Mechanism and cost of synchronous hatching

Nest predation is an essential factor affecting bird population density, reproductive ecology and life cycle. However, there are still contradictory results about how nest predation pressure changes with urbanization. One of the reasons for the controversy is that previous studies often use artificial nests to investigate nest predation, but the result of artificial nests is often biased and contrary to that of natural nests. Therefore, it is important to perform nest predation experiments simultaneously with both natural and artificial nests. In this study, we examined the change of nest predation pressure on the Chinese Bulbul (Pycnonotus sinensis) along the urbanization gradient in Hangzhou, China. We detected 148 natural nests of Chinese Bulbuls and continuously monitored them in two continuous breeding seasons from 2012 to 2013. After the breeding season, we placed artificial eggs in natural nests of Chinese Bulbuls to investigate the changes in nest predation pressure and used infrared cameras to record the predators. We then calculated the urbanization synthetic index (USI) as the measure of the degree of urbanization and examined its relationship with nest predation pressure. We found that no matter whether natural nests or artificial eggs were used, the nest predation pressure always decreased significantly with the degree of urbanization. The average height of shrubs within 10 ​m of the nest differed significantly between the predated and unpredated nests, which also affected the risk of nest predation. Our study highlights the importance of using natural and artificial nests simultaneously to conduct nest predation experiments, which can reduce the bias or errors caused by only using artificial nests. We also recommend the use of infrared cameras in future nest predation studies, which has obvious advantages in monitoring and identifying potential predators.

The fiddler crab, Uca beebei, lives in individually defended burrows, in mixed-sex colonies on intertidal mud flats. Avian predation is common, especially of crabs unable to escape into burrows. Mating pairs form in two ways. Females either mate on the surface at their burrow entrance ('surface mating') or leave their own burrow and sequentially enter and leave ('sample') courting males' burrows, before staying in one to mate underground ('burrow mating'). We tested whether perceived predation risk affects the relative frequency of these mating modes. We first observed mating under natural levels of predation during one biweekly, semi-lunar cycle. We then experimentally increased the perceived predation risk by attracting grackles (Quiscalus mexicanus) to each half of the study site in two successive biweekly cycles. In each experimental cycle, crabs were significantly less likely to mate on the side with more birds. Moreover, on the side with elevated predation risk, the number of females leaving burrows to sample was greatly reduced relative to the number of females that surface-mated. Males waved less and built fewer mud pillars, which attract females, when birds were present. We discuss several plausible proximate explanations for these results and the effect of changes in predation regime on sexual selection.

Predation risk is vital in determining habitat quality. But landscapes of fear are complex and dynamic, such as temperate deciduous forests characterized by intermittent, synchronized production of large seed crops (mast peaks). These “resource pulses” elevate the magnitude of predation risk via direct or indirect effects, possibly altering also predator avoidance strategies of a prey. Yet, how animals manage predation risk via informed habitat selection and how this depends on ecological context remains poorly explored. We studied risk-sensitive settlement behavior of wood warbler Phylloscopus sibilatrix. We tested the hypothesis that warblers flexibly avoid predators (Eurasian jays Garrulus glandarius) during habitat selection but that their interactions are mediated by mast seeding, which is known to elevate the overall nest predation risk for warblers through its cascading effects on population dynamics of rodents and generalist predators. We used a playback experiment to manipulate warblers’ perceived risk of nest predation by jays. We replicated the experiment in two consecutive years, followed by high or low seed crop. In the spring following mast trough, risky jay plots featured low abundance, late settlement, and poor pairing success compared to other treatments. In contrast, no response differences were found in the spring following mast seeding. Our results suggest that the way wood warblers adjust habitat choices to jay predation risk is indirectly mediated by mast seeding. This highlights plasticity in predator avoidance during prey habitat selection in complex and dynamic landscapes of fear, and the potential effects of mast seeding on animal behavior via indirect interactions.Significance statementAvoiding places with high nest predation risk during settlement decisions is believed to be widespread in birds. However, the mechanistic understanding of how birds acquire and use information about ambient nest predation risk to fine-tune settlement decisions is still limited. Using a playback experiment, we show that wood warblers eavesdrop on predator calls and avoid settling at sites with high perceived risk of nest predation by Eurasian jays. However, the way warblers adjust habitat choices to the nest predation risk posed by jays appears to be affected by tree mast seeding. This is potentially due to cascading effects of masting on the population dynamics of rodents and generalist predators, creating temporal peaks of elevated nest predation risk for wood warblers. Our study highlights plasticity in predator avoidance during prey settlement behavior in temporally variable, pulsed-resources ecosystems.

