Heritability of parental effort in a passerine bird

Abstract The study of the evolution of parental care is central to our understanding of social systems, sexual selection, and interindividual conflict, yet we know virtually nothing about the genetic architecture of parental care traits in natural populations. In this paper, we use data from a long term field study of a passerine bird, the long-tailed tit ( Aegithalos caudatus ), to examine the heritability of the rate at which parents feed offspring. This measure of effort is positively related to offspring survival, is repeatable within individuals, and does not appear to be confounded by environmental effects. Using both parent-offspring regression, and an animal model approach, with a pedigree derived from ringing data, we show that our measure of effort has a significant heritable component.     

Dispersal of sibling coalitions promotes helping among immigrants in a cooperatively breeding bird

Kin selection is a major force in social evolution, but dispersal is often assumed to reduce its impact by diluting kinship. In most cooperatively breeding vertebrates, in which more than two individuals care for young, juveniles delay dispersal and become helpers in family groups. In long-tailed tits (Aegithalos caudatus), however, offspring disperse to breed and helpers are failed breeders that preferentially aid kin. Helping also occurs among immigrants, but their origins are unknown and cooperation in these cases is poorly understood. Here, we combine long-term demographic and genetic data from our study population to investigate immigration and helping in this species. We first used a novel application of parentage analysis to discriminate between immigrants and unknown philopatric recruits. We then cross-checked sibship reconstruction with pairwise relatedness estimates to show that immigrants disperse in sibling coalitions and helping among them is kin biased. These results indicate that dispersal need not preclude sociality, and dispersal of kin coalitions may help maintain kin-selected cooperation in the absence of delayed dispersal.     

Social genetic and social environment effects on parental and helper care in a cooperatively breeding bird

Phenotypes expressed in a social context are not only a function of the individual, but can also be shaped by the phenotypes of social partners. These social effects may play a major role in the evolution of cooperative breeding if social partners differ in the quality of care they provide and if individual carers adjust their effort in relation to that of other carers. When applying social effects models to wild study systems, it is also important to explore sources of individual plasticity that could masquerade as social effects. We studied offspring provisioning rates of parents and helpers in a wild population of long-tailed tits Aegithalos caudatus using a quantitative genetic framework to identify these social effects and partition them into genetic, permanent environment and current environment components. Controlling for other effects, individuals were consistent in their provisioning effort at a given nest, but adjusted their effort based on who was in their social group, indicating the presence of social effects. However, these social effects differed between years and social contexts, indicating a current environment effect, rather than indicating a genetic or permanent environment effect. While this study reveals the importance of examining environmental and genetic sources of social effects, the framework we present is entirely general, enabling a greater understanding of potentially important social effects within any ecological population.     

Severe recent decrease of adult body mass in a declining insectivorous bird population

Migratory bird species that feed on air-borne insects are experiencing widespread regional declines, but these remain poorly understood. Agricultural intensification in the breeding range is often regarded as one of the main drivers of these declines. Here, we tested the hypothesis that body mass in breeding individuals should reflect habitat quality in an aerial insectivore, the tree swallow (Tachycineta bicolor), along a gradient of agricultural intensity. Our dataset was collected over 7 years (2005–2011) and included 2918 swallow captures and 1483 broods. Analyses revealed a substantial decline of the population over the course of the study (−19% occupancy rate), mirrored by decreasing body mass. This trend was especially severe in females, representing a total loss of 8% of their mass. Reproductive success was negatively influenced by intensive agriculture, but did not decrease over time. Interestingly, variation in body mass was independent of breeding habitat quality, leading us to suggest that this decline in body mass may result from carry-over effects from non-breeding areas and affect population dynamics through reduced survival. This work contributes to the growing body of evidence suggesting that declines in migratory aerial insectivores are driven by multiple, complex factors requiring better knowledge of year-round habitat use.     

