Born to cope with climate change? Experimentally manipulated hatching time does not affect duckling survival in the mallard <Emphasis Type="Italic">Anas platyrhynchos</Emphasis>

Two main hypotheses proposed to explain the seasonal decline in reproductive performance in birds are (1) deterioration of environmental conditions and (2) lower parental quality of late breeders. Previous experimental work addressing these hypotheses generally have problematic biases pertaining to delay of hatching, costs of re-laying and incubation, as well as variation in the quality of eggs, territories, offspring and parental traits. We address these biases in an experimental test of the timing hypothesis (i.e. (1) above) in a precocial bird. Using a 2-year cross-over design and game-farm mallard (Anas platyrhynchos) eggs originating from a number of hens and a standardised delay procedure, we introduced early and late broods with a foster female onto boreal oligotrophic lakes and monitored subsequent duckling survival. Standardised invertebrate sampling was done concurrently to get a measure of lake-level abundance of aquatic prey, a likely causative agent of putative seasonal difference in duckling survival. Survival data and covariates (duckling age; days) were analysed by an information theoretic approach. There was no effect of treatment (i.e. manipulation of hatching date) on duckling survival, which was higher in 2005 than in 2004. In contrast to observational studies from more seasonal wetlands, our experiment demonstrates that duckling survival on boreal lakes was not affected by a 12-day delay in hatching date. Since we did not find any consistent trends in abundance of aquatic prey, i.e. neither clear peaks nor differences between treatment periods, we hypothesise that moderate climate change has minor effects on resource abundance and hence also on mallard duckling survival in boreal environments.     

Effects of radiotransmitters and plasticine bands on Mallard duckling survival

ABSTRACT Individual markers are essential for measuring survival rates, but few reliable techniques exist for marking young waterfowl. We banded 572‐d‐old Mallard (Anas platyrhynchos) ducklings from 85 broods with plasticine‐filled leg bands in 2006–2007 and marked 82 hatchling Mallards from 43 broods with 1.8‐g back‐mounted prong‐and‐suture transmitters in 2006. We compared survival probabilities of marked ducklings and 621 unmarked ducklings from 110 broods where only the female was radio‐marked using the nest survival module in Program MARK. The best‐supported model recognized deleterious effects of both radiotransmitters (β=?0.82, 85% CI =?1.32 to ?0.33) and plasticine bands (β=?0.29, 85% CI =?0.55 to ?0.03). Cumulative survival to 30 d of age for unmarked ducklings averaged 0.22 (process variation; σ= 0.11) in 2006 and 0.20 (σ= 0.11) in 2007, whereas 30‐d survival of banded ducklings during these 2 yr was 0.14 (σ= 0.09) and 0.13 (σ= 0.09), respectively. Radio‐marked ducklings averaged 0.04 (σ= 0.03) cumulative survival in 2006. Although capturing ducklings after they leave nests is not always possible, post hoc analysis showed that marker effects decreased with duckling age and waiting until ducklings were at least 4 d old before marking would have negated these effects. Because marking hatchling Mallards with either radiotransmitters or plasticine bands negatively impacted their survival, we do not recommend these marking techniques for newly hatched ducklings.     

The diet of Grey Herons Ardea cinerea breeding at Loch Leven, Scotland, and the importance of their predation on ducklings

