Experimentally reduced male attractiveness increases parental care in the pied flycatcher Ficedula hypoleuca

This study reports effects of experimental manipulations ofreproductive effort and the size of the male's white foreheadpatch (a secondary sexual trait), on provisioning rates, reproductivesuccess, and parental breeding dispersal distance in the piedflycatcher, Ficedula hypoleuca. Parents caring for enlargedbroods resulting from manipulated clutches provisioned nestsat higher rates than parents with reduced broods. Males with a reduced forehead patch fed their nestlings more in relationto males with an unmanipulated forehead patch, and their youngfledging with a longer tarsi. This suggests that males witha reduced attractiveness may perceive their own attractivenessand they devote more time available for parental effort given their poorer prospects in male contest competition and/or femaleattraction for extra-pair copulations. However, their femalesdid not alter their provisioning effort and this runs counterto both the differential allocation and the partner-compensationhypotheses. An artificial decrease in a male secondary sexualtrait led to a wider breeding dispersal distance between successiveyears.     

Reduced breeding success of Pied Flycatchers Ficedula hypoleuca along regulated rivers

Most large rivers in northern Sweden are regulated to produce hydropower, with subsequent effects on flow dynamics and aquatic insect communities. Several studies have shown that aquatic and terrestrial systems are intimately connected via the export of emergent aquatic insects, but few have assessed how human modifications of aquatic habitats may influence this connection. We compared breeding success of the insectivorous Pied Flycatcher Ficedula hypoleuca in near‐riparian upland forests along two regulated and two free‐flowing large rivers in northern Sweden over 3 years. The regulated rivers showed lower aquatic insect export to the surroundings, as a consequence of regulation‐induced loss of suitable aquatic insect habitats. Survival of Pied Flycatcher nestlings was 10–15% greater along the free‐flowing rivers. Females breeding near the free‐flowing rivers also started egg‐laying earlier and with greater synchrony than those at the regulated rivers, and showed a smaller decrease in weight during breeding than did females along the regulated rivers. However, there were no differences in occupation rate, clutch size or number of successfully hatched juveniles between regulated and free‐flowing rivers. As regulated rivers showed lower abundance of flying aquatic insects, which may also reduce the abundance of terrestrial invertebrate prey, regulation‐induced changes in the export of emergent aquatic insects may explain both directly and indirectly the observed reduction in Pied Flycatcher breeding success along regulated rivers. Large‐scale river regulation may therefore impair the breeding success of insectivorous birds through impacts on prey availability.     

Cross‐fostering eggs reveals that female collared flycatchers adjust clutch sex ratios according to parental ability to invest in offspring

Across animal taxa, reproductive success is generally more variable and more strongly dependent upon body condition for males than for females; in such cases, parents able to produce offspring in above‐average condition are predicted to produce sons, whereas parents unable to produce offspring in good condition should produce daughters. We tested this hypothesis in the collared flycatcher (Ficedula albicollis) by cross‐fostering eggs among nests and using the condition of foster young that parents raised to fledging as a functional measure of their ability to produce fit offspring. As predicted, females raising heavier‐than‐average foster fledglings with their social mate initially produced male‐biased primary sex ratios, whereas those raising lighter‐than‐average foster fledglings produced female‐biased primary sex ratios. Females also produced male‐biased clutches when mated to males with large secondary sexual characters (wing patches), and tended to produce male‐biased clutches earlier within breeding seasons relative to females breeding later. However, females did not adjust the sex of individuals within their clutches; sex was distributed randomly with respect to egg size, laying order and paternity. Future research investigating the proximate mechanisms linking ecological contexts and the quality of offspring parents are able to produce with primary sex‐ratio variation could provide fundamental insight into the evolution of context‐dependent sex‐ratio adjustment.     

Evolution of parental incubation behaviour in dinosaurs cannot be inferred from clutch mass in birds

A recent study proposed that incubation behaviour (i.e. type of parental care) in theropod dinosaurs can be inferred from an allometric analysis of clutch volume in extant birds. However, the study in question failed to account for factors known to affect egg and clutch size in living bird species. A new scaling analysis of avian clutch mass demonstrates that type of parental care cannot be distinguished by conventional allometry because of the confounding effects of phylogeny and hatchling maturity. Precociality of young but not paternal care in the theropod ancestors of birds is consistent with the available data.     

