EVOLUTION OF OBLIGATE SIBLICIDE IN BOOBIES. 2: FOOD LIMITATION AND PARENT–OFFSPRING CONFLICT

Proximate limitation on parental food delivery has long been invoked to explain the evolution of single-chick broods of pelagic seabirds such as masked boobies (Sula dactylatra). A second possible proximate limit on brood size is siblicide driven by genetic parent–offspring conflict (POC) over brood size, if siblicidal offspring can reduce brood size to one even if the parents' optimal brood size is greater than one. I tested these two hypotheses by experimentally suppressing obligate siblicide in masked booby broods and comparing breeding parameters of these broods with unmanipulated single-chick control broods. Per capita mortality rate of experimental nestlings was higher than that of controls, but this deficit was more than made up by larger brood size. Parents of experimental broods brought more food to offspring, had higher fledging success, and apparently incurred no additional major short-term cost of reproduction, relative to parents of control broods, thus refuting the food limitation hypothesis. Estimates of inclusive fitness of chicks in experimental broods were higher than were those of control nestlings, a result inconsistent with the POC hypothesis that the siblicidal offspring's optimal brood size is one while the parents' optimum is greater than one. This discrepency between natural brood size and apparent brood size optima might be resolved in several ways: experimental artifacts may give misleading estimates of optimal brood size; experimental and control offspring may have different reproductive values at the time of fledging; nestling masked boobies may face a special frequency-dependent case of POC in which the high risk of sharing a nest with a siblicidal sibling makes invasion of other behavioral genotypes difficult even when offspring and parent inclusive fitnesses are higher from a nonsiblicidal brood of two than from a brood of one.     

Interactions between common buzzard Buteo buteo and goshawk Accipiter gentilis: trade-offs revealed by a field experiment

I examined the behavioural interactions between common buzzard Buteo buteo and goshawk Accipiter gentilis and their effects on buzzard breeding success and brood defence with a two-year field experiment using dummies and playback calls. A priori I showed through an extensive nest site analysis that there is considerable nesting habitat overlap between the two species and hence potential for interspecific competition for prime nesting habitat. Buzzards had a significantly lower breeding success when presented with a goshawk dummy compared to control broods but there was no effect of buzzard dummies on reproductive success. Buzzards failing with their breeding attempt tended to select another nest site while successful buzzards more frequently used the same nest again. Buzzard pairs were less often attacked by common crows Corvus corone while exposed to goshawk dummies compared to buzzard dummies. The decision to desert a nest seems to be a trade-off between predation risk on the one hand and protection against crows on the other. Goshawks proved far more aggressive against an intraspecific dummy than buzzards. Buzzards adjusted their level of brood defence against both intra- and interspecific dummies according to the age of offspring but not offspring number, with an increasing brood defence level with increasing offspring age. Thus the behaviour of buzzards towards goshawks is a result of a complex system of trade-offs between predation risk, competition for prime nesting habitat and protection from crows on which brood value acts as a temporal modifier.     

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.     

Group Mobbing Behaviour and Nest Defence in a Cooperatively Breeding Australian Bird

Cooperatively breeding noisy miners (Manorina melanocephala) are well known in Australia for their persistent and very vocal group mobbing of heterospecifics. Here I investigated the nature of this extraordinary behaviour, in particular its role in nest defence, in a colour banded population of noisy miners in south‐east Queensland, Australia. I focused on two questions. First, did the intensity of mobbing vary according to factors such as the threat to the nest, or the ‘value’ of a clutch? Secondly, what role did group mobbing play in the success of a nest? To answer these questions, I experimentally manipulated the nest defence behaviour by placing one of three stuffed models near active noisy miner nests. The response of noisy miners to intruders was not indiscriminate. However, I found that the number of birds that mobbed a model did not simply reflect the potential threat posed. The response of noisy miners to raptors and other potential nest predators may have reflected their rarity as well as the threat posed. The number of mobbers did not vary with the age or size of a brood. In this study, the fate of nests was independent of the number of mobbers or visitors at nests. Finally, up to 80% of mobbers were never seen to make any other type of contribution to a nest, and many could not be related to the brood that they were ‘defending’. Hence, for some noisy miner ‘helpers’ the benefits that they accrued were probably not wholly dependent on the survival of the broods. I suggest that, in this gregarious species, mobbing behaviour at the nest may be a display of social status or individual quality. This hypothesis warrants further investigation.     

