Offspring thermal demands and parental brooding efficiency differ for precocial birds living in contrasting climates.

BACKGROUND: Chicks of precocial birds hatch well-developed and can search actively for food but their homeothermy develops gradually during growth. This makes them dependent on heat provided by parents ("brooding"), which is then traded off against other activities, mainly foraging. Although brooding has been documented in many precocial birds, little is known about the differences in the amount and efficiency of brooding care, brooding diel rhythmicity, and impact on the chick's growth, particularly between species living in different climatic conditions. RESULTS: We used multisensory dataloggers to evaluate brooding patterns in two congeneric species inhabiting contrasting climate zones: temperate Northern lapwing (Vanellus vanellus) and desert Red-wattled lapwing (Vanellus indicus). In accordance with our expectation, the adult desert lapwings brooded the chicks slightly less compared to the adult temperate lapwings. However, the desert lapwings brooded their chicks in higher ambient temperatures and less efficiently (i.e. they could not reach the same brooding temperature as the temperate lapwings), which are new and hitherto unknown brooding patterns in precocial birds. In both species, night brooding prevailed even during warm nights, suggesting a general brooding rule among birds. Although the high rates of brooding can reduce the time spent by foraging, we found no negative effect of the high brooding rate on the growth rate in either species. CONCLUSIONS: Our data suggest that the chicks of species breeding in colder climates may reduce their thermal demands, while their parents may increase the efficiency of parental brooding care. More research is however needed to confirm this as a rule across species.

Aggressiveness in a subtropical shorebird's nest defense is adjusted to the predator species and shared by conspecifics.

Aggression is an important component of an animal's defense when protecting offspring from predators. Ground nesting birds use a variety of defense strategies. However, their choice according to situation context is poorly known, especially in nonpasserines and in the subtropics and tropics. The ability to distinguish between differently dangerous predator species and the opportunity to share defense with conspecifics are potentially important but little-studied aspects of nest defense strategy. We experimentally studied the nest defense of Red-Wattled Lapwing in an individually marked population in a desert area near Dubai, UAE. We used three stuffed models representing 1) a predator dangerous both to adults and to nests (a cat), 2) a nest predator (a raven), and 3) a harmless reference model (a moorhen). We confirmed that the lapwings distinguished between predator species (being most aggressive toward the cat, and least aggressive toward the moorhen) and adjusted their defense strategy accordingly. In addition, conspecific visitors play a variety of roles in parents' defense strategy. They can strengthen the parental reaction, or they can assist in distracting a predator. The visitors included not only nesting neighbors but also nonbreeding floaters. Both parents participated in nest defense to a similar extent, regardless of incubation stage and ambient temperature. This study provides new insight into the complexity of the defensive patterns in ground-nesting birds inhabiting a hot environment. Comparative experimental research on a range of environments, with various bird species and predator models, can help us to understand the drivers of these defensive behavioral patterns.

Diel timing of nest predation changes across breeding season in a subtropical shorebird.

Predation is the most common cause of nest failure in birds. While nest predation is relatively well studied in general, our knowledge is unevenly distributed across the globe and taxa, with, for example, limited information on shorebirds breeding in subtropics. Importantly, we know fairly little about the timing of predation within a day. Here, we followed 444 nests of the red-wattled lapwing (Vanellus indicus), a ground-nesting shorebird, for a sum of 7,828 days to estimate a nest predation rate, and continuously monitored 230 of these nests for a sum of 2,779 days to reveal how the timing of predation changes over the day and season in a subtropical desert. We found that 312 nests (70%) hatched, 76 nests (17%) were predated, 23 (5%) failed for other reasons, and 33 (7%) had an unknown fate. Daily predation rate was 0.95% (95%CrI: 0.76% - 1.19%), which for a 30-day long incubation period translates into ~25% (20% - 30%) chance of nest being predated. Such a predation rate is low compared to most other avian species. Predation events (N = 25) were evenly distributed across day and night, with a tendency for increased predation around sunrise, and evenly distributed also across the season, although night predation was more common later in the season, perhaps because predators reduce their activity during daylight to avoid extreme heat. Indeed, nests were never predated when midday ground temperatures exceeded 45℃. Whether the diel activity pattern of resident predators undeniably changes across the breeding season and whether the described predation patterns hold for other populations, species, and geographical regions await future investigations.

