Growth of nestling sparrowhawks (Accipiter nisus)

Growth rates, mortality and parental care of nestling sparrowhawks were studied in southwest Scotland. Ae Forest was a conifer plantation 200–400 metres above sea-level, while the Annan valley consisted of farmland, woods, and small plantations on low ground.
Nestling sparrowhawks were measured daily from hatching for 21–24 days. Weight, tarsus length and outermost primary feather length were recorded. Nestlings could be sexed by the age of 16 days from the larger size of females, which were significantly heavier than males at one day, had longer tarsi at nine days, and longer primaries at 18 days. Growth rates were calculated using linear regression over standardized periods of about 10 days.
Growth rates were independent of brood size, and were negatively correlated with hatching date in one area. Hatching order and growth rate were correlated within broods. The greatest differences in growth rates were found between zones of Ae Forest, and between forest and valley.
Twenty-one per cent of nestlings over two days old died. Causes of mortality were starvation, wet weather, predation and desertion. Most of the mortality occurred in parts of Ae Forest remote from valley woodlands.
The presence or absence of the adult female was noted on nest visits. When habitually brooding, until the young were 11 days old, the hen was present on about 85% of visits. By fledging this figure fell to 66% in the valley, and to 32% in the forest.
The development of sexual dimorphism is discussed; females gained weight and body size faster than males, which developed various skills sooner.
It is suggested that differences in growth rates between parts of the study area were related to food supply. Poor growth rates, high mortality, and lack of parental care all occurred in areas which were remote from sources of abundant prey, as measured by song-bird censuses.     

The effects of hatching asynchrony on growth and mortality patterns in Eurasian Hoopoe Upupa epops nestlings

Growth is a fundamental life history trait in all organisms and is closely related to individual fitness. In altricial birds, growth of many traits is restricted to the short period between hatching and fledging and strongly depends on the amount of food that parents deliver and the extent of hatching asynchrony. However, empirical studies of energy allocation to growth of different body size traits as a function of hatching asynchrony are scarce. We studied growth and mortality of Eurasian Hoopoe Upupa epops, a species with a long breeding season and high brood size variance, whose nestlings show pronounced hatching asynchrony, in order to test how hatching asynchrony affects different growth traits in the context of territory quality, season and brood size. The growth of five body traits (body mass, and lengths of tarsus, third primary, bill and longest crest feather) was investigated to understand how it was affected by brood size, hatching date and order, and territory quality. In total, 241 nestlings from 39 nests were measured every 4 days in 2014 in south‐western Switzerland. Brood size, hatching date and hatching order had the strongest influence on growth trajectories, although tarsus growth was only marginally affected by these variables. Nestlings that hatched earlier than their siblings were heavier and had longer third primaries, bills and crest feathers compared with later‐hatched siblings. In territories of high quality, hatching order differences disappeared for body mass growth, but persisted for lengths of third primary, bill and crest feathers. Brood size was inversely associated with third primary, bill and crest feather lengths, but positively associated with body mass. Nestling mortality was higher in later‐hatched nestlings and in broods that were raised in territories of lower quality. Our study shows that in nestlings, energy was allocated differentially between body traits and this allocation interacted with hatching order and territory quality. Rapid mass gain by nestlings was prioritized in order to increase competitive ability. Our results provide support for the brood reduction hypothesis as an explanation of hatching asynchrony in Hoopoes.     

Reversed sexual dimorphism in tawny owls, Strix aluco, correlates with duty division in breeding effort

