Adjustment of incubation according to the threat posed: a further signal of enemy recognition in the Blackcap Sylvia atricapilla?

Nest predation and brood parasitism are costly for nest owners, and natural selection should therefore favour the evolution of parental counterdefences. We addressed the question of whether Blackcaps Sylvia atricapilla change their incubation behaviour in response to various nest intruders and whether this adjustment matches the intensity of mobbing exhibited towards these intruders. Near focal nests, we successively exposed a dummy of a brood parasite, nest predator and an innocuous species. After the parents had responded, we removed the dummy and filmed their incubation. The most aggressive response towards the Common Cuckoo Cuculus canorus and high nest attendance after its disappearance indicated recognition of the brood parasite. Low-intensity response to the Jay Garrulus glandarius, together with reduced subsequent parental care, suggested that Blackcaps perceived it either as less deleterious at the egg stage than the Cuckoo or as a danger to themselves. Almost no aggression towards the Turtle Dove Streptopelia turtur, along with the resumption of incubation after its removal, implied that Blackcaps recognised it as harmless. In addition, we found that the level of aggression positively correlated with nest attendance, suggesting a link between the intensity of mobbing and subsequent parental care. Altogether, our results demonstrate that the issue of enemy recognition may be viewed as a complex of both aggressive and post-presentation behaviours.     

Circulating androgens are influenced by parental nest defense in a wild teleost fish

While social interactions influence vertebrate endocrine regulation, the dynamics of regulation in relation to specific behaviors have not been clearly elucidated. In the current study, we investigated whether androgens (testosterone) or glucocorticoids (cortisol) play a functional role in aggressive offspring defense behavior in wild smallmouth bass (Micropterus dolomieu), a teleost fish with sole paternal care. We measured circulating testosterone and cortisol concentrations in plasma samples taken from parental males following a simulated nest intrusion by a common nest predator, the bluegill sunfish (Lepomis macrochirus). To understand whether endocrine regulation changes across the parental care period, we looked both at males guarding fresh eggs and at males guarding hatched embryos. Plasma testosterone levels increased in males subjected to a simulated nest intrusion when compared to sham controls. Testosterone concentrations in males guarding embryos were lower than in males guarding fresh eggs, but circulating testosterone was positively correlated with the level of aggression towards the nest predator at both offspring development stages. However, there was no increase in cortisol levels following a simulated nest intrusion, and no relationship between cortisol and any measured parameter. These results suggest that androgens play an important role in promoting aggressive nest defense behavior in teleost fish.     

Repeatability of Host Female and Male Aggression Towards a Brood Parasite

Current research on behavioural consistency showed that various types of animal behaviour are highly repeatable in the context of mate choice, exploration and parental care, including nest protection. However, the repeatability of aggressive nest defence has not yet been studied in hosts of brood parasites, although host aggression against adult parasites represents a crucial line of antiparasitic defences. Here, we investigated the between‐season repeatability of the great reed warbler (Acrocephalus arundinaceus) aggression towards a stuffed dummy of the brood parasitic common cuckoo (Cuculus canorus). We found that under the relatively stable risk of brood parasitism across breeding seasons, female responses to the cuckoo were highly repeatable, whereas male responses were variable. We suggest that the potential explanation for the observed patterns of female and male behaviours may lie in female's prominent roles in offspring care and nest protection, and in her lower renesting potential in comparison with that of males. However, further studies on the relationship between host aggression and other types of behaviours (host personality) and their fitness consequences under the fluctuating parasitism pressures are required to clarify the adaptive significance of aggressive behaviour observed in hosts of brood parasites.     