The Defence of First and Second Broods by Great Tit (Parus major) Parents: A Test of Predictive Sociobiology1

We examined the extent to which parental investment, as measured by brood defence, is determined by selection via life history in a short-lived bird, the great tit (Parus major). Great tit parents tending 1st and 2nd broods of the season were used to test five predictions of a cost/benefit model of brood defence based on the species average demography. The benefit was envisaged as the brood's contribution to a parent's fitness, and the cost as the potential loss if the defender dies in the act of defence; this loss was mirrored by Residual Reproductive Value plus a fraction of the brood dying as a consequence of the defender's death. Univariate and multivariate procedures were applied to six measures of defence response in 221 experimentally naive great tit pairs with nestlings. Stimuli consisted of a live raptor and a multi-species taped mobbing chorus (both of which triggered strong alarm), the latter alone, and a novel, visual stimulus inhibiting nest visits for some time. As predicted by the model, response strength and associated risk increased with (1) advancing time in the breeding season, (2) the age of young, and (3) the number of young in 2nd broods. The last finding, being non-trivial, is the strongest and the only unequivocal support for the notion that life-history has moulded parental investment also in a short-lived species; findings (1) and (2) could be alternatively accounted for by simply assuming that parents tailor their defence to environmental conditions permitting a new breeding episode. The model failed to predict the change of defence behaviour between 1st and 2nd broods. Instead, proximate factors coupled to the precise breeding area proved to be of prime importance in determining defence level, thereby falsifying the idea of a rigidly preprogrammed change of response level based on population parameter means. Instead, parents might gauge their defence to quality of young, as also allowed for in the model, or they may sometimes be constrained to display the full defence. As a by-product, the experiments permit us to narrow down the range of selective agents producing the male parent's stronger defence. The sex difference, being prevalent in 1st broods and abating in 2nd broods, indicates that the male, by its defence behaviour, not only invests in its offspring but also in its home range and/or its mate. The latter interpretation is supported most directly by the male's defence behaviour in 1st broods.     

Parental investment and brood defence by made and female great skuas Catharacts skua: the influence of food supply, laying date, body size and body condition

This paper tests the predictions of parental investment theory and other hypotheses relating to variation in brood defence by examining aggression displayed by great skuas Catharacta skua towards intruders within their territories. Aggression serves the function of nest defence; hatching success of adults breeding in Shetland increased with aggression displayed during incubation, and the correlation between aggression and hatching success was apparent in three separate age classes. Adults displayed higher levels of aggression and greater parental investment in reproduction in years of poor food supply. This was not due to an increase in adults' expectations of future benefits of brood defence with increased investment. since hatching success was unaffected by food supply, and breeding success was lower in years of poor food supply. The observed increase in aggression therefore supports parental investment theory. Aggression increased with body condition index (in terms of mass corrected for body size) for females, but decreased with increasing body condition index for males. This probably reflects size-specific differences in the relative benefits of weight and manoeuvrability as means of promoting effective brood defence and reducing the risk of injury to parents. Aggression may also reflect adult quality, with body condition reflecting quality in opposite ways in males and females, as a result of their different roles during the breeding season.     

Nosy neighbours: large broods attract more visitors. A field experiment in the pied flycatcher, <Emphasis Type="Italic">Ficedula hypoleuca</Emphasis>

Life is uncertain. To reduce uncertainty and make adaptive decisions, individuals need to collect information. Individuals often visit the breeding sites of their conspecifics (i.e., “prospect”), likely to assess conspecifics’ reproductive success and to use such information to identify high-quality spots for future breeding. We investigated whether visitation rate by prospectors and success of visited sites are causally linked. We manipulated the reproductive success (enlarged, reduced, and control broods) in a nest-box population of migratory pied flycatchers, Ficedula hypoleuca, in Finland. We measured the visitation rates of prospectors at 87 nest-boxes continuously from manipulation (day 3 after hatching) to fledging. 302 adult pied flycatchers prospected 9194 times on these manipulated nests (at least 78% of detected prospectors were successful breeders). While the number of visitors and visits was not influenced by the relative change in brood size we induced, the resulting absolute brood size predicted the prospecting behaviour: the larger the brood size after manipulation, the more visitors and visits a nest had. The parental provisioning rate at a nest and brood size pre-manipulation did not predict the number of visitors or visits post-manipulation. More visitors, however, inspected early than late nests and broods in good condition. Our study suggests that individuals collect social information when visiting conspecific nests during breeding and provides evidence that large broods attract more visitors than small broods. We discuss the results in light of individual decision-making by animals in their natural environments.     