Daily Rhythms of Female Self-maintenance Correlate with Predation Risk and Male Nest Attendance in a Biparental Wader.

Parents make tradeoffs between care for offspring and themselves. Such a tradeoff should be reduced in biparental species, when both parents provide parental care. However, in some biparental species, the contribution of one sex varies greatly over time or between pairs. How this variation in parental care influences self-maintenance rhythms is often unclear. In this study, we used continuous video recording to investigate the daily rhythms of sleep and feather preening in incubating females of the Northern Lapwing (Vanellus vanellus), a wader with a highly variable male contribution to incubation. We found that the female's sleep frequency peaked after sunrise and before sunset but was low in the middle of the day and especially during the night. In contrast, preening frequency followed a 24-h rhythm and peaked in the middle of the day. Taken together, incubating females rarely slept or preened during the night, when the predation pressure was highest. Moreover, the sleeping and preening rhythms were modulated by the male contribution to incubation. Females that were paired with more contributing males showed a stronger sleep rhythm but also a weaker preening rhythm. If more incubating males also invest more in nest guarding and deterring daylight predators, their females may afford more sleep on the nest during the day and preen more when they are off the nest. Whether the lack of sleep in females paired with less caregiving males has fitness consequences awaits future investigation.

Parental incubation exchange in a territorial bird species involves sex-specific signalling.

BACKGROUND: Effective communication between sexual partners is essential for successful reproduction. Avian parents with biparental incubation need to know how to negotiate, when and who will incubate, and how to harmonize partner exchange at the nest. Although considerable effort has been dedicated to studies of incubation rhythms, few studies have investigated how behavioural signals serve to tighten cooperation between parents. Moreover, existing studies are almost exclusively restricted to species in which long distances between incubating and non-incubating parents prevent continuous communication during incubation. Thus, the most frequently described parental exchange system is a simple model characterized by the return of the non-incubating parent to the nest itself. Here, we propose more complex parental exchange behaviour in the Northern Lapwing (Vanellus vanellus), a territorial species capable of continuous partner communication during incubation and with a highly variable male contribution to incubation. RESULTS: Northern Lapwing females regularly vocalized shortly before departing from the nest, while males mostly left the nest quietly. Responsiveness of the male to female vocalization, perhaps in combination with her flying away from the nest, helped to synchronize incubation care by increasing the probability of exchange, and also by shortening the exchange gaps. In contrast, a male-to-female exchange gap most often occurred after the male quietly flew away from the nest. The frequency of female vocal signalling was not correlated with the male incubation effort on a between-nest scale, but the highest probability of a female-to-male exchange occurred after vocal signalling by females with the most nest-attentive males. Conversely, lowered effort by females to vocalize in the night was accompanied by lower willingness of males to incubate. CONCLUSIONS: Our results suggest that (1) that the incubating parent can communicate with the non-incubating partner using sex-specific behavioural signals, and this helps to synchronize parental exchange on the nest, (2) this signalling may combine acoustic and visual cues, and (3) the efficiency of this signalling might influence the overall nest attendance. The presumption that the repertoire of behavioural signals during reproduction will be much more complex in territorial species that are capable of continuous communication between the partners during the incubation period should be further tested.

Diversity of incubation rhythms in a facultatively uniparental shorebird - the Northern Lapwing.

