Plasticity of parental care under the risk of predation: how much should parents reduce care?

Predation can be an important agent of natural selection shaping parental care behaviours, and can also favour behavioural plasticity. Parent birds often decrease the rate that they visit the nest to provision offspring when perceived risk is high. Yet, the plasticity of such responses may differ among species as a function of either their relative risk of predation, or the mean rate of provisioning. Here, we report parental provisioning responses to experimental increases in the perceived risk of predation. We tested responses of 10 species of bird in north temperate Arizona and subtropical Argentina that differed in their ambient risk of predation. All species decreased provisioning rates in response to the nest predator but not to a control. However, provisioning rates decreased more in species that had greater ambient risk of predation on natural nests. These results support theoretical predictions that the extent of plasticity of a trait that is sensitive to nest predation risk should vary among species in accordance with predation risk.     

Comparative manipulation of predation risk in incubating birds reveals variability in the plasticity of responses

The evolution of different parental care strategies is thoughtto result from variation in trade-offs between the costs andbenefits associated with providing care. However, changingenvironmental conditions can alter such fitness trade-offsand favor plasticity in the type or amount of parental care provided. Avian incubation is a form of parental care whereparents face changing environmental conditions, including variationin the risk of nest predation. Because parental activity candraw attention to the location of the nest, a reduction innest visitation rates is a predicted response to an increased,immediate predation risk. Here, we experimentally increasedthe risk of nest predation using model presentations at nestsof five coexisting species that differ in their ambient levelsof nest predation. We examined whether individuals detect changesin nest predation risk and respond by reducing visitation tothe nest. We also tested whether this behavioral response differsamong species relative to differences in their ambient risk of nest predation. We found that males of all species detectedthe increased predation risk and reduced the rate at whichthey visited the nest to feed incubating females, and the magnitudeof this change was highly correlated with differences in therisk of nest predation across species. Hence, as the vulnerabilityto nest predation increases, males appear more willing to trade the cost of reduced food delivery to the female against thebenefit of reduced predation risk. Our results therefore suggestthat nest predators can have differential effects on parentalbehaviors across species. We discuss how the comparative natureof our results can also provide insight into the evolution of behavioral plasticity.     

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.     

Sexual pigmentation and parental risk‐taking in yellow warblers Setophaga petechia

Adult‐directed predation risk imposes important behavioral constraints on parents and might thus alter relationships between costly sexual ornaments and parental performance. For instance, under low predation risk, highly ornamented individuals might display better parental performance than others, as predicted by ‘good parent’ models of sexual selection. However, under high risk of predation, highly ornamented individuals might abandon parental effort if conspicuous to predators, or if social partners are more willing to take parental risks when paired with highly ornamented mates. We experimentally elevated perceived adult‐directed predation risk near nests to explore how carotenoid‐ and phaeomelanin‐based pigmentation in both sexes relate to parental risk‐taking for offspring in the yellow warbler Setophaga petechia. Compared to other males, males with more intense carotenoid‐based pigmentation maintained higher levels of paternal effort under predation risk at highly concealed nests, but reduced nestling provisioning rate more at exposed nests. Further, when faced with predation risk, females with more phaeomelanin‐based pigmentation reduced nestling provisioning rate less than other females, regardless of nest concealment. Females displayed higher parental effort across treatments when paired to males with more colorful carotenoid pigmentation. However, birds did not reduce parental effort under risk less when paired to a highly ornamented mate, suggesting that predation risk did not accentuate differential allocation. Males did not take fewer parental risks than females. Results indicate that nest concealment modifies parental risk‐taking by males with colorful carotenoid‐based pigmentation, and suggest that female melanin‐based pigmentation may indicate boldness and greater a propensity to take parental risks.     

Parental care of a cowbird host: caught between the costs of egg-removal and nest predation

Avian brood parasites reduce host fitness through the addition of parasitic eggs and the removal of host eggs. Both parasitic egg-addition and host egg-removal may be important sources of selection on host behaviour, creating fitness trade-offs with selection imposed by nest predation. However, the relative costs hosts suffer from egg-addition and host egg-removal and the responses to these costs are largely unstudied. Through experimental manipulations and observations, we demonstrate that increased nest attentiveness by female yellow warblers (Dendroica petechia) reduces the cost of brood parasitism by reducing egg-removal by brown-headed cowbirds (Molothrus ater). However, female attentiveness does not reduce the addition of parasitic eggs. Experimentally parasitized females respond to the threat of egg-removal by increasing nest attentiveness. Increased attentiveness, however, reduces time for females to gather food and requires males to visit the nest more often to feed incubating females. This increased activity in turn increases the risk of nest predation. Thus, brood parasitism (specifically egg-removal) and nest predation produce conflicting selection on incubation strategies, as parasitized hosts are caught between the costs of egg-removal by brood parasites, and the costs of increased nest predation if the female spends more time on the nest to reduce egg-removal.     