Patterns of natural selection on morphology of male and female collared flycatchers (Ficedula albicollis)

Natural selection may act in different directions among years, life stages, or classes of individuals. Fluctuating selection of this kind is potentially an important mechanism by which additive genetic variation for quantitative traits is maintained, and can prevent populations reaching local adaptive peaks. We analysed natural selection acting on three morphological traits of male and female collared flycatchers via both fecundity and survival, using 15 years' data from a large isolated population on Godand, Sweden. We particularly investigated variation in the direction and magnitude of selection acting: (1) among years over the study period; (2) on different life stages and (3) the consistency of observed patterns of selection with sexual size dimorphism (SSD) in this population. We found little evidence of natural selection on these traits over the study period. Evidence for directional, stabilizing and disruptive selection was found for some year-trait combinations, but these patterns were inconsistent with respect to both the magnitude and form of selection found. Consequently, our results, based on the detailed analysis of natural selection in a large wild population over a period of 15 years, provide evidence for the common assumption that forces of selection acting on quantitative traits are generally weak. They are also consistent with the suggestion that environmental stability is an important determinant of the degree to which organisms fit their environment.     

Triparental care in the collared flycatcher (Ficedula albicollis): Cooperation of two females with a cuckolded male in rearing a brood

Certain predominant forms of mating and parental care systems are assumed in several model species among birds, but the opportunistic and apparently infrequent variations of “family structures” may often remain hidden due to methodological limitations with regard to genetic or behavioral observations. One of the intensively studied model species, the collared flycatcher (Ficedula albicollis), is usually characterized by social monogamy with polyterritorial, facultative social polygyny, and frequent extrapair mating and extrapair paternity. During a brood‐size manipulation experiment, we observed two females and a male delivering food at an enlarged brood. A combination of breeding phenology data (egg laying and hatching date), behavioral data (feeding rates) from video recordings at 10 days of nestling age, and microsatellite genotyping for maternity and paternity suggests a situation of an unrelated female helping a pair in chick rearing. Such observations highlight the relevance of using traditional techniques and genetic analyses together to assess the parental roles within a population, which becomes more important where individuals may dynamically switch from their main and presupposed roles according to the actual environmental conditions.     

Red coloration and male parental behaviour in the threespine stickleback

In an anadromous population of the threespine stickleback, male red coloration was correlated with agonistic and egg care behaviour of parental males, but not with hatching success. When one male took over a neighbour's territory, the dominant male had more intense red coloration.     

The male and female perspective in the link between male infant care and mating behaviour in Barbary macaques

Infant care from adult males is unexpected in species with high paternity uncertainty. Still, males of several polygynandrous primates engage in frequent affiliative interactions with infants. Two non‐exclusive hypotheses link male infant care to male mating strategies. The paternal investment hypothesis views infant care as a male strategy to maximize the survival of sired offspring, while the mating effort hypothesis predicts that females reward males who cared for their infant by preferably mating with them. Both hypotheses predict a positive relationship between infant care and matings with a particular female. However, the paternal investment hypothesis predicts that increased matings come before infant care whereas the mating effort hypothesis predicts that infant care precedes an increase in matings. Both hypotheses are usually tested from the perspective of the proportion of matings and care that individual females engage in and receive, rather than from the perspective of the care and mating behaviour of individual males. We tested the relationships between care and mating from both female and male perspectives in Barbary macaques. Mating predicted subsequent care and care predicted subsequent mating when viewed from the male but not the female perspective. Males mainly cared for infants of their main mating partners, but infants were not mainly cared for by their likely father. Males mated more with the mothers of their favourite infants, but females did not mate more with the main caretakers of their infants. We suggest that females do not choose their mating partners based on previous infant care, increasing paternity confusion. Males might try to increase paternal investment by distributing the care according to their own instead of female mating history. Further, males pursue females for mating opportunities based on previous care.     