In birds, incubation by both parents is a common form of care for eggs. Although the involvement of the two parents may vary dramatically between and within pairs, as well as over the course of the day and breeding season, detailed descriptions of this variation are rare, especially in species with variable male contributions to care. Here, we continuously video-monitored 113 nests of Northern Lapwings Vanellus vanellus to reveal the diversity of incubation rhythms and parental involvement, as well as their daily and seasonal variation. We found great between-nest variation in the overall nest attendance (68-94%; median = 87%) and in how much males attended their nests (0-37%; median = 13%). Notably, the less the males attended their nests, the lower was the overall nest attendance, even though females partially compensated for the males' decrease. Also, despite seasonal environmental trends (e.g. increasing temperature), incubation rhythms changed little over the season and 27-day incubation period. However, as nights shortened with the progressing breeding season, the longest night incubation bout of females shortened too. Importantly, within the 24h-day, nest attendance was highest, incubation bouts longest, exchange gaps shortest and male involvement lowest during the night. Moreover, just after sunrise and before sunset males attended the nest the most. To conclude, we confirm substantial between nest differences in Lapwing male nest attendance, reveal how such differences relates to variation in incubation rhythms, and describe strong circadian incubation rhythms modulated by sunrise and sunset.

Flexible parental care: Uniparental incubation in biparentally incubating shorebirds.

The relative investment of females and males into parental care might depend on the population's adult sex-ratio. For example, all else being equal, males should be the more caring sex if the sex-ratio is male biased. Whether such outcomes are evolutionary fixed (i.e. related to the species' typical sex-ratio) or whether they arise through flexible responses of individuals to the current population sex-ratio remains unclear. Nevertheless, a flexible response might be limited by the evolutionary history of the species, because one sex may have lost the ability to care or because a single parent cannot successfully raise the brood. Here, we demonstrate that after the disappearance of one parent, individuals from 8 out of 15 biparentally incubating shorebird species were able to incubate uniparentally for 1-19 days (median = 3, N = 69). Moreover, their daily incubation rhythm often resembled that of obligatory uniparental shorebird species. Although it has been suggested that in some biparental shorebirds females desert their brood after hatching, we found both sexes incubating uniparentally. Strikingly, in 27% of uniparentally incubated clutches - from 5 species - we documented successful hatching. Our data thus reveal the potential for a flexible switch from biparental to uniparental care.

Unexpected diversity in socially synchronized rhythms of shorebirds.

The behavioural rhythms of organisms are thought to be under strong selection, influenced by the rhythmicity of the environment. Such behavioural rhythms are well studied in isolated individuals under laboratory conditions, but free-living individuals have to temporally synchronize their activities with those of others, including potential mates, competitors, prey and predators. Individuals can temporally segregate their daily activities (for example, prey avoiding predators, subordinates avoiding dominants) or synchronize their activities (for example, group foraging, communal defence, pairs reproducing or caring for offspring). The behavioural rhythms that emerge from such social synchronization and the underlying evolutionary and ecological drivers that shape them remain poorly understood. Here we investigate these rhythms in the context of biparental care, a particularly sensitive phase of social synchronization where pair members potentially compromise their individual rhythms. Using data from 729 nests of 91 populations of 32 biparentally incubating shorebird species, where parents synchronize to achieve continuous coverage of developing eggs, we report remarkable within- and between-species diversity in incubation rhythms. Between species, the median length of one parent's incubation bout varied from 1-19 h, whereas period length-the time in which a parent's probability to incubate cycles once between its highest and lowest value-varied from 6-43 h. The length of incubation bouts was unrelated to variables reflecting energetic demands, but species relying on crypsis (the ability to avoid detection by other animals) had longer incubation bouts than those that are readily visible or who actively protect their nest against predators. Rhythms entrainable to the 24-h light-dark cycle were less prevalent at high latitudes and absent in 18 species. Our results indicate that even under similar environmental conditions and despite 24-h environmental cues, social synchronization can generate far more diverse behavioural rhythms than expected from studies of individuals in captivity. The risk of predation, not the risk of starvation, may be a key factor underlying the diversity in these rhythms.