Even though most bird species with a raptorial feeding habit express varying extents of reversed sexual dimorphism (RSD: females bigger than males), the evolutionary basis for its maintenance, as well as its possible secondary consequences for the ecological adaptations of the different sexes, is debated. We studied pairs of tawny owls, Strix aluco (females 20% heavier than males), throughout the year by telemetry to test whether any inter-sexual differences in movement patterns, resource partitioning and breeding effort correlated with RSD. Females were larger than males in all body size measures and were 16% heavier than would be expected from the difference in wing length alone. In accordance with predictions from flight economics, males moved longer distances per time unit than females, in particular during the post-fledging season, when they also fed chicks more often than the females. Males had larger home ranges than females during the post-fledging period, whereas the sexes had home ranges of equal size during the non-breeding season. Until 10 days after fledging, females foraged much closer to the offspring than males, apparently balancing their distance to offspring between the needs of offspring guarding and foraging. In males, the parent–offspring distance only increased with decreasing brood condition. The sexes did not differ in habitat use or feeding habits, rendering no indications of food niche partitioning. The study provides further evidence that selection for males to be light and energetically efficient foragers is the main evolutionary force behind RSD in raptorial birds, even when the prey base is confined by territoriality.     

Relationships between body composition, body size and alternative reproductive tactics in a lekking sandpiper, the Ruff (Philomachus pugnax)

1. The Ruff is a lekking sandpiper in which males have two genetically determined alternative mating strategies: 'satellite' or 'independent'. Satellite males are non-territorial, following groups of females on and off leks. Independent males attempt to establish territories on leks and can be either 'resident' territory holders, or non-territorial 'marginals'. The time budgets of the three behavioural tactics (resident, marginal and satellite) differ notably in time allocated to foraging, aggression and display activity. These differences have led to the suggestion that the benefits of size and energy stores may vary with mating tactic.
2. In this paper in vivo estimates of body composition (fat, fat-free mass) for breeding male Ruffs using total body electrical conductivity (TOBEC) are presented.
3. Satellite males have significantly shorter tarsi and wings than independent males.
4. After correcting for size, independent males are significantly heavier and fatter than satellites, and marginals in particular are heavier and fatter than both residents and satellites.
5. Estimates of energy expenditure during flight suggest that satellites may maintain reduced energy reserves to minimize flight costs, while the larger fat stores of independent males are consistent with the benefits of endurance as a territory holder.     

The individual counts: within sex differences in foraging strategies are as important as sex‐specific differences in masked boobies Sula dactylatra

Understanding how animals allocate their foraging time is a central question in behavioural ecology. Intrinsic factors, such as body mass and size differences between sexes or species, influence animals’ foraging behaviour, but studies investigating the effects of individual differences in body mass and size within the same sex are scarce. We investigated this in chick‐rearing masked boobies Sula dactylatra, a species with reversed sexual dimorphism, through the simultaneous deployment of GPS and depth‐acceleration loggers to obtain information on foraging movements and activity patterns. Heavier females performed shorter trips closer to the colony than lighter females. During these shorter trips, heavier females spent higher proportions of their flight time flapping and less time resting on the water than lighter females did during longer trips. In contrast, body mass did not affect trip duration of males, however heavier males spent less time flapping and more time resting on the water than lighter males. This may occur as a result of higher flight costs associated with body mass and allow conservation of energy during locomotion. Body size (i.e. wing length) had no effect on any of the foraging parameters. Dive depths and dive rates (dives h^?1) were not affected by body mass, but females dived significantly deeper than males, suggesting that other factors are important. Other studies demonstrated that females are the parent in charge of provisioning the chick, and maintain a flexible investment under regulation of their own body mass. Variation in trip length therefore seems to be triggered by body condition in females, but not in males. Consequently, shorter trips are presumably used to provision the chick, while longer trips are for self‐maintenance. Our findings underline the importance of accounting for the effects of body mass differences within the same sex, if sex‐specific foraging parameters in dimorphic species are being investigated.     

The effect of parental quality and malaria infection on nestling performance in the Collared Flycatcher (Ficedula albicollis)