Regurgitations from nestling Grey Herons Ardea cinerea at Loch Leven, Scotland, April-July 1981–83, contained mainly three types of prey:perch, brown trout and ducklings. The diet varied markedly through the season but not between years. As the heron breeding season progressed, perch occurred in fewer regurgitations and ducklings in more. There was no seasonal variation in the occurrence of brown trout. Regular collections of duckling down from the heronry suggested that the consumption of ducklings peaked in early June. The ducklings taken by herons were less than 10 days old, Mallard ducklings predominating before mid June and Tufted ducklings later. Most ducklings were taken by only a few herons:those that bred earliest and which initially fed their young on perch. Ducklings became a major part of their diet in the late nestling period and broods fed on ducklings fledged no more or fewer young than others. Herons feeding their young on brown trout took ducklings rarely, if ever. It is argued that variation in the contents of regurgitations resulted from three interacting variables; the type of feeding habitat used by individual Grey Herons, the date at which they bred and the date that regurgitations were produced by their nestlings. The numbers of ducklings taken by herons were calculated to be about 230 in 1981 and 291 in 1982, about 5% of Mallard and 3:b of Tufted ducklings estimated to have hatched annually. Assuming herons continued to consume ducklings at the same rate after their young had dispersed, the figures for Tufted ducklings would be higher but still only about 4% in 1981 and 6% in 1982. Compared with total duckling losses of over 75%, predation by Grey Herons was minor and did not affect duckling production in the years concerned.     

Survival of Wood Duck Ducklings and Broods in Mississippi and Alabama

ABSTRACT Although North American wood ducks (Aix sponsa) are well-studied throughout their range, researchers know little about demographic and environmental factors influencing survival of ducklings and broods, which is necessary information for population management. We studied radiomarked female and duckling wood ducks that used nest boxes and palustrine wetlands at Noxubee National Wildlife Refuge (NNWR) in Mississippi, USA, in 1996–1999, and riverine wetlands of the Tennessee-Tombigbee Rivers and Waterway (TTRW) system in Alabama in 1998–1999. We estimated survival of ducklings and broods and evaluated potentially important predictors of duckling survival, including age and body mass of brood-rearing females, hatch date of ducklings, duckling mass, brood size at nest departure, inter-day travel distance by ducklings, site and habitat use, and daily minimum air temperature and precipitation. At NNWR, survival of 300 radiomarked ducklings ranged from 0.15 (95% CI = 0.04-0.27) to 0.24 (95% CI = 0.13-0.38) and was 0.21 (95% CI = 0.15-0.28) for 1996–1999. Our overall estimate of brood survival was 0.64 (n = 91; 95% CI = 0.54-0.73). At TTRW, survival of 129 radiomarked ducklings was 0.29 in 1998 (95% CI = 0.20-0.41) and 1999 (95% CI = 0.13-0.45) and was 0.29 (95% CI = 0.20-0.40) for 1998–1999. Our overall estimate of brood survival was 0.71 (n = 38; 95% CI = 0.56-0.85). At NNWR, models that included all predictor variables best explained variation in duckling survival. Akaike weight (w_i) for the best model was 0.81, suggesting it was superior to other models (<0.01 < w_i < 0.18). We detected 4 competing models for duckling survival at TTRW. Inter-day distance traveled by ducklings was important as this variable appeared in all 4 models; duckling survival was positively related to this variable. Patterns of habitat-related survival were similar at both study areas. Ducklings in broods that used scrub-shrub habitats disjunct from wetlands containing aggregations of nest boxes had greater survival probabilities than birds remaining in wetlands with such nest structures. Managers may increase local wood duck recruitment by promoting availability of suitable brood habitats (e.g., scrub-shrub wetlands) without aggregations of nest boxes that may attract predators and by dispersing nest boxes amid or adjacent to these habitats. We did not determine an optimal density of nest boxes relative to local or regional population goals, which remains important research and conservation needs.     

Social and maternal factors affecting duckling survival in eiders Somateria mollissima

