate per nestling and with no obvious differences in load mass per visit. However, lower rates of food mass delivery per nestling in large broods did not result in any measurable reduction in nestling growth (i.e. "long-term need") or in any increase in the begging effort per individual nestling whilst in the nest. Mid-way through the nestling period we also used hand-feeding laboratory trials to assess in more detail individual begging behaviour and digestive performance of the three mid-ranking nestlings from each brood. More food items were required at the start of each trial to satiate nestlings from large broods, but despite this initial control for "short-term need", nestlings from large broods went on to beg at consistently higher rates and at different acoustic frequencies. Large brood nestlings also produced smaller faecal sacs, which were quantitatively different in content but did not differ in frequency. We suggest that different nutritional histories can produce cryptic changes in nestling digestive function, and that these can lead to important differences in begging signals despite controlling for short term need.