Hatchlings of the North American painted turtle (Chrysemys picta) spend their first winter of life inside a shallow, subterranean hibernaculum (the natal nest) where they may be exposed for extended periods to ice and cold. Hatchlings seemingly survive exposure to such conditions by becoming supercooled (i.e., by remaining unfrozen at temperatures below the equilibrium freezing point for body fluids), so we investigated the role of their integument in preventing ice from penetrating into body compartments from surrounding soil. We first showed that hatchlings whose epidermis has been damaged are more likely to be penetrated by growing crystals of ice than are turtles whose cutaneous barrier is intact. We next studied integument from a forelimb by light microscopy and discovered that the basal part of the alpha-keratin layer of the epidermis contains a dense layer of lipid. Skin from the forelimb of other neonatal turtles lacks such a layer of lipid in the epidermis, and these other turtles also are highly susceptible to inoculative freezing. Moreover, epidermis from the neck of hatchling painted turtles lacks the lipid layer, and this region of the skin is readily penetrated by growing crystals of ice. We therefore conclude that the resistance to inoculation imposed by skin on the limbs of hatchling painted turtles results from the presence of lipids in the alpha-keratin layer of the epidermis. Neonates apparently are able to avoid freezing during winter by drawing much of the body inside the shell, leaving only the ice-resistant integument of the limbs exposed to ice in the environment. The combination of behavior and skin morphology enables overwintering hatchlings to exploit an adaptive strategy based on supercooling.

Evolutionary theory predicts that when phenotypic variation arises during development that differently influences the fitness of each sex, selection should favor the maternal ability to match offspring phenotype to the sex that incurs a fitness benefit from that phenotype. In reptiles with temperature-dependent sex determination, the temperatures experienced during incubation can influence numerous phenotypic parameters, including sex. To mimic more naturalistic conditions, this experiment examined how variation in temperature fluctuations affects offspring sex as well as a suite of phenotypic parameters having putative fitness consequences in the Northern Painted Turtle ( Chrysemys picta (Schneider, 1783)). We also characterized variation in natural nest temperatures, including the daily temperature range, related to the vegetation cover surrounding the nest. We found that temperature fluctuations did not affect hatchling morphology, immune response, or behavior, but did significantly affect offspring sex ratios. Thermal profiles of natural nests were related to the amount of surrounding vegetation. Results suggest that nest-site choice by females could influence the sex of their offspring, but we found no evidence that variation in temperature fluctuations adaptively matches offspring sex and phenotype.

Avoidance of predators has long been regarded as a major benefit in colonial breeding. Nevertheless, field and comparative studies have not shown a clear relationship between predation and coloniality. In the present study, we examine the association between aerial egg predation on the Common Tern (Sterna hirundo), and sub-colony size, nest aggregation and reproductive synchrony. Fieldwork was carried out at the Ebro Delta colony (north-western Mediterranean), where Common Terns breed syntopically with potential predators. Sub-colonies placed in a small area were used instead of distant colonies in an attempt to minimize site effects, e.g., the abundance of predators. We used logistic regression with random effect (i.e., sub-colony) to test simultaneously the effect of the studied factors on the risk of predation. In addition, the random effect allowed us to account for the extra-binomial variability due to the potential non-independence of nests of the same sub-colony (clustered observations). Our results support the contention that both breeding in large colonies and in aggregated territories confers protection against aerial predators. In addition, synchrony in relation to the whole colony had no effect on the risk of egg predation. However, birds breeding asynchronously earlier in the season than the average in their own sub-colony were more likely to suffer egg predation than eggs from late asynchronous nesters.

Forming interspecific associations is one of many strategies adopted by primates in order to avoid predation. In addition to improved predator detection and avoidance, benefits of interspecific associations relate to improved foraging efficiency. In this study we tested these two hypotheses explaining associations between the endangered golden-headed lion tamarin, Leontopithecus chrysomelas and the sympatric Wied's marmoset, Callithrix kuhlii. We estimated predation risk by recording the number of encounters between lion tamarins and potential predators in cabruca agroforest (shaded cacao plantation) and in mosaic forest (a mix of cabruca, primary and secondary forest). To evaluate if the association between the two species was related to foraging benefits we recorded the number of associations between the two species when the lion tamarins were eating and when they were not eating. To test if the association occurred to improve predator detection and avoidance, we evaluated if associations between the species were more frequent in areas with higher predation risk and during the part of the day when predation risk is higher. We also compared the number of associations 3 months before birth events and 3 months after, when the lion tamarins are more susceptible to predation. Predation risk, mainly by raptors, was significantly higher in cabruca than in mosaic forest (0.17 and 0.05 encounters with predators per hour of observation, respectively). Associations were significantly more frequent after birth events and during the part of the day when predation risk was also higher (5-6 am until noon). We did not observe any direct evidence of foraging-related advantages of interspecific associations for the lion tamarins. The tamarins did not associate more when they were foraging. Our findings suggest that lion tamarins are more exposed to predation in cabruca than in mosaic forest and associations between lion tamarins and Wied's marmosets are related to predation avoidance.