High brood patch temperature of less colourful,less pheomelanic female Barn Swallows Hirundo rustica

In birds, even a minor difference in egg temperature (1–1.5 °C) has been shown to affect the fitness of offspring by changing hatching success, incubation period and nestling quality. Female, but not male, passerines develop brood patches. Thus if there are traits, such as plumage ornamentation, that indicate optimal egg temperature, males should pair with females that exhibit those traits. However, no study has yet investigated the relationship between female brood patch temperature, which would directly affect egg temperature, and female plumage ornamentation. In this study, we examined the surface temperature of female brood patches during nocturnal incubation and examined its relationship with female plumage ornaments in Asian Barn Swallows Hirundo rustica gutturalis. After controlling for ambient air temperature, brood patch temperature was negatively associated with colour saturation of the female throat patch. No other female ornaments, such as tail‐length, white tail spots or throat patch size, predicted brood patch temperature. When oral (mouth) temperature was statistically controlled, females with less colourful throats and longer tails showed higher brood patch temperature, indicating that these females had hotter brood patches in relation to the temperature of other body parts. Furthermore, we found a negative relationship between pheomelanin pigmentation and brood patch temperature after controlling for ambient air temperature or oral temperature. To our knowledge, this is the first study to show that female ornaments can predict the absolute/relative thermal investment in brood patches. This relationship, together with other aspects of female quality, may affect male mate preference and female ornamentation.     

Adaptation to monogamy influences parental care but not mating behavior in the burying beetle,Nicrophorus vespilloides

The mating system is expected to have an important influence on the evolution of mating and parenting behaviors. Although many studies have used experimental evolution to examine how mating behaviors evolve under different mating systems, this approach has seldom been used to study the evolution of parental care. We used experimental evolution to test whether adaptation to different mating systems involves changes in mating and parenting behaviors in populations of the burying beetle, Nicrophorus vespilloides. We maintained populations under monogamy or promiscuity for six generations. This manipulation had an immediate impact on reproductive performance and adult survival. Compared to monogamy, promiscuity reduced brood size and adult (particularly male) survival during breeding. After six generations of experimental evolution, there was no divergence between monogamous and promiscuous populations in mating behaviors. Parents from the promiscuous populations (especially males) displayed less care than parents from the monogamous populations. Our results are consistent with the hypothesis that male care will increase with the certainty of paternity. However, it appears that this change is not associated with a concurrent change in mating behaviors.     

Flexible parents: joint effects of handicapping and brood size manipulation on female parental care in Nicrophorus vespilloides

Parental care is highly variable, reflecting that parents make flexible decisions in response to variation in the cost of care to themselves and the benefit to their offspring. Much of the evidence that parents respond to such variation derives from handicapping and brood size manipulations, the separate effects of which are well understood. However, little is known about their joint effects. Here, we fill this gap by conducting a joint handicapping and brood size manipulation in the burying beetle Nicrophorus vespilloides. We handicapped half of the females by attaching a lead weight to their pronotum, leaving the remaining females as controls. We also manipulated brood size by providing each female with 5, 20 or 40 larvae. In contrast to what we predicted, handicapped females spent more time provisioning food than controls. We also found that handicapped females spent more time consuming carrion. Furthermore, handicapped females spent a similar amount of time consuming carrion regardless of brood size, whereas controls spent more time consuming carrion as brood increased. Females spent more time provisioning food towards larger broods, and females were more likely to engage in carrion consumption when caring for larger broods. We conclude that females respond to both handicapping and brood size manipulations, but these responses are largely independent of each other. Overall, our results suggest that handicapping might lead to a higher investment into current reproduction and that it might be associated with compensatory responses that negate the detrimental impact of higher cost of care in handicapped parents.     

Risk taking during parental care: a test of the harm-to-offspring hypothesis

Amount of risk taking during parental care is often explainedin relation to the reproductive value of the offspring. The"harm-to-offspring hypothesis" focuses on the relative harma period of no parental care can do to the offspring. Accordingto this hypothesis, parents should take greater risks for offspringin poor condition than for offspring in good condition. We manipulatedoffspring condition in the pied flycatcher (Ficedula hypoleuca)and tested the harm-to-offspring hypothesis by exposing parentsto a predator model (a sparrowhawk, Accipiter nisus). Time elapseduntil a parent first entered the nest-box was used as a risk-takingmeasure. Parents spent significantly shorter time until first nestvisit for offspring in poor condition than for offspring ingood condition. Hence, the harm-to-offspring hypothesis wassupported.     

