Family break-up in Black and Red Kites Milvus migrans and M. milvus: is time of independence an offspring decision?

I observed the natural process of family break-up in 13 families of Black Kites Milvus migrans and five families of Red Kites M. milvus in which fledglings had been individually marked. In other broods, I performed experiments which modified the parental investment fledglings received by supplementing nests of both species with food or transferring Black Kite chicks to nests with younger or older chicks of the same species.
The time of family break-up in the Black Kite is mainly an offspring decision which is not affected by an artificial increase of parental investment. The duration of the post-fledging period was not increased in Black Kites that were given supplementary food. Chicks transferred to nests with a younger chick did not extend the post-fledging period, nor did chicks transferred to nests with an older chick shorten the post-fledging period. In Red Kites, parental investment seemed to have more influence on the timing of the family break-up. Red Kite adults invested less as parents than did Black Kites during the post-fledging dependence period, and in nests where supplementary food was given, Red Kite fledglings stayed attached for a longer period.     

Females compensate for moult‐associated male nest desertion in Hooded Warblers

Uniparental offspring desertion occurs in a wide variety of avian taxa and usually reflects sexual conflict over parental care. In many species, desertion yields immediate reproductive benefits for deserters if they can re‐mate and breed again during the same nesting season; in such cases desertion may be selectively advantageous even if it significantly reduces the fitness of the current brood. However, in many other species, parents desert late‐season offspring when opportunities to re‐nest are absent. In these cases, any reproductive benefits of desertion are delayed, and desertion is unlikely to be advantageous unless the deserted parent can compensate for the loss of its partner and minimize costs to the current brood. We tested this parental compensation hypothesis in Hooded Warblers Setophaga citrina, a species in which males regularly desert late‐season nestlings and fledglings during moult. Females from deserted nests effectively doubled their provisioning efforts, and nestlings from deserted nests received just as much food, gained mass at the same rate, and were no more likely to die from either complete nest predation or brood reduction as young from biparental nests. The female provisioning response, however, was significantly related to nestling age; females undercompensated for male desertion when the nestlings were young, but overcompensated as nestlings approached fledging age, probably because of time constraints that brooding imposed on females with young nestlings. Overall, our results indicate that female Hooded Warblers completely compensate for male moult‐associated nest desertion, and that deserting males pay no reproductive cost for desertion, at least up to the point of fledging. Along with other studies, our findings support the general conclusion that late‐season offspring desertion is likely to evolve only when parental compensation by the deserted partner can minimize costs to the current brood.     

Nest as an extended phenotype signal of female quality in the great reed warbler

Extended phenotypes with signalling function are mostly restricted to animal taxa that use construction behaviour during courtship displays. However, they can be used also as post‐mating signals of mate quality, allowing individuals to obtain reliable information about their partners. Nest size may have such a signalling function and a lot of indirect evidence supports this view. However, direct evidence based on an experimental approach is still widely missing. Here we test the role of nest size in post‐mating signalling of mate quality in the great reed warbler Acrocephalus arundinaceus, a passerine with female‐restricted nest‐building behaviour. Based on observational data, clutch size, nestling weight, brood size and fledglings’ propensity to return to their natal site positively correlated with nest size. Moreover, we experimentally enlarged great reed warbler nests to investigate whether this manipulation affects male investment in feeding. We found that males fed their nestlings significantly more intensively on enlarged nests than those on control nests. This suggests that nest size in this species serves as a signal of female quality or willingness to invest in reproduction and that it pays males to enhance their feeding effort according to this signal. Thus, we provide convincing evidence that animal communication takes place through the extended phenotypes and that post‐mating signalling of quality is not restricted only to males, but may function equally well in females.     

The design and function of birds' nests

All birds construct nests in which to lay eggs and/or raise offspring. Traditionally, it was thought that natural selection and the requirement to minimize the risk of predation determined the design of completed nests. However, it is becoming increasingly apparent that sexual selection also influences nest design. This is an important development as while species such as bowerbirds build structures that are extended phenotypic signals whose sole purpose is to attract a mate, nests contain eggs and/or offspring, thereby suggesting a direct trade‐off between the conflicting requirements of natural and sexual selection. Nest design also varies adaptively in order to both minimize the detrimental effects of parasites and to create a suitable microclimate for parents and developing offspring in relation to predictable variation in environmental conditions. Our understanding of the design and function of birds' nests has increased considerably in recent years, and the evidence suggests that nests have four nonmutually exclusive functions. Consequently, we conclude that the design of birds' nests is far more sophisticated than previously realized and that nests are multifunctional structures that have important fitness consequences for the builder/s.     