1. With the aid of a novel survivorship model, an 8-year field study of social and maternal factors affecting duckling survival in eiders (Somateria mollissima) revealed that duckling survival probability varies in accordance with maternal brood-rearing strategy. This variability in survival provides compelling evidence of different annual fitness consequences between females that share brood-rearing and those that tend their broods alone. Consequently, as prebreeding survival is often a major source of individual variation in lifetime reproductive success, a female's annual, state-dependent (e.g. condition) choice of a brood-rearing strategy can be a critical fitness decision. 2. Variance in duckling survival among lone tender broods was best explained by a model with significant interannual variability in survival, and survivorship tending to increase with increasing clutch size at hatch. Clutch size was correlated positively with female condition. Hatch date and female body condition together affected duckling survival, but their contributions are confounded. We were unable to identify a relationship between female age or experience and duckling survival. 3. Variance in duckling survival among multifemale brood-rearing coalitions was best explained by a model that included the number of tenders, the number of ducklings and interannual variation in how their ratio affected survivorship. Hatch date did not significantly influence survival. 4. Expected duckling survival is higher in early life for lone tenders when compared with multifemale brood-rearing coalitions. However, as ducklings approach 2-3 weeks of age, two or three females was the optimal number of tenders to maximize daily duckling survival. The survivorship advantage of multifemale brood-rearing coalitions was most evident in years of average survival. 5. The observed frequency distribution of female group sizes corresponds with the distribution of offspring survival probabilities for these groups. Evidence for optimal group sizes in nature is rare, but the most likely candidates may be groups of unrelated animals where entry is controlled by the group members, such as for female eiders. 6. Our study demonstrates that differences in social factors can lead to different predictions of lifetime reproductive success in species with shared parental care of self-feeding young.     

Environmental and not maternal effects determine variation in offspring phenotypes in a passerine bird

Maternal and environmental effects can profoundly influence offspring phenotypes, independent of genetic effects. Within avian broods, both the asymmetric post‐hatching environment created by hatching asynchrony and the differential maternal investment through the laying sequence have important consequences for individual nestlings in terms of the allocation of resources to body structures with different contributions to fitness. The purpose of this study was to evaluate the relative importance of post‐hatching environmental and maternal effects in generating variation in offspring phenotypes. First, an observational study showed that within blue tit, Cyanistes caeruleus, broods, late‐hatched nestlings allocated resources to tarsus development, maintained mass gain and head‐bill growth and directed resources away from the development of fourth primary feathers. Second, a hatching order manipulation experiment resulted in nestlings from first‐laid eggs hatching last, thereby allowing comparison with both late and early‐hatched nestlings. Experimental nestlings had growth patterns which were closer to late‐hatched nestlings, suggesting that within‐brood growth patterns are determined by post‐hatching environmental effects. Therefore, we conclude that post‐hatching environmental effects play an important role in generating variation in offspring phenotypes.     

Within‐female plasticity in sex allocation is associated with a behavioural polyphenism in house wrens

Sex allocation theory assumes individual plasticity in maternal strategies, but few studies have investigated within‐individual changes across environments. In house wrens, differences between nests in the degree of hatching synchrony of eggs represent a behavioural polyphenism in females, and its expression varies with seasonal changes in the environment. Between‐nest differences in hatching asynchrony also create different environments for offspring, and sons are more strongly affected than daughters by sibling competition when hatching occurs asynchronously over several days. Here, we examined variation in hatching asynchrony and sex allocation, and its consequences for offspring fitness. The number and condition of fledglings declined seasonally, and the frequency of asynchronous hatching increased. In broods hatched asynchronously, sons, which are over‐represented in the earlier‐laid eggs, were in better condition than daughters, which are over‐represented in the later‐laid eggs. Nonetheless, asynchronous broods were more productive later within seasons. The proportion of sons in asynchronous broods increased seasonally, whereas there was a seasonal increase in the production of daughters by mothers hatching their eggs synchronously, which was characterized by within‐female changes in offspring sex and not by sex‐biased mortality. As adults, sons from asynchronous broods were in better condition and produced more broods of their own than males from synchronous broods, and both males and females from asynchronous broods had higher lifetime reproductive success than those from synchronous broods. In conclusion, hatching patterns are under maternal control, representing distinct strategies for allocating offspring within broods, and are associated with offspring sex ratios and differences in offspring reproductive success.     