Parents, predators, parasites, and the evolution of eggshell colour in open nesting birds

The colourful surface of birds’ eggshells varies dramatically between species, but the selective pressures driving this variation remain poorly understood. We used a large comparative dataset to test several hypotheses proposed to explain the evolution of eggshell colouration. We tested the hypothesis that predation pressure might select for cryptic eggshells by examining the relationship between predation rate and egg colouration. We found that predation rates were positively related to eggshell brightness. The blackmail hypothesis suggests that females lay colourful eggshells to coerce males into providing additional care during incubation to keep colourful eggs covered. According to this hypothesis, conspicuous eggs should be found in situations with high risk of visual detection from predators or brood parasites. In support of this hypothesis, proportional blue-green chroma was positively related to parasitism risk, and eggs with higher proportional blue-green chroma or higher ultraviolet chroma received higher combined parental nest attendance during the incubation period. The sexual signalling hypothesis states that blue-green colour indicates female quality; however, we did not find that blue-green eggshell colour was greater in species where males participate in any form of parental care, and relative male provisioning was unrelated to blue-green eggshell chroma. We found some support for the hypothesis that brood parasitism may select for high inter-clutch variation in eggshell colour to facilitate egg recognition. In our dataset, parasitism risk was negatively related to inter-clutch repeatability of blue-green chroma. Our study highlights the diversity of selection pressures acting on the evolution of eggshell colour in birds and provides suggestions for novel areas of future key research direction.     

Elevating perceived predation risk modifies the relationship between parental effort and song complexity in the song sparrow Melospiza melodia

Adult‐directed predation risk elevates costs of parental care, and may modify relationships between sexually selected ornaments and parental effort by accentuating the tradeoff between survival and parental investment. We assessed multiple hypotheses regarding the relationship between maternal effort, paternal effort, and the sexually selected trait of male song complexity in the song sparrow Melospiza melodia. Further, we explored whether experimentally elevating perceived adult‐directed predation risk near nests affected these relationships. We quantified two dimensions of song complexity: song repertoire size and residual syllable number (the relative number of syllables for a given song repertoire size). Under elevated perceived predation risk, but not in the absence of the predator stimuli, females mated to males with higher residual syllable number displayed higher nestling provisioning rates and performed a greater proportion of nestling provisioning trips. In other words, elevating perceived predation risk induced a pattern of maternal investment consistent with differential allocation. In contrast, under elevated perceived predation risk, only, females performed a lesser proportion of provisioning trips when mated to males with large song repertoire sizes. Further, consistent with the good parent hypothesis, males with large song repertoire sizes displayed lower latencies to return to the nest, independent of the predator stimuli. Results suggest that residual syllable number may reflect some aspect of male genetic quality, such that females are more willing to maintain maternal effort while facing heightened predation risk. On the other hand, females may gain paternal benefits when mated to males with large song repertoires. Our study supports the hypothesis that increased costs of parental care associated with predation risk may induce relationships between sexually selected traits and parental behavior, which may increase the strength of sexual selection. Additionally, results suggest that different aspects of song complexity may fulfill non‐equivalent signaling roles.     

Facultative nest patch shifts in response to nest predation risk in the Brewer’s sparrow: a “win-stay, lose-switch” strategy?