Parental Care and Mate Choice in the Giant Water Bug Belostoma lutarium

Parental care and sexual selection are highly interrelated. Understanding the evolution of sex‐specific patterns of parental care and sexual selection is a major focus of current evolutionary ecology research and requires empirical studies that simultaneously quantify components of both parental care and sexual selection in a single species. In this study, we quantify the dynamics of paternal care and sexual selection in the giant water bug Belostoma lutarium. Specifically, we examined (1) which sex potentially experiences sexual selection, (2) which traits, if any, are associated with attaining a mate by males and/or females (i.e. which traits are potentially under selection), and (3) which male and female traits, if any, relate to paternal care and offspring survival. Our findings suggest that (1) males are likely the choosier sex and that heavier females are more likely to mate than smaller females, (2) that female body weight is under selection if female weight is a trait that is stable within a given individual and (3) body size is sexually dimorphic, with females being the larger sex in this species. There was no evidence of male or female traits being linked to offspring survival in this species, although this is potentially due to the lack of egg predators in our study. We discuss our findings in relation to the evolution of sex roles and future avenues of research in this species.     

Components of care: are they honest signals of parental care quality?

It has been argued that male parental care provides direct benefits to females and therefore should be under sexual selection. Given this, we expect signals that honestly indicate the quality of care to be favoured by selection. One such potential signal is care itself. Fish have several features that make them excellent model systems for studying the evolution and dynamics of parental care. We use the flagfish, Jordanella floridae , as a model to evaluate these general ideas. Males of this species guard, clean and fan empty nests and then eggs. Females prefer males that fan more (1) before spawning and (2) when eggs are newly received. When single males and females were paired, males that fanned and visited their nests more prior to spawning were more likely to be mated. Furthermore, among successful males, rates of fanning in the first day after spawning were correlated with the number of eggs received in the future (but not current egg numbers). We then considered whether these two putative signals were correlated and whether males that fan more in these contexts actually have higher egg survivorship. We found no correlation between nest fanning rates before and after spawning and neither 'signal' was predictive of variation in egg survivorship among mated males. We further considered whether pre‐spawning fanning rates were predictive of hatching success in an experiment in which single males were allowed to establish nests and provided eggs. We found little evidence that fanning is an honest signal of care quality and discuss alternative explanations. In particular, we discuss patterns of care elaboration in light of our results.     

A test of the endogenous and exogenous selection hypotheses for the maintenance of a narrow avian hybrid zone

The contributions of genetic and environmental factors to differential reproductive success across hybrid zones have rarely been tested. Here, we report a manipulative experiment that simultaneously tested endogenous (genetic-based) and exogenous (environmental-based) selection within a hybrid zone. We transplanted mated pairs of two chickadee species (Poecile atricapilla and P. carolinensis) and their hybrids into isolated woodlots within their hybrid zone and monitored their reproductive success. Although clutch sizes were similar, based on an estimate of the genetic compatibility of a pair, hybrid pairs produced fewer nestlings and fledglings than did pairs of either parental species. According to a linear model generated from the data, a pure pair of either parental species would be expected to produce 1.91-2.48 times more fledglings per nesting attempt, respectively, than the average or least compatible pair in the experiment. Our result of decreased reproduction for hybrid pairs relative to parental species pairs within same environment (the hybrid zone in this experiment) support the endogenous selection hypothesis for maintenance of this hybrid zone. Because the experiment was conducted entirely within the hybrid zone (i.e., the same environment for parental and hybrid pairings), our data do not support the exogenous selection hypothesis as it predicts either all pairings doing poorly or the hybrid pairs more successful than the parental pairs.     