Survival of common goldeneye Bucephala clangula ducklings in relation to weather, timing of breeding, brood size, and female condition

The survival of common goldeneye Bucephala clangula ducklings during their first week of life was studied in relation to hatching date, brood size, female condition, and weather (temperature and precipitation) during the first week post-hatch by using data from radio-marked females and their broods. Also, age-specific variation in the survival of the young was determined until fledging (over 50 d of age). Survival was lowest in the first week after hatching. Hatching date, brood size, and first week temperature and precipitation were poor predictors of duckling survival during the first week after nest exodus. Instead, the ducklings of females in a better body condition survived better in their first week of life. The results suggest that weather does not have a direct effect on downy ducklings' survival, but the condition of the female seems to be an important determinant of the survival of common goldeneye ducklings.     

Effects of variation in egg size and hatching date on survival of Lesser Scaup Aythya affinis ducklings

The consequences of avian egg-size variation on offspring quality and survival remain unclear. We evaluated the effects of egg-size and hatch-date variation on survival of Lesser Scaup Aythya affinis ducklings in the wild. Duckling mass at hatching increased significantly with increasing egg size. Ducklings from larger eggs survived better than those from smaller eggs. We suspect that ducklings from larger eggs survived better because of advantages associated with larger or more efficient utilization of nutrient reserves, or both. We were unable to detect any within-clutch differences in egg size of survivors and non-survivors, nor any consistent direction in the difference in egg size between survivors and nonsurvivors within clutches. This suggests that within-clutch variation may be insufficient to have survival consequences for offspring. In addition, ducklings that hatched later in the breeding season had a higher probability of survival. We suggest a food-dependent hypothesis as an explanation for the seasonally increasing survival and for later nest initiation of Lesser Scaup compared with other North American ducks.     

Seasonal decline in cell-mediated immunity of collared flycatcher Ficedula albicollis nestlings: does the sex of offspring matter?

The seasonal decline in offspring performance is a frequently reported phenomenon in species breeding in a temperate zone, but the potential effect of brood sex ratio on such declines has not been studied. Here, we predicted that this decline may occur if the sex that exhibits the lower immune response or lower survival rate tends to be more frequent among late broods. The seasonal patterns of four performance parameters of collared flycatcher Ficedula albicollis nestlings have been examined during 4 years. Sex was assigned to all studied individuals using molecular techniques. We found significant seasonal decline in cell-mediated immune response, tarsus length and survival of the chicks. The lack of interactions between gender and hatching date revealed that both sexes contributed equally to the observed decline. The brood sex ratio did not vary with the laying date. On the basis of available data, we suggest that the breeding date may only exceptionally induce female-driven sex allocation in species with only slight sexual size dimorphism. In consequence, we suggest that seasonal sex ratio shifts do not account for seasonally declining fitness of nestlings in passerines.     

Incubation temperature influences locomotor performance in young wood ducks (Aix sponsa)

Incubation temperature is an important maternal effect in birds that can influence numerous offspring traits. For example, ducklings from eggs incubated at lower temperatures have lower growth rates, protein content, and are in poorer body condition than ducklings from eggs incubated at higher temperatures. Based on these observations, we predicted that incubation temperature would indirectly influence performance through its direct effects on body size. Wood duck (Aix sponsa) eggs were incubated at three ecologically relevant temperatures (35, 35.9, 37°C). After hatching, all ducklings were housed under identical conditions and were subjected to aquatic and terrestrial racing trials at 15 and 20 days posthatch (dph). Contrary to our prediction, incubation temperature did not influence most duckling body size parameters at 15 or 20?dph. However, incubation temperature did have a strong influence on locomotor performance independent of body size and body condition. Ducklings hatched from eggs incubated at the lowest temperature had significantly reduced maximum aquatic swim velocity than ducklings from higher temperatures. Maximum terrestrial sprint velocity followed a similar pattern, but did not differ statistically among incubation treatments. To our knowledge, this is the first study to demonstrate that slight changes in incubation temperature can directly affect locomotor performance in avian offspring and thus provide a significant source of phenotypic variation in natural wood duck populations.     