The effects of predation on Common Redshanks Tringa totanus incubation patterns and behaviour was investigated in 2006 and 2007 at three mainland study sites at the Jadebusen and at one study site on the island Wangerooge, Wadden Sea National Park Lower Saxony (Germany), using temperature data loggers. At these sites, breeding Redshanks were naturally exposed to different predation pressures (as revealed by different hatching success) that varied between 10 and 90% amongst study sites. In areas with a higher predation risk, incubating Redshanks showed a lower nest attendance than in areas where predation risk was low. On the mainland, two behavioural strategies could be distinguished. Some nests were unattended during the night for several hours whereas other nests were incubated for most of the night. Nocturnal absence behaviour was found only on the mainland, and then only in areas easily reached by ground predators. Since ground predators are often active during the night, and dangerous for the incubating birds, this behaviour has probably become established to avoid direct mortality of adult birds rather than avoidance of egg predation. This results from a trade-off between current and future reproduction. By leaving the nest unattended during the night, these birds probably put their own survival before reproduction to maximise their lifetime reproductive success. Since such long incubation recesses may have negative consequences for embryo development, the results of this investigation provide an example of an indirect effect of predation. However, the behavioural trait of nocturnal absence from the nest may directly reduce reproduction and thus fitness of these individuals.

Floating aquatic vegetation and algal blooms are increasing with global warming, potentially reducing UVB exposure and, consequently, vitamin D (vit-D) synthesis in freshwater turtles. Vit-D mediates calcium metabolism and overall health, yet the effects of floating aquatic vegetation on vit-D levels remain unclear, as is whether turtles actively avoid habitats with abundant floating vegetation. Here, we address these questions by quantifying vit-D3 levels in the blood of adult female painted turtles (Chrysemys picta) exposed to high-vegetation (darker/colder) or clear-water (lighter/warmer) treatments for one month outdoors and one month indoors at a single temperature during late summer and early fall. The observed circulating vit-D3 levels resembled those reported for other freshwater turtles, declined over time in both treatments, and were marginally lower under high vegetation after 60 days compared to clear water. However, this difference disappeared after correcting for lymph contamination and multiple comparisons, suggesting that perhaps adult females are robust to the effect of floating vegetation, but whether they were buffered by vit-D3 stores in lipids is unclear. Additionally, in subsequent years, females were exposed to habitat choice experiments and exhibited a strong preference for high floating vegetation over clear water, both as a group (outdoors) and individually (outdoors, and indoors at 21 °C and 26 °C), consistent with known benefits conferred by floating vegetation (food, predator avoidance). While no ill effects of high vegetation nor behavioral avoidance were detected here, longer experiments at different seasons on both sexes and varying ages are warranted before concluding whether painted turtles are truly resilient in their vit-D levels or if, instead, a tradeoff exists between the known benefits of floating vegetation and potential [yet unidentified] detrimental effects (lower dissolved oxygen or vit-D) when vegetation is overgrown for extended periods.

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.

Timing of emergence from the natal nest is a variable trait in the life history of turtles. In theory, hatchling turtles that emerge synchronously, within and among nests, should gain a survival advantage over hatchlings that emerge independently. We examined emergence patterns for seven species of freshwater turtles that use a common nesting area in northern Indiana, USA. Hatchlings of four species emerged from the nest exclusively in late summer or early fall. However, hatchlings of three species usually overwintered in the nest chamber and emerged the following spring. Interspecific and intraspecific emergence from the nest was more synchronous in fall than in spring. Mean date of fall emergence from the nest did not vary among species; however, a species-specific pattern of emergence was observed in spring. Midland Painted Turtles (Chrysemys picta marginata Agassiz, 1857) emerged in late March and early April and, on average, these hatchlings left their nests 2 weeks earlier than Northern Map Turtles (Graptemys geographica (Le Sueur, 1817)) and 4 weeks earlier than Red-eared Sliders (Trachemys scripta elegans (Wied-Neuwied, 1839)). Although hatchlings of C. p. marginata are smaller than those of G. geographica and T. s. elegans, presumably they gain a survival or growth advantage by emerging earlier.

