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Prevalence of different modes of parental care in birds.

Estimates of the incidence of major classes of parental care by birds are drawn from classical studies that preceded both the publication of a massive secondary literature and the revolution driven by molecular approaches to avian phylogeny. Here, I review this literature in the light of new phylogenetic hypotheses and estimate the prevalence of six distinct modes of care: use of geothermal heat to incubate eggs, brood parasitism, male only care, female only care, biparental care and cooperative breeding. Female only care and cooperative breeding are more common than has previously been recognized, occurring in 8 and 9% of species, respectively. Biparental care by a pair-bonded male and female is the most common pattern of care but at 81% of species, the pattern is less common than once believed. I identify several problems with existing hypotheses for the evolution of parental care and highlight a number of poorly understood contrasts which, once resolved, should help elucidate avian social evolution.

Animals↗

Polygyny and its fitness consequences for primary and secondary female pied flycatchers.

In polygynous species with biparental care, the amount of paternal support often varies considerably. In the pied flycatcher (Ficedula hypoleuca), females mated with monogamous males receive more male assistance during the nestling phase than females mated with bigynous males, as the latter have to share their mates with another female. Bigynous males, however, give more support to their primary broods than to their secondary broods. Using a long-term dataset (31 years), the present study revealed that direct reproductive success, i.e. number of fledglings, was lower in females that mated with bigynous males, especially in secondary broods without male assistance, than in females that mated with monogamous males. Secondary broods with male assistance were more affected than primary broods. Female survival was independent of mating status. In primary broods, a delayed compensation for inferior direct reproductive success was found in terms of the number of grandoffspring, a phenomenon that did not occur in secondary broods. Delayed compensation in primary broods refers to indirect effects, i.e. good genes. According to the sexy son hypothesis, genetically superior (i.e. sexy) males may have sons with a higher number of broods belonging to a polygynous breeding status than do sons from broods with a monogamous father. This was indeed the case for sons descending from primary broods, but not for sons descending from secondary broods.

Animals↗

Cryptic gentes revealed in pallid cuckoos Cuculus pallidus using reflectance spectrophotometry.

Many cuckoo species lay eggs that match those of their hosts, which can significantly reduce rejection of their eggs by the host species. However, egg mimicry is problematic for generalist cuckoos that parasitize several host species with different egg types. Some generalist cuckoos have overcome this problem by evolving several host-specific races (gentes), each with its own, host-specific egg type. It is unknown how generalist cuckoos lacking gentes are able to avoid egg rejection by hosts. Here we use reflectance spectrophotometry (300-700 nm) on museum egg collections to test for host-specific egg types in an Australian generalist cuckoo reported to have a single egg type. We show that the colour of pallid cuckoo (Cuculus pallidus) eggs differed between four host species, and that their eggs closely mimicked the eggs of the host they parasitized. These results reveal that pallid cuckoos have host-specific egg types that have not been detected by human observation, and indicate that gentes could be more common than previously realized.

Adaptation, Physiological↗

Surplus nest boxes and the potential for polygyny affect clutch size and offspring sex ratio in house wrens.

Females of many species can gain benefits from being choosy about their mates and even exhibit context-dependent investment in reproduction in response to the quality of their breeding situation. Here, we show that if a male house wren is provided with surplus nest boxes in his territory, his mate lays a larger clutch with a significantly higher proportion of sons. This response to a territory characteristic directly associated with male competitive ability, and ultimately to male reproductive success, suggests that male competition over access to high-quality territories with surplus nest boxes (i.e. those able to support polygyny) may influence female reproductive investment decisions. The results of this study have interesting implications, particularly considering the important role that studies of cavity nesting birds utilizing nest boxes have played in advancing our understanding of behaviour, ecology and evolution.

Animals↗

Maternal developmental stress reduces reproductive success of female offspring in zebra finches.

Environmental factors play a key role in the expression of phenotypic traits and life-history decisions, specifically when they act during early development. In birds, brood size is a main environmental factor affecting development. Experimental manipulation of brood sizes can result in reduced offspring condition, indicating that developmental deficits in enlarged broods have consequences within the affected generation. Yet, it is unclear whether stress during early development can have fitness consequences projecting into the next generation. To study such trans-generational fitness effects, we bred female zebra finches, Taeniopygia guttata, whose mothers had been raised in different experimental brood sizes. We found that adult females were increasingly smaller with increasing experimental brood size in which their mother had been raised. Furthermore, reproductive success at hatching and fledging covaried negatively with the experimental brood size in which their mothers were raised. These results illustrate that early developmental stress can have long-lasting effects affecting reproductive success of future generations. Such trans-generational effects can be life-history responses adapted to environmental conditions experienced early in life.

Animals↗

A UV signal of offspring condition mediates context-dependent parental favouritism.