The post‐fledging period in a tropical bird: patterns of parental care and survival

How environmental conditions affect the timing and extent of parental care is a fundamental question in comparative studies of life histories. The post‐fledging period is deemed critical for offspring fitness, yet few studies have examined this period, particularly in tropical birds. Tropical birds are predicted to have extended parental care during the post‐fledging period and this period may be key to understanding geographic variation in avian reproductive strategies. We studied a neotropical passerine, the western slaty‐antshrike Thamnophilus atrinucha, and predicted greater care and higher survival during the post‐fledging period compared to earlier stages. Furthermore, we predicted that duration of post‐fledging parental care and survival would be at the upper end of the distribution for Northern Hemisphere passerines. Correspondingly, we observed that provisioning continued for 6–12 weeks after fledging. In addition, provisioning rate was greater after fledging and offspring survival from fledging to independence was 75%, greater than all estimates from north‐temperate passerines. Intervals between nesting attempts were longer when the first brood produced successful fledglings compared to nests where offspring died either in the nest or upon fledging. Parents delayed initiating second nests after the first successful brood until fledglings were near independence. Our results indicate that parents provide greater care after fledging and this extended care likely increased offspring survival. Moreover, our findings of extended post‐fledging parental care and higher post‐fledging survival compared to Northern Hemisphere species have implications for understanding latitudinal variation in reproductive effort and parental investment strategies.     

Video nest monitoring reveals male coloration-dependant nest predation and sex differences in prey size delivery in a bird under high sexual selection

Increased predation risk should select for reduced parental activity to decrease the probability of visually hunting predators discovering the nest. Parental activity and conspicuousness are known to increase predation risk. Here, we test for sex differences in parental visitation rate (number of visits), time seen at the nest (time at the nest × adult visibility), and food delivery (prey size) using continuous video recordings at nests. We test the role of these variables for predation outcome in the Superb Fairy-wren (Malurus cyaneus). The study species is sexually dimorphic: males have iridescent blue plumage while females have brown plumage. The results showed that nest predation was predicted by male time seen at the nest (but not visitation rate), but not female time seen at the nest (or visitation rate). Contrary to our expectation that males would have lower visitation rates than females, our analysis of video images showed that male and female visitation was comparable but that males consistently brought smaller prey items to the nest than females. These findings are discussed in the light of morphological differences between male and female beak size. We conclude that sexual selection has favoured conspicuous male signalling in this system, and that natural selection should select for reduced parental care for the conspicuous sex.     

Effects of extra-pair paternity and maternity on the provisioning strategies of the Azure-winged Magpie Cyanopica cyanus

Altricial birds show enormous intraspecific diversity in their provisioning strategies, in terms of both the provisioning rate and the amount of food delivered per feeding bout. Extra-pair copulations (EPCs), which result in either extra-pair paternity (EPP) or maternity (EPM), provide an opportunity to demonstrate why provisioning strategies vary among individuals. Because EPP-cuckolded males and EPM-cuckolded females must raise unrelated young, whereas EPM-cuckolded males and EPP-cuckolded females need not, we hypothesized that the first two categories of breeders would reduce parental effort, whereas the latter two categories of breeders would increase parental effort. We tested this hypothesis in the Azure-winged Magpie Cyanopica cyanus by comparing the number and body mass of fledglings, provisioning rates and food amount delivered per feeding bout between EPP- and EPM-cuckolded breeders and faithful breeders. We found that (1) the number of fledglings did not differ significantly between cuckolded and faithful breeders, and (2) fledglings raised by EPM-cuckolded males and EPP-cuckolded females did not differ from faithful breeders, whereas fledglings raised by EPP-cuckolded males and EPM-cuckolded females were significantly smaller than those raised by faithful breeders. Compared with faithful breeders, cuckolded breeders increased parental efforts in nests that contained no unrelated young; hence, their loss in parentage may be compensated for by the enhanced quality of fledglings. In nests that contained unrelated young, cuckolded breeders did not reduce parental efforts; hence, their own offspring would not starve and could survive in competition with mixed brood-mates. Our findings suggest that the differences in parentage created by EPP and EPM scenarios and the potential fitness return of raising a brood for the cuckolded breeders can explain the intraspecific variation in provisioning strategies of altricial birds.     