Plumage ornamentation often signals the quality of males and, therefore, female birds may choose elaborately ornamented mates to increase their fitness. Such mate choice may confer both direct and indirect benefits to the offspring. Males with elaborate ornaments may provide good genes, which can result in better nestling growth, survival or resistance against parasitic infections. However, these males may also provision their offspring with more food or food of better quality, resulting in nestlings growing at a higher rate or fledging in better condition. In this study, we examined if there was an association between male ornamentation and malaria infection in Collared Flycatchers (Ficedula albicollis). We also investigated offspring performance in relation to malaria infection in the parents and the quality of the genetic and rearing fathers (assessed by the size of two secondary sexual characters) under simulated good and bad conditions (using brood size manipulation). We found that secondary sexual characters did not signal the ability of males to avoid parasitic infections, and malaria infection in the genetic and the rearing parents had no effect on nestling growth and fledging size. Our results do show, however, that it may be beneficial for the females to mate with males with a large forehead patch because wing feathers of nestlings reared by large-patched males grew at a higher rate. Fast feather growth can result in earlier fledging which, in turn, could improve nestling survival in highly variable environments or under strong nest predation.     

Sex allocation in the sexually monomorphic fairy martin

Offspring sex ratios were examined at the population and family level in the sexually monomorphic, socially monogamous fairy martin Petrochelidon ariel at five colony sites over a 4-year period (1993–1996). The sex of 465 nestlings from 169 broods was determined using sex-specific PCR at the CHD locus. In accordance with predicted sex allocation patterns, population sex ratios at hatching and fledging did not differ from parity in any year and the variance in brood sex ratios did not deviate from the binomial distribution. Further, brood sex ratio did not vary with hatching date during the season, brood number, brood size or colony size. The sex ratio of broods with extra-pair young did not differ from those without, while the sex ratio of broods fathered by males that gained extra-pair fertilizations did not differ from broods fathered by other males. Extra-pair chicks were as likely to be male as female. Neither the total number of feeding visits to the brood nor the relative feeding contribution by the sexes varied significantly with brood sex ratio. Brood sex ratios were also unrelated to paternal size, condition and breeding experience or maternal condition and breeding experience. However, contrary to our prediction, brood sex ratio was negatively correlated with maternal size. Generally, these results were consistent with our expectations that brood sex ratios would not vary with environmental factors or parental characteristics, and would not influence the level of parental provisioning. However, the finding that females with longer tarsi produced an excess of daughters is difficult to reconcile with our current understanding of fairy martin life history and breeding ecology.     

Development of intraspecific size variation in black coucals,white-browed coucals and ruffs from hatching to fledging

Most studies on sexual size dimorphism address proximate and functional questions related to adults, but sexual size dimorphism usually develops during ontogeny and developmental trajectories of sexual size dimorphism are poorly understood. We studied three bird species with variation in adult sexual size dimorphism: black coucals (females 69% heavier than males), white-browed coucals (females 13% heavier than males) and ruffs (males 70% heavier than females). Using a flexible Bayesian generalized additive model framework (GAMM), we examined when and how sexual size dimorphism developed in body mass, tarsus length and bill length from hatching until fledging. In ruffs, we additionally examined the development of intrasexual size variation among three morphs (Independents, Satellites and Faeders), which creates another level of variation in adult size of males and females. We found that 27–100% of the adult inter- and intrasexual size variation developed until fledging although none of the species completed growth during the observational period. In general, the larger sex/morph grew more quickly and reached its maximal absolute growth rate later than the smaller sex/morph. However, when the daily increase in body mass was modelled as a proportion, growth patterns were synchronized between and within sexes. Growth broadly followed sigmoidal asymptotic models, however only with the flexible GAMM approach, residual distributions were homogeneous over the entire observation periods. These results provide a platform for future studies to relate variation in growth to selective pressures and proximate mechanisms in these three species, and they highlight the advantage of using a flexible model approach for examining growth variation during ontogeny.     

Fitness and feather wear in the Collared Flycatcher Ficedula albicollis

Individual variation in the degree of feather wear is potentially a useful marker of individual quality or fitness, but next to nothing is known about causes and fitness consequences of feather wear in birds. We studied the effects of sex, age, year and experimental manipulation of brood size on primary feather wear in Collared Flycatchers Ficedula albicollis , and related variation in degree of feather wear to differences in fitness (viz. recruitment, survival). At the end of the breeding period, females and young birds had more worn flight feathers than males and adult birds, and the sexual difference in the degree of feather wear was particularly pronounced in one of the two study years. Experimental reduction of brood size reduced the degree of primary feather wear, whereas experimental enlargement of brood size did not lead to increased feather wear. In both sexes, there was a clear tendency for very old (>5 years old) birds to have more worn feathers than middle aged birds. The individual differences in the degree of feather wear were not correlated with individual differences in recruitment rate of young, but survival probability to the next breeding season increased with increasing degree of feather wear.     