Evolution of single-chick broods in the Swallow-tailed Gull Creagrus furcatus

Swallow-tailed Gulls lay single-egg clutches, and so raise single-chick broods. As they are pelagic seabirds, this small brood size is expected to relate to proximate food limitation owing to infrequent food deliveries. However, a previous brood doubling experiment detected an 82% increase in fledging success from experimentally doubled broods compared to controls. We repeated the brood doubling experiment, and found that none of 50 enlarged broods produced more than one independent offspring. Control and experimental parents produced fledglings of similar body size, which also had indistinguishable rates of fledging and subsequent survival and reproduction. A variety of parameters estimating survival and breeding costs of reproduction showed no treatment effect. Since two-chick broods yield dramatically higher fledging rates at some times, apparently without excess costs of reproduction, selection on brood size appears to favour a two-chick brood. However, selection may not favour a two-egg clutch if egg production is very costly. Additionally, our estimates of reproductive success do not incorporate the performance of experimental and control offspring as adults, which could differ, since growth of chicks differed slightly by treatment.     

Temporal Variation in Decisions about Parental Care in Bluegill, Lepomis macrochirus

Parental investment theory states that an individual will trade‐off present and future reproductive potential to maximize lifetime reproductive success. Only when parental care is costly in terms of reduced future reproductive potential should individuals be sensitive to changes in the value of current offspring and adjust their care. Here, we examine temporal variation in parental care decision‐making in bluegill (Lepomis macrochirus), in which care is provided by males called ‘parentals’. Previous research has shown that parentals that nest early in the breeding season are in higher energetic condition than those that nest later, and early nesting males appear not to pay an opportunity cost to their care in terms of reduced future reproductive potential. Early nesting males also may have higher paternity in their broods than later nesting males. To examine the parental care decisions made by early and mid‐season nesting parentals, we experimentally reduced males’ perceived paternity by swapping eggs between nests. We found that experimental males that nested early in the breeding season adjusted their brood defence behaviour similarly to control males, which had sham egg swaps performed. Conversely, experimental males that nested mid‐season significantly decreased their brood defence behaviour after the manipulation as compared with control males. Thus, unlike mid‐season nesting males, early nesting males appear relatively insensitive to changes in brood value (paternity), possibly because early nesting males pay little cost in terms of reduced future reproductive potential to providing full care or because these males have a predisposition to high paternity.     

Parental Investment Theory and Nest Defence by Imperial Shags: Effects of Offspring Number,Offspring Age,Laying Date and Parent Sex

Nest defence is a common form of parental care employed by birds to improve the survival of their offspring. Theory predicts that parents should adjust their nest defence according to the value of the brood at stake, defending more intensively broods with high survival and reproductive prospects. We evaluated the influence of offspring number, offspring age, laying date and parent sex on nest‐defence intensity (NDI) of the Imperial Shag Phalacrocorax atriceps, a sexually dimorphic seabird with seasonal decline in offspring survival and very limited renesting potential. We also evaluated whether NDI was correlated within pairs and whether NDI of both members of the pair was correlated with incubation and breeding success. To elicit defensive behaviour, we simulated predation attempts using a Kelp Gull Larus dominicanus model. As predicted by theory, NDI was positively correlated with the number of offspring in the nest and offspring age. NDI during chick rearing was higher than that at early and late incubation, while no differences were found between incubation stages. Contrary to our prediction, we did not find differences in NDI according to laying date. NDI for males was higher than females, while NDI was also positively correlated within pairs. NDI was not statistically related to incubation or breeding success. These results suggest that other factors, such as laying date or parental quality and age, play a much larger role in determining the outcome and productivity of a nesting attempt. Our results provide partial support for parental investment theory; while parental defence increased with brood value according to offspring number and age, parental defence was not related to laying date, a factor strongly affecting offspring survival and recruitment probabilities in this species.     

