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.

Global population trends in shorebirds: migratory behaviour makes species at risk.

Linking population trends to species' traits is informative for the detection of the most important threatening factors and for assessing the effectiveness of conservation measures. Although some previous studies performed such an analysis at local to continental scales, the global-scale focus is the most relevant for conservation of the entire species. Here we evaluate information on global population trends of shorebirds, a widely distributed and ecologically diversified group, where some species connect different parts of the world by migration, while others are residents. Nowadays, shorebirds face rapid environmental changes caused by various human activities and climate change. Numerous signs of regional population declines have been recently reported in response to these threats. The aim of our study was to test whether breeding and non-breeding habitats, migratory behaviour (migrants vs. residents) and migration distance, breeding latitude, generation time and breeding range size mirror species' global population trends. We found that a majority of shorebird species have declined globally. After accounting for the influence of traits and species taxonomy, linear mixed-effects models showed that populations of migratory shorebirds decreased more than populations of residents. Besides that, declines were more frequent for species breeding at high latitudes of the Northern Hemisphere, but these patterns did not hold after excluding the non-migratory species. Our findings suggest that factors linked to migration, such as habitat loss as well as deterioration at stop-over or wintering sites and a pronounced climate change impact at high latitudes, are possible drivers of the observed worldwide population decreases.

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.

Response to Comment on "Global pattern of nest predation is disrupted by climate change in shorebirds".

Bulla et al dispute our main conclusion that the global pattern of nest predation is disrupted in shorebirds. We disagree with Bulla et al's conclusions and contest the robustness of their outcomes. We reaffirm our results that provide clear evidence that nest predation has increased significantly in shorebirds, especially in the Arctic.

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.

How long do floods throughout the millennium remain in the collective memory?

Is there some kind of historical memory and folk wisdom that ensures that a community remembers about very extreme phenomena, such as catastrophic floods, and learns to establish new settlements in safer locations? We tested a unique set of empirical data on 1293 settlements founded in the course of nine centuries, during which time seven extreme floods occurred. For a period of one generation after each flood, new settlements appeared in safer places. However, respect for floods waned in the second generation and new settlements were established closer to the river. We conclude that flood memory depends on living witnesses, and fades away already within two generations. Historical memory is not sufficient to protect human settlements from the consequences of rare catastrophic floods.

Global pattern of nest predation is disrupted by climate change in shorebirds.

Ongoing climate change is thought to disrupt trophic relationships, with consequences for complex interspecific interactions, yet the effects of climate change on species interactions are poorly understood, and such effects have not been documented at a global scale. Using a single database of 38,191 nests from 237 populations, we found that shorebirds have experienced a worldwide increase in nest predation over the past 70 years. Historically, there existed a latitudinal gradient in nest predation, with the highest rates in the tropics; however, this pattern has been recently reversed in the Northern Hemisphere, most notably in the Arctic. This increased nest predation is consistent with climate-induced shifts in predator-prey relationships.

Equilibrium dynamics of European pre-industrial populations: the evidence of carrying capacity in human agricultural societies.

Human populations tend to grow steadily, because of the ability of people to make innovations, and thus overcome and extend the limits imposed by natural resources. It is therefore questionable whether traditional concepts of population ecology, including environmental carrying capacity, can be applied to human societies. The existence of carrying capacity cannot be simply inferred from population time-series, but it can be indicated by the tendency of populations to return to a previous state after a disturbance. So far only indirect evidence at a coarse-grained scale has indicated the historical existence of human carrying capacity. We analysed unique historical population data on 88 settlements before and after the Thirty Years War (1618-1648), one the longest and most destructive conflicts in European history, which reduced the population of Central Europe by 30-50%. The recovery rate of individual settlements after the war was positively correlated with the extent of the disturbance, so that the population size of the settlements after a period of regeneration was similar to the pre-war situation, indicating an equilibrium population size (i.e. carrying capacity). The carrying capacity of individual settlements was positively determined mostly by the fertility of the soil and the area of the cadastre, and negatively by the number of other settlements in the surroundings. Pre-industrial human population sizes were thus probably controlled by negative density dependence mediated by soil fertility, which could not increase due to limited agricultural technologies.

Mate choice for major histocompatibility complex complementarity in a strictly monogamous bird, the grey partridge (Perdix perdix).

BACKGROUND: Sexual selection has been hypothesised as favouring mate choice resulting in production of viable offspring with genotypes providing high pathogen resistance. Specific pathogen recognition is mediated by genes of the major histocompatibility complex (MHC) encoding proteins fundamental for adaptive immune response in jawed vertebrates. MHC genes may also play a role in odour-based individual recognition and mate choice, aimed at avoiding inbreeding. MHC genes are known to be involved in mate choice in a number of species, with 'good genes' (absolute criteria) and 'complementary genes' (self-referential criteria) being used to explain MHC-based mating. Here, we focus on the effect of morphological traits and variation and genetic similarity between individuals in MHC class IIB (MHCIIB) exon 2 on mating in a free-living population of a monogamous bird, the grey partridge. RESULTS: We found no evidence for absolute mate choice criteria as regards grey partridge MHCIIB genotypes, i.e., number and occurrence of amino acid variants, though red chroma of the spot behind eyes was positively associated with male pairing success. On the other hand, mate choice at MHCIIB was based on relative criteria as females preferentially paired with more dissimilar males having a lower number of shared amino acid variants. This observation supports the 'inbreeding avoidance' and 'complementary genes' hypotheses. CONCLUSIONS: Our study provides one of the first pieces of evidence for MHC-based mate choice for genetic complementarity in a strictly monogamous bird. The statistical approach employed can be recommended for testing mating preferences in cases where availability of potential mates (recorded with an appropriate method such as radio-tracking) shows considerable temporal variation. Additional genetic analyses using neutral markers may detect whether MHC-based mate choice for complementarity emerges as a by-product of general inbreeding avoidance in grey partridges.

Different temperature and cooling patterns at the blunt and sharp egg poles reflect the arrangement of eggs in an avian clutch.

Incubation is an energetically demanding process during which birds apply heat to their eggs to ensure embryonic development. Parent behaviours such as egg turning and exchanging the outer and central eggs in the nest cup affect the amount of heat lost to the environment from individual eggs. Little is known, however, about whether and how egg surface temperature and cooling rates vary among the different areas of an egg and how the arrangement of eggs within the clutch influences heat loss. We performed laboratory (using Japanese quail eggs) and field (with northern lapwing eggs) experiments using infrared imaging to assess the temperature and cooling patterns of heated eggs and clutches. We found that (i) the sharp poles of individual quail eggs warmed to a higher egg surface temperature than did the blunt poles, resulting in faster cooling at the sharp poles compared to the blunt poles; (ii) both quail and lapwing clutches with the sharp poles oriented towards the clutch centre (arranged clutches) maintained higher temperatures over the central part of the clutch than occurred in those clutches where most of the sharp egg poles were oriented towards the exterior (scattered clutches); and (iii) the arranged clutches of both quail and lapwing showed slower cooling rates at both the inner and outer clutch positions than did the respective parts of scattered clutches. Our results demonstrate that egg surface temperature and cooling rates differ between the sharp and blunt poles of the egg and that the orientation of individual eggs within the nest cup can significantly affect cooling of the clutch as a whole. We suggest that birds can arrange their eggs within the nest cup to optimise thermoregulation of the clutch.