Sex‐Specific Parental Care in Response to Predation Risk in the European Roller,Coracias garrulus

Sublethal effects of predation constitute an important part of predation effects, which may modulate prey population and community dynamics. In birds, the risk of nest predation may cause a reduction in parental activity in the care of offspring to reduce the chance of being detected by predators. In addition, parents may modify their parental food allocation preferences within the brood in response to predation risk. Our aim in this study was to evaluate the effects of risk of nest predation on parental care and within‐nest food allocation in the European Roller (Coracias garrulus), an asynchronously hatching bird. We manipulated brood predation risk by placing a snake model near the nests that simulates the most common nest predator in the Mediterranean region. Our results show that males but not females increased their provisioning rate when they were exposed to the model and that despite this, nestlings’ body mass decreased in response to this temporary increase in predation risk. We did not find evidence that parents changed their food allocation strategy towards senior or junior nestlings in their nests in response to predation risk. These results show that the European roller modifies parental care in response to their perception of predation risk in the nest and a sex‐specific sensitivity to the threat, which suggests a different perception of offspring reproductive value by parents. Finally, our results show that changes in parental behaviour in response to nest predation risk might have consequences for nestling fitness prospects.     

Parental defence of offspring: A model and an example

Defence of offspring against predators is an important form of parental investment in many species. We derive a model for the optimal level of parental defence during a predator attack. A higher level of defence increases offspring security, but it also exposes the parent to a higher risk. Other conditions being equal, the model predicts that the optimal level of defence increases with offspring age. This is because the relative difference between parent and offspring in expected future survival decreases with increasing offspring age. Compared with the parent itself, the relative importance of the offspring for parental inclusive fitness therefore increases. The risk that the parent should take in defending offspring therefore increases with its age. The model is applied to fieldfare (Turdus pilaris) nest defence. As predicted, parent fieldfares increase their defence throughout the nest period. The model also predicts the observed decrease in parental defence after the hatching and scattering of a precocial brood of young.     

Increasing the perceived predation risk changes parental care in female but not in male Great Tits Parus major

In birds, little is known about how the presence of predators alters parental food distribution decisions among nestlings. We found that experimentally increasing perceived predation risk changed parental care in female but not in male Great Tits Parus major. Females fed the lightest and average nestlings at similar rates under control conditions when predation risk was not manipulated but ignored the lightest nestling under increased perceived predation risk. Moreover, females reduced the duration of nest visits greatly after encountering a model predator, suggesting that the perception of predators may facilitate brood reduction mechanisms.     

Causes and consequences of voluntary anorexia during the parental care period of wild male smallmouth bass (Micropterus dolomieu)

By definition, parental care behaviors increase offspring survival, and individual fitness, at some cost to the parent. In smallmouth bass (Micropterus dolomieu), parental males provide sole care for the developing brood that includes an increase in activity during brood defense and decreased foraging resulting in a decline in endogenous energy reserves. No mechanisms have been proposed for cessation of voluntary foraging, though regulation of appetite hormones such as ghrelin have been documented to affect feeding behavior in other fishes. We documented baseline fluctuations in plasma ghrelin concentrations across parental care. Plasma ghrelin concentrations were lowest during the early stages of parental care before increasing as the brood developed to independence. Additionally, we performed an intervention experiment whereby plasma ghrelin levels were artificially increased through an injection of rodent ghrelin at the onset of parental care. Despite measuring a significant increase in plasma ghrelin approximately 1 week after injection, we noted no differences in plasma-borne indicators of recent foraging activity indicating that voluntary anorexia is possibly reinforced by receptor insensitivity to appetite hormones. Finally, we assessed the ultimate consequences of foraging during parental care by feeding fish to satiation and measuring post-prandial changes in swimming performance and aggression. Fish fed to satiation showed significant decreases in burst swimming ability and aggressiveness towards potential brood predators. Voluntary anorexia during smallmouth bass parental care is an adaptive behavior that avoids potentially deleterious declines in swimming performance and aggression apparently through a modulation of production and reception of appetite hormones including ghrelin.     

Parental care and the ontogeny of predator-avoidance in two species of centrarchid fish

Stamps (1978) suggested that the type and extent of parental care can influence the behavioural ontogeny of the offspring. Largemouth bass (Micropterus salmoides) males guard the fry for up to a month after they leave the nest while rock bass (Ambloplites rupestris) males guard the fry only until they leave the nest. In laboratory experiments, naive largemouth fry displayed reduced predator-avoidance responses to large and small predators during their first 3 weeks of free-swimming. Rock bass fry, on the other hand, avoided predators throughout the study period. From 6 weeks (largemouth) and 5 weeks (rock bass) of age the fry showed a significantly greater response to the large predator than the small predator. The responses of laboratory-reared largemouth bass fry to the small predator changed from avoidance to association as the fry grew. The size of the fry relative to predator size was found to be important.     