Mid-contract management of Conservation Reserve Program grasslands provides benefits for ring-necked pheasant nest and brood survival

Conservation Reserve Program (CRP) fields may provide good habitat for nesting and brood-rearing ring-necked pheasants (Phasianus colchicus) during early stages of succession. But, the success of hens in early successional CRP, relative to late successional CRP and other grassland habitats, has yet to be evaluated. The reproductive period is especially critical for populations of pheasants, and CRP's benefits to hens and chicks may decrease as fields age because of loss of vegetative diversity, decrease in vegetation density, and accumulation of residual litter. During 2005–2006, we evaluated spatial and temporal variation in nest and brood survival for radio-marked hen pheasants in areas of northeastern Nebraska where portions of CRP fields had been recently disced and interseeded (DICRP) with legumes. Nests in DICRP tended to have a higher daily survival rate (0.984; 95% CI: 0.957–0.994) than nests in grasslands (including CRP) that were unmanaged (0.951; 95% CI: 0.941–0.972). The probability of 23-day nest success was 0.696 (95% CI: 0.631–0.762) for DICRP and 0.314 (95% CI: 0.240–0.389) for unmanaged grasslands. Daily brood survival rates varied by habitat type, brood age, and date of hatch. The probability of a brood surviving to day 21 was 0.710 (95% CI: 0.610–0.856). Brood survival rates increased with time spent in DICRP and as the brood aged. Survival decreased as broods spent more time in cropland and peaked seasonally with broods that hatched on 15 June. Brood survival probability, to 21 days, would be reduced to 0.36 (95% CI: 0.100–0.701) if broods in our sample had not used DICRP. We combined nest and brood survival in a productivity model that suggested 2,000 hens, in a landscape with no DICRP, would produce 1,826 chicks, whereas the same hens in a landscape of 100% DICRP would produce 5,398 chicks. Production of first-year roosters more than doubled when hens nested in DICRP. Without DICRP, population growth rates of pheasant populations usually declined; with DICRP, populations stabilized with at annual survival rates of 0.3 or greater. The positive response of nest and brood survival to discing and interseeding CRP provides further evidence that CRP fields must be managed to optimize wildlife benefits. © 2012 The Wildlife Society.     

Increased nest defence of upland‐nesting ducks in response to experimentally reduced risk of nest predation

Parent birds should take greater risks defending nests that have a higher probability of success. Given high rates of mammalian nest predation, therefore, parents should risk more for nests in areas with a lower risk of mammalian predation. We tested this hypothesis using nest defence data from over 1300 nests of six species of dabbling ducks studied in an area where predation risk had been reduced through removal of mammalian predators. When predator removal reduced nest predation, the ducks increased risk taking as predicted. Also as predicted, risk taking varied inversely with body size, an index of annual survival, among species. For ducks to vary nest defence in response to variation in predation risk they must be able to assess the risk of nest predation. Because ducks modified nest defence in the breeding season immediately following predator removal, ducks may be able to assess predator abundance indirectly (e.g. by UV reflection from urine) rather than by seeing or interacting directly with the predators.     

Effectiveness of nest defence in the Acadian Flycatcher Empidonax virescens

We used presentations of models to determine the effectiveness of nest defence in the Acadian Flycatcher Empidonax virescens against a nest predator (Blue Jay Cyanocitta cristata ) and a brood parasite (Brown-headed Cowbird Molothrus ater ). Principal components analysis (PCA) of four component variables of nest defence (call rate, swoop rate, closest approach and number of adults) generated a measure of overall nest defence (aggression). We determined effectiveness of defence by looking for correlations between measures of defence and measures of nest success (nest predation and brood parasitism). We also determined whether nest defence increased with clutch size, nestling age and time in the breeding season. Defence against model Brown-headed Cowbirds did not correlate with levels of parasitism, clutch size, age of young or time of breeding. There was, however, a strong, but insignificant, trend for nests with high levels of all measures of defence to suffer less from brood parasitism. Aggression, vocalization rate, closest approach and number of adults defending against models of predatory Blue Jays correlated positively with nesting success during the egg stage but not the nestling stage of the nesting cycle. Aggression, vocalization rate, closest approach correlated with clutch size and age of the brood. These results suggest that nest defence can effectively deter nest predators, but may be less effective against brood parasites. Different behavioural components of nest defence may work at different stages of the nest cycle and against different nest predators. The components of nest defence that correlated with nest success also correlated with clutch value, a result consistent with hypotheses on the evolution of nest defence.     