Facultative shifts in nesting habitat selection in response to perceived predation risk may allow animals to increase the survival probability of sessile offspring. Previous studies on this behavioral strategy have primarily focused on single attributes, such as the distance moved or changes in nesting substrate. However, nest site choice often encompasses multiple habitat elements at both the nest site and nest patch scales. We studied the within-season re-nesting strategy of a multi-brooded songbird, the Brewer’s sparrow (Spizella breweri), to determine whether pairs utilized a “win-stay, lose-switch” decision rule with respect to inter-nest distance, nest substrate and/or nest patch characteristics in response to previous nest fate. Pairs moved sequential nest sites slightly farther following nest predation versus success. When inter-nest distance was controlled, however, pairs changed nest patch attributes (shrub height, potential nest shrub density) associated with probability of nest predation to a greater extent following nest predation than success. The strategy appeared to be adaptive; daily nest survival probability for previously depredated pairs increased with greater Euclidian habitat distances between attempts, whereas previously successful pairs were more likely to fledge second attempts when nest sites were similar to those of previous attempts. Our results suggest that nesting birds can use prior information and within-season plasticity in response to nest predation to increase re-nesting success, which may be a critical behavioral strategy within complex nest predator environments. Re-nesting site selection strategies also appeared to integrate multiple habitat components and inter-nest distances. The consideration of such proximate, facultative responses to predation risk may clarify often unexplained variation in habitat preferences and requirements.     

Incubation recess behaviors influence nest survival of Wild Turkeys

In ground nesting upland birds, reproductive activities contribute to elevated predation risk, so females presumably use multiple strategies to ensure nest success. Identification of drivers reducing predation risk has primarily focused on evaluating vegetative conditions at nest sites, but behavioral decisions manifested through movements during incubation may be additional drivers of nest survival. However, our understanding of how movements during incubation impact nest survival is limited for most ground nesting birds. Using GPS data collected from female Eastern Wild Turkeys (n = 206), we evaluated nest survival as it relates to movement behaviors during incubation, including recess frequency, distance traveled during recesses, and habitat selection during recess movements. We identified 9,361 movements off nests and 6,529 recess events based on approximately 62,065 hr of incubation data, and estimated mean nest attentiveness of 84.0%. The numbers of recesses taken daily were variable across females (range: 1?7). Nest survival modeling indicated that increased cumulative distance moved during recesses each day was the primary driver of positive daily nest survival. Our results suggest behavioral decisions are influencing trade‐offs between nest survival and adult female survival during incubation to reduce predation risk, specifically through adjustments to distances traveled during recesses.     

Helper contributions in the cooperatively breeding laughing kookaburra: feeding young is no laughing matter

I studied the contributions of individuals to incubation and nestling feeding in a population of cooperatively breeding laughing kookaburras, Dacelo novaeguineae. In most cooperatively breeding birds where nest success is limited by nestling starvation, related helpers increase the overall level of provisioning to the nest, thus boosting the production of nondescendent kin. However, although partial brood loss is the largest cause of lost productivity in kookaburra nests, additional helpers failed to increase overall provisioning. Instead, all group members, but especially helpers, reduced their feeding contributions as group size increased. Breeders and helpers reduced the size of prey delivered, and helpers also reduced the number of feeding visits. An important benefit of helping in kookaburras may be to allow all group members to reduce their effort. Within groups, contributions to care depended on status, sex, group size and the brood size. Breeding males delivered the most food. Breeding females provisioned less than their partner, but their effort was comparable to that of male helpers. Female helpers contributed the least food. Incubation effort followed similar patterns. The relatedness of helpers to the brood had no impact on their provisioning. Across all group sizes, helpers generally brought larger items to the nest than breeders. Copyright 2000 The Association for the Study of Animal Behaviour.     

Predation risk is unlikely to account for the failure of subordinate speckled warblers Chthonicola sagittata to help at the nest

The predator avoidance hypothesis suggests that the failure of subordinate birds to provision nestlings in communally breeding species is a consequence of increased predation risk. Parents exclude subordinates from the nest area and thus reduce the frequency of predator-attracting visits when the nest is most vulnerable, leading to increased reproductive success. I evaluated this hypothesis for the speckled warbler Chthonicola sagittata , a group-living member of the Pardalotidae in which subordinate males never feed nestlings or fledglings even though they are unrelated to the primary pair, compete for copulations and sometimes sire young in the brood. Parents did not modify provisioning behaviour relative to the risk of nest predation; provisioning rates to 10 d-old nestlings were similar on high and low risk territories. Furthermore, there was no evidence that parents modified the timing of deliveries or adjusted the relative size of deliveries in relation to predation risk. The condition (residual mass) of nestlings differed between high and low risk territories because nestlings on high risk territories had smaller tarsi but similar body mass to those at low risk. Tarsus length was the result of parental phenotype, not modified provisioning behaviour. Given that parents were unresponsive to predation risk, it seems unlikely that predation can account for the failure of subordinates to provision at the nest.     