Factors affecting offspring survival and development in a cooperative bird: social, maternal and environmental effects

1. In many noncooperative vertebrates, maternal effects commonly influence offspring survival and development. In cooperative vertebrates, where multiple adults help to raise young from a single brood, social effects may reduce or replace maternal effects on offspring. 2. Factors affecting offspring survival and development at different stages (fledging, nutritional independence and adulthood) were tested in the cooperatively breeding Arabian babbler to determine the relative importance of social, maternal and environmental factors at each stage. An influence of maternal effects was found during the nestling stage only. 3. Social factors affected the survival and development of young at all stages. The amount of food received from helpers influenced post-fledging weight gain, development of foraging skills, and survival to reproductive age. Environmental effects were also important, with groups occupying high-quality territories more likely to produce young that survived to maturity. 4. The strong influence of helper contributions on the survival and development of young at all stages from hatching to maturity suggests social factors may have important long-term effects on offspring fitness in cooperative societies. Traditional measures of offspring survival in cooperative birds, which commonly measure survival to fledging age only, may underestimate the significant benefit of helper contributions on the survival and development of young.     

Carry-over effects of the social environment on future divorce probability in a wild bird population

Initial mate choice and re-mating strategies (infidelity and divorce) influence individual fitness. Both of these should be influenced by the social environment, which determines the number and availability of potential partners. While most studies looking at this relationship take a population-level approach, individual-level responses to variation in the social environment remain largely unstudied. Here, we explore carry-over effects on future mating decisions of the social environment in which the initial mating decision occurred. Using detailed data on the winter social networks of great tits, we tested whether the probability of subsequent divorce, a year later, could be predicted by measures of the social environment at the time of pairing. We found that males that had a lower proportion of female associates, and whose partner ranked lower among these, as well as inexperienced breeders, were more likely to divorce after breeding. We found no evidence that a female''s social environment influenced the probability of divorce. Our findings highlight the importance of the social environment that individuals experience during initial pair formation on later pairing outcomes, and demonstrate that such effects can be delayed. Exploring these extended effects of the social environment can yield valuable insights into processes and selective pressures acting upon the mating strategies that individuals adopt.     

环境温度和体内储备物共同影响煤山雀夜间体重的下降(英文)

环境温度和体内储备物水平被认为是鸟类在静止状态下能量利用的重要调节因子(夜间体重降低).然而,以往的研究没有把环境温度和体内储备物对夜间能量维持加以明确的区分.为了研究环境温度是否是为煤山雀(Periparus ater)夜间体重调节的直接因子,在自由取食条件下,实验室控制日-日和日-夜环境温度.温度变化模拟西班牙中部地中海山区秋季日-夜温度的变化.夜间体重取决于黄昏时的体重以及前一天体重的增加值.当前一日白天煤山雀体重增加最大时,记录夜间体重最大降低的比率.然而,环境温度的不可预见性没有影响煤山雀夜间体重降低,可以解释煤山雀内在的生理能量平衡.这些结果提示,当一些环境因子如温度变得不可预见时,鸟类在狭小范围内保持体内储备物     

A review of the social behaviour of the giraffe Giraffa camelopardalis: a misunderstood but socially complex species

1. Until around 2000, giraffe Giraffa camelopardalis were believed to have no social structure. Despite a resurgence of interest in giraffe behaviour in around 2010, most studies are of isolated populations, making it difficult to draw general conclusions. Although it is now well established that giraffe social organisation is non-random, there is little consensus as to what influences preferred and avoided associations or the underpinning mechanisms.
2. We test two hypotheses: first, giraffe have a complex cooperative social system, exhibited by 1) stable groups of females, 2) offspring that stay in their natal group for part or all of their lives, 3) support by non-mothers in rearing young, and 4) non-reproductive females in the group; and second, giraffe form matrilineal societies, evidenced by 1) male dispersal, 2) female philopatry, 3) assistance in raising or protecting offspring, and 4) individual benefits gained from social foraging.
3. We reviewed 404 papers on giraffe behaviour and social organisation; captive studies were included where they supplemented information from free-living populations.
4. We show that giraffe exhibit many of the features typical of mammals with complex cooperative social systems and matrilineal societies. However, the social complexity hypothesis posits that such species also require complex communication systems to regulate interactions and relations among group members; giraffe communication systems are poorly understood.
5. Quantifying the fitness and survival benefits of the giraffe’s social organisation is necessary to ensure its long-term survival. Giraffe numbers have declined by 40% since 1985, they have been declared extinct in seven (possibly nine) countries and are listed as Vulnerable by the International Union for Conservation of Nature. We identify research areas that will advance our understanding of giraffe behaviour and conservation requirements.