Parental and chick responses of Laughing Gulls (Leucophaeus atricilla) to increase of flight costs and brood size

One main line of thought in life history theory is that investment in offspring must be balanced to minimize negative impacts on adult survival and future breeding. Seabirds have been regarded as fixed investors, although they exhibit a whole gradient of life history traits. We studied the consequences of brood size (one and three chicks) and of increased flight costs to one mate of a pair (3- and 5-cm trimming of the edge of the primary feathers) on parental response and on survival and body condition of chicks of Laughing Gulls (Leucophaeus atricilla). Parent gulls modified their nest attendance when their mate was handicapped, in a pattern dependent on the sex of the latter. Trimming of males affected chicks more severely than that of females. On its part, brood size affected amount of feeding sessions. Both chick body condition and survival were negatively affected by larger broods and by increased flight costs of one of their parents, especially when it was the male. Overall, parental inversion exhibited adjustments depending on the requirements of the brood and the fact that males compensated better the increased flying costs of their mates than vice versa. Despite a certain capacity by the males to compensate for the increased flight costs of the females, compensation was insufficient, and much less so in females, especially in larger broods, affecting chicks’ body condition and survival.     

Genetic and environmental components of growth in nestling blue tits (Parus caeruleus)

We investigated the effect of brood‐size mediated food availability on the genetic and environmental components of nestling growth in the blue tit (Parus caeruleus), using a cross‐fostering technique. We found genetic variation for body size at most nestling ages, and for duration of mass increase, but not of tarsus growth. Hence, nestling growth in our study population seems to have the potential to evolve further. Furthermore, significant genotype–environment interactions indicated heritable variation in reaction norms of growth rates and growth periods, i.e. that our study population had a heritable plasticity in the growth response to environmental conditions. The decreasing phenotypic variance with nestling age indicated compensatory growth in all body traits. Furthermore, the period of weight increase was longer for nestlings growing up in enlarged broods, while there was no difference to reduced broods in the period of tarsus growth. At fledging, birds in enlarged broods had shorter tarsi and lower weights than birds in reduced broods, but there was no difference in wing length or body condition between the two experimental groups. The observed flexibility in nestling growth suggests that growing nestlings are able to respond adaptively to food constraint by protecting the growth of ecologically important traits.     

Does sibling competition have a sex‐specific effect on offspring growth and development in the burying beetle Nicrophorus vespilloides?

During offspring growth, reduction in resource availability through competitive interactions will restrict how large individuals can become. Given that selection to mature at a large size will be greater for the sex in which large size provides the highest fitness return, sex‐specific differences in response to a decrease in resource availability may be expected. Using Nicrophorus vespilloides Herbst (Coleoptera: Silphidae) we examined the sex‐specific response of offspring to resource availability through sibling competition. We found that males and females were affected similarly by an increase in the level of sibling competition as brood size increased. Interestingly, although male N. vespilloides were consistently heavier than females, over a range of brood sizes, they were only significantly heavier than females at intermediate brood sizes. At present, the causes behind this finding remain unclear.     

Female fecundity and early offspring growth in the guppy, <Emphasis Type="Italic">Poecilia reticulata</Emphasis>

We examined the relationships between family (female parentage), body size of females, brood retention time between mating and parturition, female fecundity, and early growth of offspring in the guppy Poecilia reticulata. Mature, virgin females from a single brood were mated with a single male. Results of generalized linear models indicate that the effect of the family on female fecundity and offspring growth was significant, which suggested that these traits are genetically determined to a certain extent. Larger females at the time of mating produced larger broods, although female body size at the time of parturition did not affect brood size, in contrast to the results of some previous studies in guppies. Brood size was negatively associated with the body size of neonates. Results highlighted significant associations between brood retention time and female fecundity as well as offspring growth. In addition, the interaction between the family and brood retention time was significantly associated with female fecundity and offspring growth. Females of some families had longer retention times of larger broods, whereas those of other families had shorter retention times of smaller broods. On the other hand, females with longer brood retention times produced smaller neonates with slower growth. Since the family also affected the brood retention time, selection may work against the duration of brood retention of females via the size, growth and number of offspring, depending on environmental factors such as the intensity of predation or competition in neonates.     