Females compensate for moult‐associated male nest desertion in Hooded Warblers

Uniparental offspring desertion occurs in a wide variety of avian taxa and usually reflects sexual conflict over parental care. In many species, desertion yields immediate reproductive benefits for deserters if they can re‐mate and breed again during the same nesting season; in such cases desertion may be selectively advantageous even if it significantly reduces the fitness of the current brood. However, in many other species, parents desert late‐season offspring when opportunities to re‐nest are absent. In these cases, any reproductive benefits of desertion are delayed, and desertion is unlikely to be advantageous unless the deserted parent can compensate for the loss of its partner and minimize costs to the current brood. We tested this parental compensation hypothesis in Hooded Warblers Setophaga citrina, a species in which males regularly desert late‐season nestlings and fledglings during moult. Females from deserted nests effectively doubled their provisioning efforts, and nestlings from deserted nests received just as much food, gained mass at the same rate, and were no more likely to die from either complete nest predation or brood reduction as young from biparental nests. The female provisioning response, however, was significantly related to nestling age; females undercompensated for male desertion when the nestlings were young, but overcompensated as nestlings approached fledging age, probably because of time constraints that brooding imposed on females with young nestlings. Overall, our results indicate that female Hooded Warblers completely compensate for male moult‐associated nest desertion, and that deserting males pay no reproductive cost for desertion, at least up to the point of fledging. Along with other studies, our findings support the general conclusion that late‐season offspring desertion is likely to evolve only when parental compensation by the deserted partner can minimize costs to the current brood.     

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.     

Predation by sparrowhawks Accipiter nisus on male and female pied flycatchers Ficedula hypoleuca in relation to their breeding behaviour and foraging

Bright plumage, song display, and aggressive resource defence in males may cause higher predation on males than on females during the breeding season. However, in birds, higher predation on females is sometimes observed. Parental investment may be high in females (egg-laying, incubation and feeding of offspring), which might lead to a high risk of predation. We studied predation by sparrowhawks Accipiter nisus in relation to behaviour in pied flycatchers Ficedula hypoleuca where breeding males are more conspicuous than females in plumage and behaviour. Male pied flycatchers generally occupied more exposed perches than females. Females were more mobile and foraged more than males, especially prior to and during incubation. During the incubation and nestling stages, when predation on the sexes could be directly compared, sparrowhawks took about the same number of male and female pied flycatchers. During incubation, however, females spent about 77% of the day in the nest and were 4.7 times more vulnerable than males per unit of time available (i.e. outside the nest). A comparison with the chaffinch Fringilla coelebs , where hawks took more females than males, indicates that timing of breeding, foraging behaviour and parental roles of males and females affect predation risk.     

Fitness-related consequences of egg mass in nestling house wrens

The fitness-related consequences of egg mass, independent of confounding influences associated with parental quality, remain poorly understood for wild birds in general and for passerines in particular. We performed cross-fostering experiments to test the hypothesis that egg mass, independent of parental quality, is the primary determinant of fitness-related traits in nestling house wrens (Troglodytes aedon), an insectivorous passerine. Nestling mass was significantly correlated with the mass of the eggs from which nestlings hatched early but not late in the nestling period in early-season broods. In contrast, in late-season broods, nestling mass was correlated with egg mass until nestlings achieved asymptotic mass. Neither nestling growth nor survival to nest leaving was related to egg mass in either early- or late-season broods; however, nestlings in late-season broods grew more slowly than did nestlings in early-season broods. We propose that nestling mass and egg mass remained correlated throughout the nestling period in late-season broods because decreased arthropod food resources late in the breeding season constrain parents'' ability to provision nestlings. We conclude that female house wrens in this population trade-off clutch size for greater egg mass to maximise reproductive success in late-season broods.     