     

Natural selection on the genetical component of variance in body condition in a wild bird population

Although there is substantial evidence that skeletal measures of body size are heritable in wild animal populations, it is frequently assumed that the nonskeletal component of body weight (or ‘condition’) is determined primarily by environmental factors, in particular nutritional state. We tested this assumption by quantifying the genetic and environmental components of variance in fledgling body condition index (=relative body weight) in a natural population of collared flycatchers (Ficedula albicollis), and compared the strength of natural selection on individual breeding values with that on phenotypic values. A mixed model analysis of the components of variance, based on an ‘animal model’ and using 18 years of data on 17 717 nestlings, revealed a significant additive genetic component of variance in body condition, which corresponded to a narrow sense heritability (h²) of 0.30 (SE=0.03). Nongenetic contributions to variation in body condition were large, but there was no evidence of dominance variance nor of contributions from early maternal or common environment effects (pre‐manipulation environment) in condition at fledging. Comparison of pre‐ and post‐selection samples revealed virtually identical h² of body condition index, despite the fact that there was a significant decrease (35%) in the levels of additive genetic variance from fledging to breeding. The similar h² in the two samples occurred because the environmental component of variance was also reduced by selection, suggesting that natural selection was acting on both genotypic and environmental variation. The effects of selection on genetic variance were confirmed by calculation of the selection differentials for both phenotypic values and best linear unbiased predictor (BLUP) estimates of breeding values: there was positive directional selection on condition index both at the phenotypic and the genotypic level. The significant h² of body condition index is consistent with data from human and rodent populations showing significant additive genetic variance in relative body mass and adiposity, but contrasts with the common assumption in ecology that body condition reflects an individual’s nongenetic nutritional state. Furthermore, the substantial reduction in the additive genetic component of variance in body condition index suggests that selection on environmental deviations cannot alone explain the maintenance of additive genetic variation in heritable traits, but that other mechanisms are needed to explain the moderate to high heritabilities of traits under consistent and strong directional selection.     

Variation in body mass index among Polish adults: effects of sex, age, birth cohort, and social class

Remains of 15 hominids were recovered within a Mousterian archaeological context in the cave of Qafzeh, Israel. Dated to ca. 95 kyr BP, this skeletal material has been crucial for understanding biological, chronological, and cultural aspects of anatomically modern ancient Homo sapiens. The high proportion of children (N = 8) in Qafzeh Cave is unique among Middle Palaeolithic sites and encourages the search for skeletal evidence of disease and trauma. We report on the case of one child, Qafzeh 12, ca. 3 years old (according to modern human reference standards), who manifests some outstanding skeletal abnormalities that indicate hydrocephalus.     

Daily body mass regulation in dominance-structured coal tit (Parus ater) flocks in response to variable food access: a laboratory study