Long-term consequences of early ontogeny in free-living Great Tits Parus major

Environmental factors during early development may have profound effects on subsequent life-history traits in many bird species. In wild birds, sex-specific effects of early ontogeny on natal dispersal and future reproduction are not well understood. The objective of this work was to determine whether hatching date and pre-fledging mass and condition of free-living Great Tits Parus major have any subsequent effect on individuals’ natal dispersal and reproductive performance at first breeding. Both males and females dispersed longer distances in coniferous than in deciduous forests, while dispersal was condition-dependent only in males (heavier as nestlings dispersed farther). In females, mass and condition at pre-fledging stage correlated significantly with clutch size, but not with subsequent reproductive performance as measured by fledging success or offspring quality. In contrast, heavier males as nestlings had higher future fledging success and heavier offspring in their broods compared with those in worse condition as nestlings. The hatching date of female as well as male parents was the only parental parameter related to the number of eggs hatched at first breeding. These results indicate that pre-fledging mass and condition predict subsequent fitness components in this bird species. We suggest that sex-specific relationships between a disperser’s condition and its selectivity with respect to breeding habitat and subsequent performance need to be considered in future models of life-history evolution.     

Consequences of elevating plasma testosterone in females of a socially monogamous songbird: evidence of constraints on male evolution?

To explore whether selection for testosterone-mediated traits in males might be constrained by costs of higher testosterone to females, we examined the effects of experimental elevation of plasma testosterone on physiological, reproductive, and behavioral parameters in a female songbird, the dark-eyed junco (Junco hyemalis). We used subcutaneous implants to elevate testosterone (T) in captive and free-living female juncos. In captive birds, we measured the effects of high T on body mass, feather molt, and brood patch formation. In the field, we monitored its effects on the timing of egg laying, clutch size, egg size, egg steroid levels, incubation, and nest-defense behavior. Females implanted with testosterone (T-females) had significantly higher circulating levels of testosterone than did control females (C-females). Captive T-females had lower body mass, were less likely to develop brood patches, and delayed feather molt relative to C-females. Among free-living females, the interval between nest completion and appearance of the first egg was longer for T-females than for C-females and egg yolk concentrations of testosterone were higher, but there were no significant differences in estradiol levels, clutch size, or egg size. Incubation and nest defense behavior were also similar between T- and C-females. Our results suggest that selection on males for higher testosterone might initially lead to a correlated response in females producing changes in body mass and feather molt, both of which could be detrimental. Other possible female responses would be delayed onset of reproduction, which might reduce reproductive success, and higher yolk testosterone, which might have either positive or negative effects on offspring development. We found no reason to expect reduced parental behavior by females as a negative fitness consequence of selection for higher testosterone in males.     

Unusual pattern of sex‐specific mortality in relation to initial brood sex composition in the black‐billed magpie Pica pica

In sexually size‐dimorphic species, brood sex composition may exert differential effects on sex‐specific mortality. We investigated the sex‐specific mortality and body condition in relation to brood sex composition in nestlings of the black‐billed magpie Pica pica. Neither significantly sex‐biased production at hatching nor overall sex‐biased mortality during the nestling period was found. Sex‐specific mortality as a function of brood sex composition, however, differed between female and male nestlings. We found higher mortality for females in male‐biased broods and higher mortality for males in female‐biased broods, a phenomenon that we call ‘rarer‐sex disadvantage’. As a result, fledging sex ratios became more biased in the direction of bias at hatching, a phenomenon that cannot be readily explained by previous hypotheses for sex‐specific mortality. Two temporal variables, fledging date and laying date, were also correlated with sex‐specific mortality: female nestlings in earlier broods experienced higher mortality than male nestlings whereas male nestlings in later broods experienced higher mortality. We suggest that this unusual pattern of mortality may be explained by adaptive adjustments of brood sex composition by parents, either through the effects of a slight sex difference in offspring dispersal patterns on parental fitness, or owing to sex differences as regards the benefits of early fledging.     