Partial host fidelity in nest selection by the shiny cowbird (Molothrus bonariensis), a highly generalist avian brood parasite

Obligate avian brood parasites can be host specialists or host generalists. In turn, individual females within generalist brood parasites may themselves be host specialists or generalists. The shiny cowbird Molothrus bonariensis is an extreme generalist, but little is known about individual female host fidelity. We examined variation in mitochondrial control region sequences from cowbird chicks found in nests of four common Argentinean hosts. Haplotype frequency distributions differed among cowbird chicks from nests of these hosts, primarily because eggs laid in nests of house wrens Troglodytes aedon differed genetically from those laid in nests of the other three hosts (chalk-browed mockingbird Mimus saturninus, brown-and-yellow marshbird Pseudoleistes virescens, and rufous-collared sparrow Zonotrichia capensis). These differences in a maternally inherited marker indicate the presence of a nonrandom laying behaviour in the females of this otherwise generalist brood parasite, which may be guided by choice for nest type, as house wrens nest in cavities whereas the other three species are open cup nesters.     

Nest Predation Avoidance: an Alternative Explanation for Male Incubation in Acrocephalus melanopogon

In the monogamous moustached warbler, male incubation changes from predictably variable (it is dependent on ambient temperature and time of day in April) to high average levels across the day (with no predictor variables in May) as the season progresses. In contrast, females contribute the constant incubation component from April to May. This paper investigates possible explanations for the change in male incubation effort involving changing risks to either (1) embryonic survival within the egg, and/or (2) egg predation. Using egg temperature readouts, the probability of reaching the 25 °C thermal threshold (below which embryonic development ceases) across the season against the probability of predator sightings 0–15 m from the nest was calculated. The results show an inverse relationship between these two risks. During April, male incubation correlates with egg cold stress and changeovers between males and females occur prior to egg cooling below the thermal stress line. During May, the risk of predation increases. The results show increased predator encounter rates from April to May and active nest defense by the incubating parent. Furthermore, high male incubation reduces brood predation. Selection for reduction of the costs of laying replacement clutches (after predation) is suggested given high male incubation and infrequent male-female changeovers during midday, when egg temperatures are highest, with direct benefits to females of increased foraging. Thus, the shift in male incubation across the season may be explained by minimization of changing risk to offspring survival.     

A comparison of life‐history and parental care in temperate and tropical wrens

Comparing closely related species that live in different environments is a powerful way to understand selective pressures that influence life‐history evolution. We examined a suite of life‐history traits and parental care in neotropical buff‐breasted wrens Cantorchilus leucotis and north‐temperate Carolina wrens Thryothorus ludovicianus (Family Troglodytidae), to test hypotheses about life‐history evolution. As expected, buff‐breasted wrens exhibited smaller clutch sizes and higher annual adult survival than Carolina wrens. We found minimal support for the nest predation hypothesis, as nest survival and age‐corrected provisioning rates to whole broods were similar between species, and number of breeding attempts and breeding season length were greater in temperate wrens. Critical predictions of the food limitation hypothesis were not supported; in particular age‐corrected provisioning rates per nestling were higher in the tropical than temperate species. The adult survival and offspring quality hypothesis garnered the most support, as buff‐breasted wrens exhibited greater age‐corrected provisioning rates per nestling, a longer nestling period, longer re‐nesting intervals following nest success, and lower annual fecundity than Carolina wrens. Despite similarly prolonged breeding seasons, reproductive strategies differ between species with buff‐breasted wrens investing considerably in single broods to optimize first‐year survival and Carolina wrens investing in multiple small broods to optimize annual fecundity.     

Cascading costs of reproduction in female house wrens induced to lay larger clutches

In many species, females produce fewer offspring than they are capable of rearing, possibly because increases in current reproductive effort come at the expense of a female's own survival and future reproduction. To test this, we induced female house wrens (Troglodytes aedon) to lay more eggs than they normally would and assessed the potential costs of increasing cumulative investment in the three main components of the avian breeding cycle – egg laying, incubation and nestling provisioning. Females with increased clutch sizes reared more offspring in the first brood than controls, but fledged a lower proportion of nestlings. Moreover, nestlings of experimental females were lighter than those of control females as brood size and prefledging mass were negatively correlated. In second broods of the season, when females were not manipulated, experimental females laid the same number of eggs as controls, but experienced an intraseasonal cost through reduced hatchling survival and a lower number of young fledged. Offspring of control and experimental females were equally likely to recruit to the breeding population, although control females produced more recruits per egg laid. The reproductive success of recruits from broods of experimental and control females did not differ. The manipulation also induced interseasonal costs to future reproduction, as experimental females had lower fecundity than controls when breeding at least 2 years after having their reproductive effort experimentally increased. Finally, females producing the modal clutch size of seven eggs in their first broods had the highest lifetime number of fledglings.