Reconstructing paternal genotypes to infer patterns of sperm storage and sexual selection in the hawksbill turtle

Postcopulatory sperm storage can serve a range of functions, including ensuring fertility, allowing delayed fertilization and facilitating sexual selection. Sperm storage is likely to be particularly important in wide‐ranging animals with low population densities, but its prevalence and importance in such taxa, and its role in promoting sexual selection, are poorly known. Here, we use a powerful microsatellite array and paternal genotype reconstruction to assess the prevalence of sperm storage and test sexual selection hypotheses of genetic biases to paternity in one such species, the critically endangered hawksbill turtle, Eretmochelys imbricata. In the majority of females (90.7%, N = 43), all offspring were sired by a single male. In the few cases of multiple paternity (9.3%), two males fertilized each female. Importantly, the identity and proportional fertilization success of males were consistent across all sequential nests laid by individual females over the breeding season (up to five nests over 75 days). No males were identified as having fertilized more than one female, suggesting that a large number of males are available to females. No evidence for biases to paternity based on heterozygosity or relatedness was found. These results indicate that female hawksbill turtles are predominantly monogamous within a season, store sperm for the duration of the nesting season and do not re‐mate between nests. Furthermore, females do not appear to be using sperm storage to facilitate sexual selection. Consequently, the primary value of storing sperm in marine turtles may be to uncouple mating and fertilization in time and avoid costly re‐mating.     

Postfledging parental care in Savannah sparrows: sex, size and survival

We investigated postfledging parental care in a philopatric population of Savannah sparrows,Passerculus sandwichensis , breeding on Kent Island, New Brunswick, Canada in an effort to understand the factors influencing adult birds' decisions about parental investment in offspring. Brood division was not based on offspring sex: male and female parents were equally likely to care for sons or daughters. The total duration of parental care, from hatching to independence, was similar for sons and daughters (median=23 days), regardless of the sex of the care-giving parent. The duration of parental care also corresponded closely to the time required for juveniles to acquire basic foraging skills. Despite high levels of extrapair paternity, male Savannah sparrows invested as much in postfledging care and were as effective as females in caring for fledglings, based on recruitment of fledglings into the breeding population the following year. Male parents were more likely to care for smaller fledglings and for offspring from early broods (presumably to enable females to dedicate their efforts towards second clutches). Caring for fledglings was costly for parents: survivorship decreased as a function of the duration of postfledging parental care and the number of fledglings cared for. Parental survivorship, however, was not affected by the sex of the fledglings cared for. This study suggests that sex-biased provisioning may be unlikely except in species with strongly sexually dimorphic offspring, biased offspring sex ratios and sex-biased natal dispersal. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Sexual selection and natural selection in bird speciation

The role of sexual selection in speciation is investigated, addressing two main issues. First, how do sexually selected traits become species recognition traits? Theory and empirical evidence suggest that female preferences often do not evolve as a correlated response to evolution of male traits. This implies that, contrary to runaway (Fisherian) models of sexual selection, premating isolation will not arise as an automatic side effect of divergence between populations in sexually selected traits. I evaluate premating isolating mechanisms in one group, the birds. In this group premating isolation is often a consequence of sexual imprinting, whereby young birds learn features of their parents and use these features in mate choice. Song, morphology and plumage are known recognition cues. I conclude that perhaps the main role for sexual selection in speciation is in generating differences between populations in traits. Sexual imprinting then leads to these traits being used as species recognition mechanisms. The second issue addressed in this paper is the role of sexual selection in adaptive radiation, again concentrating on birds. Ecological differences between species include large differences in size, which may in themselves be sufficient for species recognition, and differences in habitat, which seem to evolve frequently and at all stages of an adaptive radiation. Differences in habitat often cause song and plumage patterns to evolve as a result of sexual selection for efficient communication. Therefore sexual selection is likely to have an important role in generating premating isolating mechanisms throughout an adaptive radiation. It is also possible that sexual selection, by creating more allopatric species, creates more opportunity for ecological divergence to occur. The limited available evidence does not support this idea. A role for sexual selection in accelerating ecological diversification has yet to be demonstrated.     