In a dominance-structured flock, social status may determinepriority of access to food. Birds of low social status mayperceive present and future access to food as less predictable,and so have a higher risk of starvation, than birds of highsocial rank. Theoretical models predict that subordinate birdsshould carry larger fat reserves and incur higher mass-dependentcosts than dominants. However, empirical tests of the assumptionsof these models are still scarce and controversial. We investigatedthe effect of dominance rank on daily mass gain under conditionsof fluctuating food availability in a laboratory experimentusing four flocks of four coal tits (Parus ater) each. Thesame amount of food was delivered in two treatments, but inone treatment the food was offered at a constant rate betweendays (fixed treatment), while in the other treatment the dailyfood supply varied in an unpredictable sequence between days(variable treatment). All birds showed greater variance inbody mass in the variable treatment than in the fixed treatment.Body mass within birds showed the same variability at dawn thanat dusk in the fixed treatment, but less variability at dawnthan at dusk in the variable treatment. This may be a mechanismto reduce the immediate risk of starvation at the beginningof the day, when fat reserves are at their lowest and the aggressionbetween flock members when feeding highest. Subordinate birdswere excluded from the feeders by dominants more often in theearly morning than in the rest of the day, and they showedmore variability in daily mass gain and body mass at dawn thandominant birds. These results support the hypothesis that subordinatebirds have a reduced probability of surviving when food availabilitychanges unexpectedly compared to dominants.     

Predators and the breeding bird: behavioral and reproductive flexibility under the risk of predation

A growing body of work suggests that breeding birds have a significant capacity to assess and respond, over ecological time, to changes in the risk of predation to both themselves and their eggs or nestlings. This review investigates the nature of this flexibility in the face of predation from both behavioural and reproductive perspectives, and also explores several directions for future research. Most available work addresses different aspects of nest predation. A substantial change in breeding location is perhaps the best documented response to nest predation, but such changes are not always observed and not necessarily the best strategy. Changes in nesting microhabitat (to more concealed locations) following predation are known to occur. Surprisingly little work addresses the proactive avoidance of areas with many nest predators, but such avoidance is probably widespread. Individual birds could conceivably adopt anti‐predator strategies based on the nest predators actually present in an area, but such effects have yet to be demonstrated. In fact, the ways in which birds assess the risk of nest predation is unclear. Nest defence in birds has historically received much attention, but little is known about how it interacts with other aspects of decision‐making by parents. Other studies concentrate on predation risk to adults. Some findings suggest that risk to adults themselves influences territory location, especially relative to raptor nests. An almost completely unexplored area concerns the sorts of social protection from predators that might exist during the breeding season. Flocking typical of the non‐breeding season appears unusual while breeding, but a mated pair may sometimes act as a “flock of two”. Opportunistic heterospecific sociality may exist, with heterospecific protector species associations more prevalent than currently appreciated. The dynamics of singing during the breeding season may also respond to variation in predation risk, but empirical research on this subject is limited. Furthermore, a few theoretical and empirical studies suggest that changes in predation risk also influence the behaviour of lekking males. The major influence of predators on avian life histories is undoubtedly expressed at a broad phylogenetic scale, but several studies hint at much flexibility on an ecological time scale. Some species may forgo breeding completely if the risk of nest predation is too high, and a few studies document smaller clutch sizes in response to an increase in nest predation. Recent evidence suggests that a female may produce smaller eggs rather than smaller clutches following an increase in nest predation risk. Such an increase may also influence decisions about intraspecific brood parasitism. There are no clear examples of changes in clutch/egg size with changes in risk experienced by adults, but parental responses to predators have clear consequences for offspring fitness. Changes in risk to adults may also influence body mass changes across the breeding season, although research here is sparse. The topics highlighted herein are all in need more empirical attention, and more experimental field work whenever feasible.     

Culling-induced social perturbation in Eurasian badgers Meles meles and the management of TB in cattle: an analysis of a critical problem in applied ecology

The Eurasian badger (Meles meles) is implicated in the transmission of bovine tuberculosis (TB) to cattle in the UK and Republic of Ireland. Badger culling has been employed for the control of TB in cattle in both countries, with varying results. Social perturbation of badger populations following culling has been proposed as an explanation for the failure of culling to consistently demonstrate significant reductions in cattle TB. Field studies indicate that culling badgers may result in increased immigration into culled areas, disruption of territoriality, increased ranging and mixing between social groups. Our analysis shows that some measures of sociality may remain significantly disrupted for up to 8 years after culling. This may have epidemiological consequences because previous research has shown that even in a relatively undisturbed badger population, movements between groups are associated with increases in the incidence of Mycobacterium bovis infection. This is consistent with the results from a large-scale field trial, which demonstrated decreased benefits of culling at the edges of culled areas, and an increase in herd breakdown rates in neighbouring cattle.     