Spring phenology and timing of breeding in short‐distance migrant birds: phenotypic responses and offspring recruitment patterns in common goldeneyes

Understanding how organisms adjust breeding dates to exploit resources that affect fitness can provide insights into impacts of climate change on avian demography. For instance, mismatches have been reported in long‐distance migrant bird species when environmental cues experienced during spring migration are decoupled from conditions on breeding grounds. Short‐distance migrant bird species that store reproductive nutrients prior to breeding may avoid or buffer adverse phenological effects. Furthermore, reduced short‐term reproductive success could be offset by higher future recruitment of surviving offspring. We evaluated whether recruitment of locally‐hatched female offspring was related to hatching date alone or strength of mismatched breeding date for 405 individually‐marked adult female common goldeneyes Bucephala clangula (a short‐distance migrant) and their ducklings from a site in central Finland where ice‐out date has advanced by ～ 2 weeks over 24 yr. Path analyses revealed that older, early‐nesting females with good body condition and larger broods recruited the most female offspring. Offspring recruitment decreased strongly among females that bred late relative to other females in the population each year; the extent of mismatched breeding date, i.e. hatching date scaled to annual ice‐out date, was less influential. Overall, most females advanced breeding dates when ice‐out occurred earlier in spring, but some females exhibited greater flexibility in response to ice‐out conditions than did others. In general, directional selection favoured early breeding over a wide range of ice‐out dates. Our results seem most consistent with a hypothesis that some short‐distance migrant species like goldeneyes have the capacity to track and respond appropriately to changing environmental conditions prior to onset of breeding.     

Within-season sequential density dependence regulates breeding success in mallards Anas platyrhynchos

Density dependence in vital rates is a key issue in population ecology but remains largely unexplored experimentally. We studied breeding success, lake use, and prey availability in wild mallards Anas platyrhynchos on small nemoral lakes in a replicated, two-year cross-over experiment in which pair density was increased. The number of wild mallards that settled on lakes prior to introductions of extra pairs did not differ between control and introduction years. Introductions led to a lake-level reduction in the number of broods observed. However, the number of stage 2+ (almost fledged) ducklings did not differ between treatments, nor did lake utilization by nonbreeding adults, broods and ducklings. Prey resource availability differed greatly among lakes, but it did not correlate with breeding success. Partialling out the possible effect of food competition from wild adult nonbreeding mallards did not change this conclusion. Our study demonstrates sequential density dependence in breeding success; introductions caused a decrease in brood number, but despite fewer broods a similar number of nearly fledged ducklings were produced. We suggest that predation and/or lake change of broods soon after hatching created these patterns. We conclude that using a single and late measure of breeding success such as fledged birds can mask regulatory processes. Implications of density dependence and its relation to individual reproductive success are understood better if breeding success is decomposed into nest success, duckling survival and fledgling survival.     

Growth rate, size, and sex ratio of last-laid, last-hatched offspring in the tree swallow Tachycineta bicolor

In tree swallows Tachycineta bicolor, last‐laid eggs typically hatch one to two days after the other eggs in the clutch hatch, putting last‐hatched offspring at a disadvantage when competing for food delivered by parents. We studied the biology of last‐laid, last‐hatched tree swallow offspring over two years in a Wyoming, USA, population. Our first objective was to compare the growth of last‐hatched offspring to that of their earlier‐hatched nestmates. One previous study had suggested that last‐hatched, competitively disadvantaged offspring grow feathers faster than senior nestmates, even at the expense of other aspects of growth. This may allow last‐hatched offspring to fledge with senior nestmates and avoid abandonment by parents. A second objective was to determine the sex of nestlings from last‐laid eggs. If last‐laid eggs typically produce undersized, weak adults that are poor competitors for resources, and if the fitness costs of being undersized/weak are more severe for males than for females, then selection may favour having offspring from last‐laid eggs to be female. In this study, last‐laid eggs hatched in 63 of 66 (94%) nests and hatched last in 93% of cases. At hatching, offspring from last‐laid eggs weighed, on average, 63% as much as their three heaviest nestmates (range: 26–107%). Offspring from last‐laid eggs fledged from 71% of the nests that produced at least one fledgling and apparently starved to death in remaining nests. Last‐hatched offspring who were presumably at a substantial competitive disadvantage (those whose mass at hatching was no more than about 75% of the mean mass of their three heaviest nestmates), gained mass more slowly than their senior nestmates but they eventually attained the same peak mass before fledging. Last‐hatched offspring grew primary feathers more slowly than their senior nestmates although the difference in growth rate was slight (0.2 mm/d) and only marginally significant. As a group, offspring from last‐laid eggs did not differ from offspring from all other eggs in either maximum mass attained before fledging or tarsus length at fledging. This is atypical for species with asynchronous hatching and is possibly the result of another unusual trait: the tendency of parent tree swallows to distribute food equally among young within broods. The sex ratio of offspring from last‐laid eggs did not deviate from 1:1 (22 males, 21 females). Given that last‐hatched eggs do not routinely produce undersized/weak individuals in our study population, there should be little selection on parent females to bias the sex ratio of last‐laid offspring towards females.     