Brood size and the economy of brood defence: examining Lack's hypothesis in a biparental cichlid fish

Synopsis We tested the explanatory value of two hypotheses reviewed by Lack (1954) in the maintenance of brood size in free-ranging convict cichlidsCichlasoma nigrofasciatum: (1) physiological constraints on egg production, and (2) behavioural constraints imposed by brood defence. Number of free-swimming young in 13 experimental (E) broods was augmented to the upper limit of the size distribution of natural broods (150 young); 18 control (C) broods were handled in the same way but brood size was not changed (mean ± SE = 69.5 ± 11.0). E and C brood sizes were measured at 5 day intervals. At day 20 (just before independence from parental care), 50.3 ± 9.4 (n = 9) young remained in E broods and 30.8 ± 7.8 (n = 8) young remained in C broods (p> 0.05). Offspring number did not differ significantly (p> 0.05) between C and E broods after day 10. Mean growth rate of offspring was significantly lower in E broods than in C broods, perhaps in response to increased density of young in the former. Both the convergence of offspring number in E and C broods and suppression of growth in E broods support a behavioural constraint; that during the first 10 days in which the young are free swimming, two parents are unable to defend large broods as successfully as small broods. A trade-off exists in parental investment between current and future reproduction. Extra-parental investment in current reproduction (eggs) does not result in an increased number of young at independence, therefore a behavioural constraint during brood defence should stabilize the evolution of clutch size.     

Biparental care: short-term manipulation of partner contribution and brood size in the starling, Sturnus vulgaris

In species with biparental care, the evolutionarily stable investmentstrategy depends, among other factors, on the reproductive valueof the brood and on the contribution of one's partner. A shortfallin work rate by one parent should be partially compensated forby its partner, while both parents should invest more effortwhen there are more young. We simultaneously tested these predictionsby attaching weights to one member of pairs of European starlings(Sturnus vulgaris), thus reducing one partner's nest visitationrate, and by daily manipulation of brood sizes (between threeand seven chicks). Both sexes compensated to an equivalent degreefor their partner's lower work rate, while at the same timeadjusting flexibly to the daily brood size changes by increasingvisit rates to larger broods. However, this ability to compensatewas limited in the larger brood sizes, perhaps due to an upperlimit on provisioning rate. Weighted birds and their partnersshowed differences in prey types delivered, relative to controls,taking a lower proportion of leatherjackets. With regard tothe proximate cues affecting parental provisioning rate, beggingnoise levels (automatically recorded by computer from microphonesin the nest-box) increased with brood size, but average lengthof begging bout did not. As expected, visit rates per chickdecreased as brood size increased, and this was reflected inlower weight gains per day by chicks in larger broods. Theseresults agree in general with games theory models but revealimportant departures from our previous work with respect to(1) parents reaching an apparent ceiling to nest visitationrate at high brood sizes and (2) the ability to track thesebrood demands with short-term adjustments.     

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.     

Optimal brood size and its limiting factors in the Rufous Turtle DoveStreptopelia orientalis

Optimal brood size and its limiting factors of the Rufous Turtle Dove,Streptopelia orientalis, were studied at the campus of the University of Tsukuba, Japan, during the breeding season in 1990–92. The dove usually lays two eggs in a nest. I made nests of a brood size of one and three by transferring a hatchling from one nest to the other, and compared their fledging success, factors of breeding failure, weight and tarsus length at fledging, growth rate and nestling period with those of a brood of two. The index of fitness (fledgling weight multiplied by average number of fledglings per nest) was almost the same in broods of two and three. However, the highest variation in fledging weight within the brood and the extension of nestling period were observed in broods of three, which caused the extension of inter-brood interval and consequently the smaller number of broods in the total breeding season. Therefore, broods of three would not have an advantage in producing more offspring than broods of two. Crop milk production had an effect on the growth of nestlings in the early phase of the nestling period, but the rapid growth in the granivorous phase compensated for the growth delay of the smallest nestling in broods of three. Small brood size and a large number of broods in a season would also be more effective under high predation pressure.     

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.     

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.     

The influence of brood loss on nest abandonment decisions in largemouth bass Micropterus salmoides

Largemouth bass Micropterus salmoides broods were experimentally reduced in size to test whether brood size (BS) and simulated brood depredation affect the decision by a male to continue providing care for its brood or to abandon that brood prematurely before its offspring reach independence. The highest ranked of the generalized linear models predicting brood abandonment was based on the number of offspring remaining in a nest following brood devaluation, indicating that parental male fish reassess the value of a brood following perturbation. Paternal M. salmoides were more likely to abandon their broods if initial BS was small before devaluation, and if there was a greater decrease in BS, indicating a threshold for both the amount of brood loss and remaining BS. Larger, older males were also less likely to abandon their brood than smaller, younger conspecifics. These results have broad implications for determining drivers of parental care trade‐offs and how individuals assess the value of a brood.