Factors influencing tadpole deposition site choice in a frog with male parental care: An experimental field study

Parents have evolved a variety of strategies to minimize risks to their offspring, including complex choices regarding suitable rearing sites, based on abiotic and biotic factors, which differentially affect offspring survival. Because availability and quality of these sites are variable, parents may have to choose between immediately available lower‐quality rearing sites or extended search time. In some frog species with larval transport, parents are known to select bodies of water that are free of predators, cannibalistic tadpoles, or intraspecific competitors for larval deposition and rearing sites. We tested whether abiotic factors and the presence of predators and conspecific tadpoles affect tadpole deposition behavior in a population of smooth guardian frog, Limnonectes palavanensis, on the island of Borneo. Females lay eggs on land and males guard them until they hatch; after hatching, tadpoles climb onto the male's back and are subsequently transported to small pools of water on the forest floor, which are scarce and patchily distributed. We estimated the abundance of natural tadpole rearing sites in our study area and conducted experiments using artificial pools to test whether abiotic characteristics of these pools affect the probability of larval deposition. We also performed choice experiments to test whether males of L. palavanensis avoid pools with conspecific tadpoles or predators. Lastly, we tested whether the tadpoles of this species exhibit cannibalism. The abundance of natural deposition sites was low, and males readily used artificial pools for tadpole deposition. Males were less likely to deposit tadpoles in artificial pools located in steep areas, and males did not avoid depositing tadpoles in pools with conspecifics or with experimentally introduced predators. Males exhibited clutch‐partitioning behavior, dividing tadpoles between adjacent artificial pools. Pool availability, rather than the presence of potential competitors or predators in a pool, affects tadpole deposition decisions in this species.     

The origin of parental care in relation to male and female life history

The evolution of maternal, paternal, and bi‐parental care has been the focus of a great deal of research. Males and females vary in basic life‐history characteristics (e.g., stage‐specific mortality, maturation) in ways that are unrelated to parental investment. Surprisingly, few studies have examined the effect of this variation in male and female life history on the evolution of care. Here, we use a theoretical approach to determine the sex‐specific life‐history characteristics that give rise to the origin of paternal, maternal, or bi‐parental care from an ancestral state of no care. Females initially invest more into each egg than males. Despite this inherent difference between the sexes, paternal, maternal, and bi‐parental care are equally likely when males and females are otherwise similar. Thus, sex differences in initial zygotic investment do not explain the origin of one pattern of care over another. However, sex differences in adult mortality, egg maturation rate, and juvenile survival affect the pattern of care that will be most likely to evolve. Maternal care is more likely if female adult mortality is high, whereas paternal care is more likely if male adult mortality is high. These findings suggest that basic life‐history differences between the sexes can alone explain the origin of maternal, paternal, and bi‐parental care. As a result, the influence of life‐history characteristics should be considered as a baseline scenario in studies examining the origin of care.     

Brood provisioning rates and fledgling behavior of Cordilleran Flycatchers in southwestern Colorado

The behavior of young songbirds after fledging is one of the least understood phases of the breeding cycle, although parental provisioning rates and movement of fledglings are key to understanding life history evolution. We studied Cordilleran Flycatchers (Empidonax occidentalis) at two sites in southwestern Colorado, USA, from 2012 to 2017. We banded and sexed breeding adults to determine the relative contributions of males and females to nestling and fledgling care, and attached radio‐transmitters to nestlings to facilitate observations of brood behavior after fledging. Females made 60% and 78% of total observed feedings of nestlings and fledglings, respectively. Parental provisioning rates increased with nestling age, and per‐nestling provisioning rates increased with brood size. Parental provisioning rates declined just before fledging, then increased just after fledging. Fledging times of individuals in broods were asynchronous and concentrated during the late afternoon and early evening. Males stopped caring for fledglings before females even though this species is single‐brooded, with some late‐season broods being abandoned by males. Broods spent the first three weeks after fledging within 400 m of nests, after which they began to disperse. Most aspects of the breeding biology of Cordilleran Flycatchers in our study, including the duration of nestling and fledging periods, female‐dominated provisioning, and movement patterns of fledglings, were similar to those of other Empidonax species. However, the times when young fledged were not concentrated in the morning as reported in most other songbirds, and this result warrants additional study of the timing of fledging in ecologically and taxonomically similar species. The increased per‐nestling provisioning rate with increasing brood size was unexpected, and additional study is needed to determine if this increase results from a trade‐off between adult annual survival and productivity favoring increased provisioning of young in larger broods, or from the existence of high‐quality individuals where larger clutches and higher provisioning rates are linked.     