Sex-specific fledgling success and brood sex ratio in the Great Reed Warbler (Acrocephalus arundinaceus)

We examine sex ratio variation and sex-specific probability of successful fledgling in relation to hatching date across 376 broods of Great Reed Warblers (Acrocephalus arundinaceus). The sex ratio of complete broods as well as broods with partial mortality did not deviate significantly from parity (0.5 and 0.53, respectively). Variation in sex ratio between broods was not larger than expected from binomial distribution, thus females seem not to manipulate the sex ratio of their broods in the studied population. As a consequence, sex ratio did not vary in relation to hatching date, years and fishponds. Female offspring showed lower fledgling success than their brothers, but the relationship between probability of successful fledgling and hatching date differed between sexes. Fledgling success of female offspring declined with hatching date more strongly than the success of male offspring. Thus, our study shows that Great Reed Warblers do not adjust offspring sex to match observed seasonal sex-specific variation in survival.     

Hatching asynchrony and brood reduction influence immune response in Common Kestrel Falco tinnunculus nestlings

The onset of incubation before the end of laying imposes asynchrony at hatching and, therefore, a size hierarchy in the brood. It has been argued that hatching asynchrony might be a strategy to improve reproductive output in terms of quality or quantity of offspring. However, little is known about the mediating effect of hatching asynchrony on offspring quality when brood reduction occurs. Here, we investigate the relationship between phenotypic quality and hatching asynchrony in Common Kestrel Falco tinnunculus nestlings in Spain. Hatching asynchrony did not increase breeding success or nestling quality. Furthermore, hatching asynchrony and brood reduction had different effects on nestlings’ phytohaematogglutinin (PHA)‐mediated immune response and nestling growth. In asynchronous and reduced broods (in which at least one nestling died), nestlings showed a stronger PHA‐mediated immune response and tended to have a smaller body size compared with nestlings raised in synchronous and reduced broods. When brood reduction occurred in broods hatched synchronously, there was no effect on nestling size, but nestlings had a relatively poor PHA‐mediated immune response compared with nestlings raised in asynchronous and reduced broods. We suggest that resources for growth can be directed to immune function only in asynchronously hatched broods, resulting in improved nestling quality, as suggested by their immune response. We also found that males produced a greater PHA‐mediated immune response than females only in brood‐reduced nests without any effect on nestling size or condition, suggesting that females may trade off immune activities and body condition, size or weight. Overall, our results suggest that hatching pattern and brood reduction may mediate resource allocation to different fitness traits. They also highlight that the resolution of immune‐related trade‐offs when brood reduction occurs may differ between male and female nestlings.     

Rearing other peoples' young; Brood-mixing in the shelduck Tadorna tadorna

The mixing of ducklings from different broods, in several species of waterfowl, involves care of offspring other than the parents' own. In the eider, Somateria mollissima, this behaviour appears to be of benefit, in terms of increased adult and duckling survival, to both donor and recipient parents, but in the shelduck, Tadorna tadorna, brood-mixing appears not to be advantageous. Dominant males tended to lose ducklings to more subordinate ones but there was no significant difference between donor and recipient parents in the number of their own offspring reared to fledging. The apparent lack of any advantage in brood-mixing suggests that the behaviour might be a consequence of high brood density and that in the shelduck brook defence is adapted to low density.     