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.     

Nutritional condition and physiology of paternal care in two congeneric species of black bass (<Emphasis Type="Italic">Micropterus</Emphasis> spp.) relative to stage of offspring development

Parental care requires a complex integration of physiology and behaviour, yet little is known about the physiological and energetic consequences or correlates of these behaviours. Using two species of male black bass (smallmouth bass, Micropterus dolomieu; largemouth bass, M. salmoides) as a model, the focus of this study was to determine the biochemical and hematological indicators of change in nutritional status and potential for chronic stress. This was accomplished by randomly sampling individuals at four stages across parental care. Additionally, a subset of individuals was repeatedly sampled at three brood development stages to track changes in biochemical factors within the individual. Though there were changes in physiological factors across parental care in randomly sampled fish of both species (declines in plasma glucose in largemouth bass; decreases in hematocrit and plasma chloride in smallmouth bass), repeated sampling of individuals was determined to be a more appropriate sampling technique due to natural variability in biochemical factors among individual fish. Repeated sampling of smallmouth bass did not adversely influence physiological metrics or brood abandonment. However, there were higher incidences of nest abandonment in repeatedly sampled largemouth bass. Amongst the repeatedly sampled smallmouth bass, nutritional indicators such as plasma triglyceride levels decreased indicating individual fasting across the majority of parental care. Increases in plasma calcium and magnesium towards the end of care indicated that feeding most likely resumed when the brood was close to independence after ~3 weeks of care. Lastly, several indicators of chronic stress, such as plasma glucose and chloride levels, increased throughout the parental care period. These sublethal stressors are indicative of decreasing body condition associated with prolonged activity and fasting which may have marked impacts on the ability of an individual to continue parental care for the current brood and impact subsequent individual fitness. Further research into the mechanistic relationships between behaviour, physiology, and energetics during the parental care period will provide a better understanding of the decisions by individuals facing multiple trade-offs that ultimately lead to differences in individual fitness.     

Tidal cycling and parental behavior of the cichlid fish,Biotodoma cupido

Synopsis Parental behavior of the substrate-brooding cichlid, Biotodoma cupido, was studied in a small creek entering the lower Essequibo River, Guyana, where the freshwaters are affected by semi-diurnal tides. Physico-chemical variables of the tidal cycle were associated with the parental behavior of B. cupido. During late ebb and early flood tides, while off-spring were nest dependent, parents displayed intense aggression toward brood predators, mainly characins. At low tide, when the concentration of dissolved oxygen decreased to about 4 mg 1^–1 and that of free carbon dioxide increased to 28 mg 1^–1, parents entered a state of somnolence and brood predators vanished. Early flood tide brought an immediate and dramatic reversal of hypoxic and hypercarbic conditions and an associated renewal of aggressive and predatory activity. At very low tide, parents orally transferred the brood to a secondary nest depression in deeper water. The significance of water-level fluctuation to the evolution of this behavior, as well as that of parent-brood itineracy and the related phenomena of oral incubation and movable nests, is discussed.     

Two Sides of the Same Coin? Consistency in Aggression to Conspecifics and Predators in a Female Songbird

Different forms of aggression have traditionally been treated separately according to function or context (e.g. aggression towards a conspecific versus a predator). However, recent work on individual consistency in behavior predicts that different forms of aggression may be correlated across contexts, suggesting a lack of independence. For nesting birds, aggression towards both conspecifics and nest predators can affect reproductive success, yet the relationship between these behaviors, especially in females, is not known. Here we examine free-living female dark-eyed juncos (Junco hyemalis) and compare their aggressive responses towards three types of simulated intruders near the nest: a same-sex conspecific, an opposite-sex conspecific, and a nest predator. We also examine differences in the strength of response that might relate to the immediacy of the perceived threat the intruder poses for the female or her offspring. We found greater aggression directed towards a predator than a same-sex intruder, and towards a same-sex than an opposite-sex intruder, consistent with a predator being a more immediate threat than a same-sex intruder, followed by an opposite-sex intruder. We also found positive relationships across individuals between responses to a same-sex intruder and a simulated predator, and between responses to a same-sex and an opposite-sex intruder, indicating that individual females are consistent in their relative level of aggression across contexts. If correlated behaviors are mediated by related mechanisms, then different forms of aggression may be expressions of the same behavioral tendency and constrained from evolving independently.     