Overwintering survival in relation to body mass in a field population of the wolf spider (Hygrolycosa rubrofasciata)

Body size is often considered to be an important trait affecting individual fitness. In arthropods, females commonly benefit from larger size directly through increased fecundity (Roff, 1992), and males through increased mating success (Andersson, 1994). It has also been suggested that larger individuals may in general have a better survival than smaller individuals (Calder, 1983; Peters, 1983). From this suggestion it may be predicted that during stressful environmental conditions larger individuals should do better than smaller individuals.     

Constraint versus restraint in the body mass dynamics during moult of a species with precocial young

Body mass declines during wing moult in numerous, but not all, populations of Anatidae. We assessed two leading hypotheses for body mass dynamics during wing moult: (1) body mass dynamics are adapted to attain a target body mass at the end of wing moult (restraint hypothesis) vs. (2) body mass dynamics reflect environmental constraint on the nutrient–energy balance during wing moult (constraint hypothesis). We used regressions of mass of breeding female Black Brant Branta bernicla nigricans on ninth primary length (a measure of moult stage) for each of 16 years to assess mass dynamics during wing moult and used regression equations to predict mass at the beginning and end of wing moult each year. We also included gosling mass at 30 days (an indicator of forage availability) in models of adult mass to assess how mass dynamics varied as a function of foraging conditions. Predicted body mass (± 95% CI) at the start of wing moult (ninth primary = 0 mm) varied significantly among years from 1032 ± 52 to 1169 ± 27 g. Similarly, predicted mass in late wing moult (ninth primary = 142 mm) ranged from 1048 ± 25 to 1222 ± 28 g. The rate of mass gain was significantly related to gosling mass at 30 days: interaction between adult ninth primary length and gosling mass = 0.0031 ± 0.0020 (P = 0.003). Females initiated wing moult at lower body masses, gained mass more rapidly and ended with wing moult heaviest when goslings were heaviest. Body mass dynamics of female Black Brant during wing moult were consistent with the constraint hypothesis. The positive association between gosling mass and rate of body mass gain by adult females during wing moult was also consistent with the constraint hypothesis.     

The costs of keeping cool in a warming world: implications of high temperatures for foraging,thermoregulation and body condition of an arid‐zone bird

Recent mass mortalities of bats, birds and even humans highlight the substantial threats that rising global temperatures pose for endotherms. Although less dramatic, sublethal fitness costs of high temperatures may be considerable and result in changing population demographics. Endothermic animals exposed to high environmental temperatures can adjust their behaviour (e.g. reducing activity) or physiology (e.g. elevating rates of evaporative water loss) to maintain body temperatures within tolerable limits. The fitness consequences of these adjustments, in terms of the ability to balance water and energy budgets and therefore maintain body condition, are poorly known. We investigated the effects of daily maximum temperature on foraging and thermoregulatory behaviour as well as maintenance of body condition in a wild, habituated population of Southern Pied Babblers Turdoides bicolor. These birds inhabit a hot, arid area of southern Africa where they commonly experience environmental temperatures exceeding optimal body temperatures. Repeated measurements of individual behaviour and body mass were taken across days varying in maximum air temperature. Contrary to expectations, foraging effort was unaffected by daily maximum temperature. Foraging efficiency, however, was lower on hotter days and this was reflected in a drop in body mass on hotter days. When maximum air temperatures exceeded 35.5 °C, individuals no longer gained sufficient weight to counter typical overnight weight loss. This reduction in foraging efficiency is likely driven, in part, by a trade‐off with the need to engage in heat‐dissipation behaviours. When we controlled for temperature, individuals that actively dissipated heat while continuing to forage experienced a dramatic decrease in their foraging efficiency. This study demonstrates the value of investigations of temperature‐dependent behaviour in the context of impacts on body condition, and suggests that increasingly high temperatures will have negative implications for the fitness of these arid‐zone birds.