Sex of the first hatched chick influences survival of the brood in the herring gull (Larus argentatus)

Differences in the growth rate of male and female offspring can result in different parental rearing costs for sons and daughters. Such differences may also influence the survival chances of male and female offspring when conditions are unfavourable. In birds, hatching asynchrony leads to hierarchical competition for food between siblings. Therefore, the sex of the chick in the first hatched position in the brood may influence breeding success by affecting the extent to which the later hatched chicks can compete for resources. The interaction between brood sex composition and chick performance in the herring gull Larus argentatus was examined under different environmental conditions. When environmental conditions were relatively good, chick survival within broods was better when a female was first to hatch, an effect that was most obvious later in the season. When conditions were poorer however, sex of the first hatched chicks was not related to brood survival. In neither situation did the overall primary sex ratio differ from equality. However in the year of relatively good food availability, the first chick in the brood was more likely to be male early in the season, which was when the disadvantageous effects on brood survival of males being in this position are weakest.     

Family living among birds

Family cohesion in birds is often explained as a product of limitations on breeding opportunities leading to delayed dispersal among the offspring. Yet, it is not independent reproduction but floating (queuing outside the natal territory for a breeding opportunity) that is the alternative to delaying dispersal. In contrast to a strong phylogenetic bias in cooperative breeding that has been taken to indicate a hard-wired behaviour maintained by evolutionary inertia, offspring behaviour is plastic and facultative in group living in the short term, indicating that family cohesion is under selection. Non-breeding offspring could gain inclusive fitness that would promote family cohesion from delaying dispersal and providing alloparental care to subsequent broods to boost group productivity. This holds in particular for offspring hatched from early broods in multi-brooded species that have the opportunity to gain inclusive fitness from help in rearing siblings hatched from later broods in the same season. Yet, seasonality will circumscribe the potential for alloparenting to be the immediate factor selecting for family cohesion. The option of gaining inclusive fitness from providing alloparental care is not open to the offspring among single-brooded species until after they have survived a non-breeding season, rather indicating kin cooperation returning enhanced survival prospects from general group living effects as a more immediate factor selecting for family cohesion. Indeed, a variety of family groups maintained in the absence of alloparental care underlines the capacity of general group living enhancing survival as a primary agent selecting for family cohesion. These seasonal constraints on fitness components selecting for family cohesion may contribute to the large scale geographical pattern with a relative paucity of family cohesions among bird species in the northern hemisphere.     

A novel PCR‐based genetic marker shows that sex of offspring does not account for hatch‐order effects on growth,survival and dominance in Pūkeko Porphyrio melanotus melanotus

Intra‐brood competition can influence a variety of fitness‐related traits in birds. Previous research on the joint‐nesting Pūkeko Porphyrio melanotus melanotus, a New Zealand subspecies of Australasian Swamphen, showed that chicks that hatched earlier in a brood tended to grow faster, were more likely to survive and had higher dominance status as adults than later hatched nest‐mates. However, this finding could be due to changes in offspring sex ratio across hatch order (e.g. if males tend to hatch earlier), which was not previously examined because of methodological challenges associated with sexing nestling Pūkeko. Here, we report a useful PCR‐based genetic marker to determine the sex of Pūkeko. We then used new sex‐specific data to re‐examine patterns of offspring growth, survival and dominance. We found that the sex of offspring does not account for the hatching‐order patterns related to social dominance, growth or survival. Furthermore, changes in offspring sex ratio across hatching‐order were negligible and offspring sex ratios did not differ significantly between the primary female and secondary female broods (in joint‐clutch nests), or when comparing primary female and single female broods. We found no clear evidence for sex ratio bias according to hatching‐order and conclude that hatching‐order and not offspring sex explain patterns of growth, survivorship and adult dominance in Pūkeko.