Predation is one of the main drivers of behavioral adaptation in prey. In species with parental care, predation can affect survival of both adults and the offspring under their care. This effect can be further modulated by or combined with other factors that contribute to mating and reproductive success of adults, such as the quality of their breeding site. Using the nest guarding behavior of a treefrog with paternal egg attendance, Kurixalus eiffingeri, we quantify the effects of predation on the behavior of adult frogs and the survival of their offspring. We observed predator avoidance behavior of male frogs guarding empty breeding sites or breeding sites with eggs in response to the introduction of a predatory snake, Dinodon rufozonatum. We hypothesized that male nest guarding behavior would be negatively correlated with predation risk and positively correlated with their clutch sizes. Furthermore, we predicted that predator avoidance in male frog would negatively impact the survivorship of their offspring. We found that breeding site fidelity in male frogs was determined by perceived predation risk and breeding site value, with more males abandoning sites when exposed to high mortality risks (presence of a snake) and when the reproductive benefits (number of offspring) were low. In addition, we found that the willingness of males to return to their breeding sites was positively correlated with their clutch sizes. Abandoning breeding sites, however, can be costly to males, because abandoned eggs had a lower hatching rate. Our findings provide empirical evidence of how the costs and benefits of predation risk and breeding site value can determine the behavior of an amphibian with parental care. When faced with predation risks, the willingness of individuals to maintain their ownership over nest sites can have significant effects on their reproductive success. For species with parental care, nest defense can also be critical for offspring survival. Using an East Asian treefrog with male egg attendance, we hypothesized that males would maximize their reproductive output by increasing investment for nest sites with low predation risks and with more offspring. We found that males are more likely to invest in nest attendance when risks are low and benefits of offspring survival are high. Moreover, we found that the willingness of males to continue parental investment resulted in an increase in survivorship of their offspring. Our results show that perceived risks and benefits to the adults are important factors in determining offspring survival in an amphibian species with parental care.

The house-farmed edible-nest swiftlet (Aerodramus sp.) is monogamous, produces nests from hardened saliva, and uses echolocation to navigate in the darkness. Most previous studies have focused on swiftlet nest building and incubation behaviour, but nestling provisioning behaviour, crucial for the survival of offspring during the breeding season, remains unclear. This study examined the nestling provisioning behaviour of house-farmed swiftlets at a swiftlet house farm in central Peninsular Malaysia. Complete focal video footage of the provisioning phase of swiftlets was analysed in detail. The overall feeding period decreased as the chick grew older. The parents alternately visited their nest 6-14 times a day from dawn to dusk to feed the nestling. The peak of this activity was from 07:00 to 08:00 and between 19:00 and 20:00. Nestling provisioning behaviour differed between pairs. The egg-laying interval of parents with two chicks was longer and affected the development of the second chick since the first chick more successfully competed for food and grew larger in size. Most second chicks with a smaller body size did not survive, because of being either purposely or unintentionally eliminated by the parents or sibling. The smaller-sized second chicks that survived took a longer time to fledge than the first chick; therefore, parents had to increase their foraging effort to ensure the survival of their chick until fledging. These behaviours may be influenced by physiological factors, such as age, experience, individual fitness, and abiotic factors, such as weather and food resources; therefore, different parental strategies may be established during the nestling provisioning phase.

Philopatric behavior has been demonstrated in a wide taxonomic spread of animals. In temperate environments, overwinteredPolisteswasp foundresses often return to their natal nest prior to initiating colony construction. Previous research has shown that these spring foundresses can identify the natal nest in the absence of landmark and gross morphological cues. Hydrocarbons are essential recognition cues forPolistesnest and nestmate discrimination, but cuticular hydrocarbon profiles can become homogenized when foundresses overwinter in mixed colony groups. We examined the hydrocarbon profiles ofPolistes dominulusfoundresses and nests before and after an overwintering period, and found that the hydrocarbon profiles of nests remain unique over time and that this uniqueness is influenced by the original foundresses. Our data raise the possibility that in returning to the natal nest, foundresses reacquire their colony‐specific signature, which may play a role in the formation of cooperative associations.

Effects of artificial light at night and predator cues on foraging and predator avoidance in the keystone inshore mollusc Concholepas concholepas.