Paternity, parental behavior and circulating steroid hormone concentrations in nest-tending male bluegill

Like many teleost fishes, bluegill (Lepomis macrochirus) are characterized by sole male parental care of offspring. In addition, bluegill parental males experience cuckoldry by specialized parasitic male morphs. This cuckoldry has previously been shown to influence the expression of parental care behavior. To better understand some of the proximate mechanisms mediating parental behavior, we examined the relationships between circulating steroid hormones, paternity, and parental behavior during the egg and fry stages of care in parentals that spawned during the first third of the breeding season. During the egg stage of care, we found that males with higher paternity had lower levels of testosterone, but there was no relationship between paternity and either 11-ketotestosterone or cortisol. There also was no relationship between the hormones and care behavior comprising fanning of the eggs, nest rim circles, chases of brood predators, or pecking at the eggs (indicative of egg cannibalism), except for a negative relationship between cortisol and pecking behavior. During the fry stage of care, we conversely found that males with higher paternity had higher levels of testosterone and 11-ketotestosterone. There also was a negative relationship between the concentrations of these two androgens and the defensive behavior of males when exposed to a potential brood predator (a pumpkinseed, Lepomis gibbosus). We discuss these results in relation to previous work in fishes and other vertebrate taxa. Overall, our data suggest a complex relationship between circulating steroid hormone levels, paternity and parental behavior.     

The effect of partial brood loss on male desertion in a cichlid fish: an experimental test

There is little experimental evidence testing whether currentbrood size and past brood mortality influence mate desertion.In the cichlid Aequidens coeruleopunctatus both parents initiallydefend offspring. In a field study, all experimental broods,irrespective of initial brood size (222.9 ± 60.4, mean± SD), were manipulated to a size of 100 fry. Neitherthe duration nor investment of females in parental care differed between control and brood reduced pairs, even though care seemedcostly. On average, females lost 5.1 ± 4.8% of initialweight while guarding a brood until independence. In contrast,males with experimentally reduced broods guarded fry for significantlyfewer days before deserting their mate than did males fromcontrol pairs with natural-sized broods (20.5 ± 7.5 vs. 14.2 ± 6.2 days). In at least 20% of cases (n = 9/45),the deserting male immediately mated with another female. Maleswith experimentally reduced broods also spent less time guardingfry before deserting and attacked fewer brood predators thandid males with control broods. For broods manipulated to have100 fry, there was a significant negative relationship betweenthe days until male desertion and the proportion of the initialbrood removed. This indicates that male assessment of the futuresuccess of the current brood (hence its reproductive value)is based on past mortality and/or that there is variation amongmales in the expected size of future broods. Both current broodsize and brood size relative to initial brood size are thereforepredictors of male, but not female, parental behavior and matedesertion. Female care may be unaffected by brood reductiondue to limited breeding opportunities and partial compensationfor reduced male care.     

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.     

Introduced predator elicits deficient brood defence behaviour in a crater lake fish

Introduced species represent one of the most serious global threats to biodiversity. In this field-based study, we assessed behavioural responses of brood tending cichlid fish to an invasive predator of their offspring. This was achieved by comparing parental defence responses of the endangered arrow cichlid (Amphilophus zaliosus), a fish species endemic to the crater lake Apoyo in Nicaragua, towards the bigmouth sleeper (Gobiomorus dormitor), a formidable predator of cichlid fry, and all other potential fish predators of offspring. The bigmouth sleeper was recently introduced into Apoyo but naturally co-exists with cichlids in a few other Nicaraguan lakes. Arrow cichlid parents allowed bigmouth sleepers to advance much closer to their fry than other predators before initiating aggressive brood defence behaviours. Interestingly, parents of a very closely related species, A. sagittae, which has coevolved with bigmouth sleepers in crater lake Xiloá, reacted to approaching bigmouth sleepers at comparable distances as to other predators of cichlid fry. These results provide a novel demonstration of the specific mechanism (i.e. naive parental behaviour) by which invasive predators may negatively affect species that lack the adequate behavioural repertoire.     