The role of maternal and paternal effects in the evolution of parental quality by sexual selection

Genetic models of maternal effects and models of mate choice have focused on the evolutionary effects of variation in parental quality. There have been, however, few attempts to combine these into a single model for the evolution of sexually selected traits. We present a quantitative genetic model that considers how male and female parental quality (together or separately) affect the expression of a sexually selected offspring trait. We allow female choice of males based on this parentally affected trait and examine the evolution of mate choice, parental quality and the indicator trait. Our model reveals a number of consequences of maternal and paternal effects. (1) The force of sexual selection owing to adaptive mate choice can displace parental quality from its natural selection optimum. (2) The force of sexual selection can displace female parental quality from its natural selection optimum even when nonadaptive mate choice occurs (e.g. runaway sexual selection), because females of higher parental quality produce more attractive sons and these sons counterbalance the loss in fitness owing to over-investment in each offspring. (3) Maternal and paternal effects can provide a source of genetic variation for offspring traits, allowing evolution by sexual selection even when those traits do not show direct genetic variation (i.e. are not heritable). (4) The correlation between paternal investment and the offspring trait influenced by the parental effects can result in adaptive mate choice and lead to the elaboration of both female preference and the male sexually selected trait. When parental effects exist, sexual selection can drive the evolution of parental quality when investment increases the attractiveness of offspring, leading to the elaboration of indicator traits and higher than expected levels of parental investment.     

Detrimental impacts of radiotransmitters on juvenile Louisiana Waterthrushes

ABSTRACT.   The Louisiana Waterthrush (waterthrush; Seiurus motacilla ) is a forest-dwelling, Nearctic-Neotropical migratory passerine that nests along streams. We attached radiotransmitters (0.6–0.8 g) to 12 nestling waterthrushes using snug, elastic loops. At three nests, adult waterthrushes were videotaped removing radio-tagged young from the nest. In addition, we recovered nine radio-backpacks (with two still attached to the carcasses of nestlings) near nests within a few days after attaching transmitters. Only one of 12 radio-tagged young was relocated more than 24 h after attaching the transmitter. Thus, the method of transmitter attachment we used was not effective. Using snug, nonelastic loops (e.g., nylon) for the harness may reduce the loss of transmitters, but may injure the skin as fledglings grow. Other possible alternatives include (1) gluing the transmitter to skin on the back of nestlings, (2) capturing fledglings in mist nets and attaching transmitters a week or more after fledging by which time contour feathers have grown and the likelihood of a parent removing the transmitter may be reduced, or (3) attempting to monitor fledglings without attaching transmitters. The success of the latter two alternatives would likely be enhanced by attaching transmitters to adults and then tracking them to locate their still-dependent fledglings.     

Within‐season variation in sexual selection in a fish with dynamic sex roles

The strength of sexual selection may vary between species, among populations and within populations over time. While there is growing evidence that sexual selection may vary between years, less is known about variation in sexual selection within a season. Here, we investigate within‐season variation in sexual selection in male two‐spotted gobies (Gobiusculus flavescens). This marine fish experiences a seasonal change in the operational sex ratio from male‐ to female‐biased, resulting in a dramatic decrease in male mating competition over the breeding season. We therefore expected stronger sexual selection on males early in the season. We sampled nests and nest‐holding males early and late in the breeding season and used microsatellite markers to determine male mating and reproductive success. We first analysed sexual selection associated with the acquisition of nests by comparing nest‐holding males to population samples. Among nest‐holders, we calculated the potential strength of sexual selection and selection on phenotypic traits. We found remarkable within‐season variation in sexual selection. Selection on male body size related to nest acquisition changed from positive to negative over the season. The opportunity for sexual selection among nest‐holders was significantly greater early in the season rather than late in the season, partly due to more unmated males. Overall, our study documents a within‐season change in sexual selection that corresponds with a predictable change in the operational sex ratio. We suggest that many species may experience within‐season changes in sexual selection and that such dynamics are important for understanding how sexual selection operates in the wild.     