Nest survival is influenced by parental behaviour and heterospecifics in a mixed‐species colony

Studies of avian nest success often focus on examining influences of variation in environmental and seasonal factors. However, in‐depth evaluations can also incorporate variation in individual incubation behaviour to further advance our understanding of avian reproductive ecology. We examined these relationships in colonially nesting Black‐crowned Night‐Herons Nycticorax nycticorax using intensive video‐monitoring methods to quantify incubation behaviours. We modelled nest survival as a function of both extrinsic factors and incubation behaviours over a 3‐year period (2010–12) on Alcatraz Island, USA. Model‐averaged parameter estimates indicated that nest survival increased as a function of greater incubation constancy (% of time spent incubating eggs within a 24‐h period), and average daily precipitation throughout the nesting stage. Common Ravens Corvus corax are the only known nest predator of Night‐Herons on Alcatraz Island, as on many other coastal Pacific islands. We also investigated the effects of heterospecific nesting of California Gulls Larus californicus and Western Gulls Larus occidentalis in a mixed‐species colony with Night‐Herons, based on nesting proximity data collected over a 2‐year period (2011–12). This second analysis indicated that, in addition to incubation behaviours, nesting heterospecifics are an important factor for explaining variation in Night‐Heron nest survival. However, contrary to our original expectation, we found that Night‐Herons experienced increased nest survival with increasing distance from gull colony boundaries. These results may apply to other areas with multiple colonial nesting species and similar predator communities and climatic patterns.     

Division of labour during incubation in a woodpecker Colaptes auratus with reversed sex roles and facultative polyandry

The contribution of males to incubation has rarely been studied in altricial birds because the pursuit of extra mating opportunities is believed to conflict with incubation. Woodpeckers show reversed sex roles in parental care with males doing most of the nest construction, incubating and brooding of the young while females may be polyandrous. I investigated incubation by each sex at 71 monogamous and four polyandrous nests of the Northern Flicker Colaptes auratus , predicting that males would contribute to incubation according to their energy reserves (body condition) whereas females would contribute based on alternate reproductive opportunities. Nest attendance was 99% with males contributing a mean of 66% of the total incubation including nocturnal incubation. The length of daytime bouts averaged about 2 h and did not differ between the sexes. Consistent with predictions of investment strategies, structurally larger males and those in poorer body condition incubated less than smaller males, perhaps because they required more recess time to forage or to conserve energy. Older females contributed less incubation than young females and polyandrous females contributed less incubation at their secondary nests than monogamous females. Incubation period, nest depredation rate and hatching success were not influenced by bout length, number of bouts or relative contribution of the sexes. Hatching success was 86% at nests of both monogamous and polyandrous females because males compensated for reduced female participation. Because incubation of the sexes is compensatory and not additive, incubation pattern did not influence short-term reproductive success. I conclude that males invest in incubation according to their energy needs, and females may adjust their contributions based on alternate reproductive tactics.     

Fluctuations in incubation temperature affect incubation duration but not morphology,locomotion and growth of hatchlings in the sand lizard Lacerta agilis (Lacertidae)

Li, H., Zhou, Z.‐S., Ding, G.‐H. and Ji, X. 2011. Fluctuations in incubation temperature affect incubation duration but not morphology, locomotion and growth of hatchlings in the sand lizard Lacerta agilis (Lacertidae). —Acta Zoologica (Stockholm) 00 : 1–8. Studies looking for potential effects of temperature and temperature fluctuations on phenotypic traits of reptile hatchlings have shown species variation, but have not always allowed a distinction between effects of fluctuation per se and temperature extremes themselves. To examine whether incubation temperature fluctuation has a key role in influencing the phenotype of offspring, we incubated eggs of the sand lizard Lacerta agilis at one of the four temperature regimes (27, 27 ± 2, 27 ± 4 and 27 ± 6 °C). We found that: (1) hatchlings incubated under the four temperature regimes did not differ from each other in any of the morphological and physiological traits examined; (2) interactions that included temperature treatment did not affect any trait examined; (3) the mean incubation length was longer in the 27 ± 6 °C treatment than in the other three treatments; and (4) female hatchlings were shorter in head length and width but longer in snout‐vent length as well as abdomen length than males derived from the same‐sized egg. Our data show that both the type and the magnitude of temperature variation can affect incubation length. We found no evidence for phenotypic divergence in responses to temperature fluctuations during incubation and therefore suggest that temperature variation does not affect the phenotype of hatchlings in L. agilis.