Is female investment in defence correlated with breeding success in Hen Harriers Circus cyaneus

CapsuleThe intensity of parental defence, irrespective of the value of offspring, may be one of the proximate causes of breeding success, reflecting the quality of breeders.

Aims To test whether female Hen Harrier investment in defence towards a human predator influences breeding success.

Methods Generalized linear mixed models were used, with nest content, date of visit, the interaction (nest content*date), breeding success (number of fledglings/clutch size) and presence or absence of male as explanatory variables. ‘Female’ was included as a random variable in the model. Alarm call rate by a female Hen Harrier during the first three minutes after my arrival at a nest was used as a dependent variable (as an indicator of parental investment in defence towards a human predator).

Results Females were present and alarmed in 100% of visits whereas males were present in 37% of visits. The individual variations in female investment in defence, after controlling for the principal determinants of nest defence, were significantly correlated with breeding success of each pair.

Conclusion The relationship between female investment in nest defence and reproductive success can be explained through differences in female quality. Good quality females, as measured by the level of investment in nest defence, probably also invested more in other breeding aspects not controlled in this study obtaining higher reproductive success.     

Spawning and Brood Defense of Smallmouth Bass Under the Process of Invasion into a Novel Habitat

Smallmouth bass, Micropterus dolomieu, known as one of the world's most disastrous invasive species, was introduced into Lake Aoki, central Japan in the last 10 years and is of concern for the conservation of the native biodiversity. We investigated spawning and brood defense under novel conditions to devise measures to eradicate a local population of this species. Compared to their native habitat, the size distribution of nesting males was less skewed to the right in this invasive population but rather a skewed normal distribution probably due to prevention of younger cohorts entering the nesting stock. The number of offspring deposited in nests was not positively related to the size of the nesting male. These observations imply that a shortage of suitable nest sites leads to an escalation in male–male competition, followed by a relaxation of female mate choice based on male size. Further, when presented with a model of a potential predator of the offspring, nesting males reacted individually and the intensity of site tenacity was independent of the male size. We propose that the removal of cover accompanied by the use of native predators of young bass can effectively decelerate the expansion of invasive smallmouth bass.     

Hissing females of great tits (Parus major) have lower breeding success than non‐hissing individuals

Antipredator strategies vary remarkably between individuals within populations. Parents tend to take greater risks when brood value is increased. Moreover, individuals consistently differ in a whole suite of correlated behaviours that may cause distinctive responses to predators. It is likely that individual differences in antipredator behaviour may co‐vary with proxies for fitness such as reproductive success. We used a 4‐year data from wild great tits (Parus major) to test whether passive and active antipredator strategies (females with no response vs. those giving hissing calls towards a nest predator) during the incubation stage can reflect variation in breeding success. Although clutch size did not depend on hissing behaviour, the number of surviving offspring from eggs and neonates to fledglings was higher for non‐hissing than hissing birds. We conclude that females with distinct antipredator strategies can prioritize different fitness components.     

Family dynamics through time: brood reduction followed by parental compensation with aggression and favouritism

Parental food allocation in birds has long been a focal point for life history and parent–offspring conflict theories. In asynchronously hatching species, parents are thought to either adjust brood size through death of marginal offspring (brood reduction), or feed the disadvantaged chicks to reduce the competitive hierarchy (parental compensation). Here, we show that parent American coots (Fulica americana) practice both strategies by switching from brood reduction to compensation across time. Late‐hatching chicks suffer higher mortality only for the first few days after hatching. Later, parents begin to exhibit parental aggression towards older chicks and each parent favours a single chick, both of which are typically the youngest of the surviving offspring. The late‐hatched survivors can equal or exceed their older siblings in size prior to independence. A mixed allocation strategy allows parents to compensate for the costs of competitive hierarchies while gaining the benefits of hatching asynchrony.