Brood parasitism increases provisioning rate, and reduces offspring recruitment and adult return rates, in a cowbird host

Interspecific brood parasitism in birds presents a special problem for the host because the parasitic offspring exploit their foster parents, causing them to invest more energy in their current reproductive effort. Nestling brown-headed cowbirds (Molothrus ater) are a burden to relatively small hosts and may reduce fledgling quality and adult survival. We documented food-provisioning rates of one small host, the prothonotary warbler (Protonotaria citrea), at broods that were similar in age (containing nestlings 8–9 days old), but that varied in composition (number of warbler and cowbird nestlings) and mass, and measured the effect of brood parasitism on offspring recruitment and adult returns in the host. The rate of food provisioning increased with brood mass, and males and females contributed equally to feeding nestlings. Controlling for brood mass, the provisioning rate was higher for nests with cowbirds than those without. Recruitment of warbler fledglings from unparasitized nests was 1.6 and 3.7 times higher than that of fledglings from nests containing one or two cowbirds, respectively. Returns of double-brooded adult male and female warblers decreased with an increase in the number of cowbirds raised, but the decrease was more pronounced in males. Reduced returns of warbler adults and recruitment of warbler fledglings with increased cowbird parasitism was likely a result of reduced survival. Cowbird parasitism increased the warblers’ investment in current reproductive effort, while exerting additional costs to current reproduction and residual reproductive value. Our study provides the strongest evidence to date for negative effects of cowbird parasitism on recruitment of host fledglings and survival of host adults.     

Modification of adverse conditions by insects

Many insects modify their environments directly, rather than merely choosing available sites that are already favourable. The modifications are carried out by making excavations in soil and other substrates, constructing feeding or resting shelters, inducing plant responses such as galls, aggregating, building colonial nests, and through parental actions. Such environmental modifications are briefly reviewed and related to the conditions that they modify. Some of the modifications offset physical factors such as dryness or flooding and cool or freezing temperatures. Others reduce the effects of natural enemies or enhance food resources. These effects have seldom been quantified and much of the evidence is anecdotal, but preliminary generalizations are made from existing information. Although potential roles often overlap, excavations and shelters protect especially against physical factors, while aggregations, colonies and parental actions more often influence the acquisition of resources. How modifications affect the impact of natural enemies differs among different kinds of enemies and is especially difficult to test. In any event, adaptive local modifications of the environment by insects are shown to be widely distributed and important. However, their specific roles have often been assumed rather than tested, or have been overlooked along with the potential interdependence of different effects. Therefore, environmental modifications should be considered explicitly and examined with greater rigour during the study of insect life cycles.     

Sexual Selection in the Three-Spined Stickleback

The purpose of this study was to seek evidence for the influence of sexual selection on both male and female three-spined sticklebacks, a species in which the ♂ alone has parental responsibilities. We studied the intrasexual relationships of ♂♂ and ♀♀ prior to courtship, and the intrasexual and intersexual relationships of both sexes during courtship. Results demonstrated that both ♂♂ and ♀♀ compete intrasexually, and suggested that both sexes are discriminative in mate selection. These results are compatible with Trivers' (1972) model of the role of parental investment in sexual selection.     

Social and Thermal Cues Influence Nest‐site Selection in a Nocturnal Gecko,Oedura lesueurii

In oviparous species lacking parental care, successful reproduction depends on females selecting nest sites that facilitate embryonic development. Such sites may be limited in the environment, which can lead to multiple females using the same nest site simultaneously. However, there are several alternative explanations for communal nesting, including natal homing, predator satiation, and adaptive benefits to offspring. We used laboratory experiments to evaluate three hypotheses about nest‐site selection in velvet geckos (Oedura lesueurii), which often nest communally. We investigated whether the trend to nest communally is influenced by the following: (1) evidence of previous nesting (hatched eggshells); (2) body size; and/or (3) thermal regimes. When given the choice, females laid their eggs in shelters containing hatched eggshells rather than in empty shelters, and this was not influenced by body size. Females selected nest sites that were cooler than their own mean selected body temperatures, suggesting that thermal requirements of their developing embryos could outweigh their own thermoregulatory preferences. Field observations of natal homing and high predation rates on gravid females suggest that imprinting on nest sites and/or predator swamping also play roles in communal nesting. Collectively, our results suggest that female velvet geckos use multiple cues to select appropriate nest sites, and hence that multiple mechanisms result in communal nesting behavior in this species.