Maternal and paternal effects on offspring phenotype in the dung beetle Onthophagus taurus

Abstract.— Parents often have important influences on the development of traits in their offspring. One mechanism by which parents are able to influence offspring phenotype is through the level of care they provide. In onthophagine dung beetles, parents typically provision their offspring by packing dung fragments into a brood mass. Onthophagus taurus males can be separated into two discrete morphs: Large, "major" males have head horns, whereas "minor" males are hornless. Here we show that a switch in parental provisioning strategies adopted by males coincides with the switch in male morphology. Male provisioning results in the production of heavier brood masses than females will produce alone. However, unlike females in which the level of provisioning increases with body size in a continuous manner, the level of provisioning provided by males represents an "all-or-none" tactic with all major males providing a fixed level of provisioning irrespective of their body size. Offspring size is determined largely by the quantity of dung provided to the developing larvae so that paternal and maternal provisioning affects the body size and horn size of offspring produced. The levels of provisioning by individual parents are significantly repeatable, suggesting paternal and maternal effects as candidate indirect genetic effects in the evolution of horn size in the genus Onthophagus .     

Sexual dimorphism of body and relative head sizes in neonatal common garter snakes

Morphological and behavioral differences between sexes are commonplace throughout the animal kingdom. Body size is one of the most obvious sex differences frequently found in snakes. However, the developmental origins of size differences in many species, including snakes, are not well known. We examined post-natal variation in sexual size dimorphism in garter snakes Thamnophis sirtalis . The weights, body and tail lengths, and head sizes of male and female neonates born to mothers collected from ecologically dissimilar habitats on Beaver Island, Lake Michigan were compared. Sexual size dimorphism was prominent. Overall, males had significantly longer bodies and tails than females. Females were significantly heavier and had larger heads than male snakes. Maternal site affected head but not body measurements, perhaps due to differences in prey availability. The body condition of maternal females predicted neonatal body length. Significant litter variation suggests heritable variation in morphological traits possibly correlated with feeding success and survival.     

Do Seaducks Minimise the Flightless Period?: Inter- and Intra-Specific Comparisons of Remigial Moult

Remigial moult is one of the crucial events in the annual life cycle of waterfowl as it is energetically costly, lasts several weeks, and is a period of high vulnerability due to flightlessness. In waterfowl, remigial moult can be considered as an energy-predation trade-off, meaning that heavier individuals would minimise the flightless period by increasing feather growth rate and energy expenditure. Alternatively, they could reduce body mass at the end of this period, thereby reducing wing-loading to increase flight capability. We studied timing of remigial moult, primary growth rates, flightlessness duration, and the pattern of body mass variation in 5 species of captive seaducks (Melanitta fusca, M. perspicillata, Clangula hyemalis, Histrionicus histrionicus, and Somateria mollissima) ranging in size from 0.5 to 2.0 kg. Their feather growth rates weakly increased with body mass (M^0.059) and no correlation was found at the intra-specific level. Consequently, heavier seaduck species and especially heavier individuals had a longer flightless period. Although birds had access to food ad libidum, body mass first increased then decreased, the latter coinciding with maximum feather growth rate. Level of body mass when birds regained flight ability was similar to level observed at the beginning of remigial moult, suggesting they were not using a strategic reduction of body mass to reduce the flightlessness duration. We suggest that the moulting strategy of seaducks may be the result of a compromise between using an intense moult strategy (simultaneous moult) and a low feather growth rate without prejudice to feather quality. Despite the controlled captive status of the studied seaducks, all five species as well as both sexes within each species showed timing of moult reflecting that of wild birds, suggesting there is a genetic component acting to shape moult timing within wild birds.