As fitness returns during a breeding attempt are context-dependent, parents are predicted to bias their food allocation within a brood from poor towards good condition nestlings when environmental conditions deteriorate. We tested this prediction in the Alpine swift and the European starling, two migratory bird species, by modifying an ultraviolet (UV) visual signal of condition in nestlings and exploring how parents allocate food to their young as the season progresses. We show in both species that: (i) UV light reflected by the body skin of offspring positively correlates with their stature (i.e. body mass and skeletal size) and (ii) parental favouritism towards young with more UV reflective skin gradually increases as the season progresses. Early-breeding parents supplied food preferentially to UV pale (i.e. small stature) nestlings, whereas late-breeding parents favoured UV bright offspring (i.e. large stature). These results emphasize that parents use UV signals of offspring condition to adjust their feeding strategies depending on the ecological context.

Animals↗

Dark nests and egg colour in birds: a possible functional role of ultraviolet reflectance in egg detectability.

Owing to the conspicuousness of ultraviolet (UV) colour in dark environments, natural selection might have selected UV egg coloration because it would enhance egg detectability by parents in murky nests. Here, we tested this hypothesis by using comparative and experimental approaches. First, we studied variation in egg coloration of 98 species of European passerines measured using UV-visible reflectance spectrometry (300-700nm) in relation to nesting habits. Analyses based on raw data and controlling for phylogenetic distances both at the species and the family levels revealed that hole-nester species produced eggs with higher UV reflectance than those nesting in open habitats. The experimental approach consisted of the manipulation of UV reflectance of the experimental eggs introduced outside the nest-cup of the hole-nester spotless starling Sturnus unicolor and the study of the retrieval of these eggs. Ultraviolet-reflecting eggs (controls) were more frequently retrieved to the nest-cup than non-reflecting (-UV) eggs. These results were not due to '-UV' eggs being recognized by starlings as parasitic because when a parasitic egg is detected, starlings removed it from the nest-box. Therefore, these results are consistent with the hypothesis that UV egg colours are designed to provide highly detectable targets for parent birds in dark nest environments.

Animals↗

Information flow, opinion polling and collective intelligence in house-hunting social insects.

The sharing and collective processing of information by certain insect societies is one of the reasons that they warrant the superlative epithet 'super-organisms' (Franks 1989, Am. Sci. 77, 138-145). We describe a detailed experimental and mathematical analysis of information exchange and decision-making in, arguably, the most difficult collective choices that social insects face: namely, house hunting by complete societies. The key issue is how can a complete colony select the single best nest-site among several alternatives? Individual scouts respond to the diverse information they have personally obtained about the quality of a potential nest-site by producing a recruitment signal. The colony then deliberates over (i.e. integrates) different incoming recruitment signals associated with different potential nest-sites to achieve a well-informed collective decision. We compare this process in honeybees and in the ant Leptothorax albipennis. Notwithstanding many differences - for example, honeybee colonies have 100 times more individuals than L. albipennis colonies - there are certain similarities in the fundamental algorithms these societies appear to employ when they are house hunting. Scout honeybees use the full power of the waggle dance to inform their nest-mates about the distance and direction of a potential nest-site (and they indicate the quality of a nest-site indirectly through the vigour of their dance), and yet individual bees perhaps only rarely make direct comparisons of such sites. By contrast, scouts from L. albipennis colonies often compare nest-sites, but they cannot directly inform one another of their estimation of the quality of a potential site. Instead, they discriminate between sites by initiating recruitment sooner to better ones. Nevertheless, both species do make use of forms of opinion polling. For example, scout bees that have formerly danced for a certain site cease such advertising and monitor the dances of others at random. That is, they act without prejudice. They neither favour nor disdain dancers that advocate the site they had formerly advertised or the alternatives. Thus, in general the bees are less well informed than they would be if they systematically monitored dances for alternative sites rather than spending their time reprocessing information they already have. However, as a result of their lack of prejudice, less time overall will be wasted in endless debate among stubborn and potentially biased bees. Among the ants, the opinions of nest-mates are also pooled effectively when scouts use a threshold population of their nest-mates present in a new nest-site as a cue to switch to more rapid recruitment. Furthermore, the ants' reluctance to begin recruiting to poor nest-sites means that more time is available for the discovery and direct comparison of alternatives. Likewise, the retirement of honeybee scouts from dancing for a given site allows more time for other scouts to find potentially better sites. Thus, both the ants and the bees have time-lags built into their decision-making systems that should facilitate a compromise between thorough surveys for good nest-sites and relatively rapid decisions. We have also been able to show that classical mathematical models can illuminate the processes by which colonies are able to achieve decisions that are relatively swift and very well informed.

Animal Communication↗

Nested allosteric interactions in the cytoplasmic chaperonin containing TCP-1.

Initial rates of ATP hydrolysis by the chaperonin containing TCP-1 (CCT) from bovine testis were measured as a function of ATP concentration. Two allosteric transitions are observed: one at relatively low concentrations of ATP (<100 microM) and the second at higher concentrations of ATP. The data suggest that CCT has positive intra-ring cooperativity and negative inter-ring cooperativity in ATP hydrolysis, with respect to ATP, as previously observed in the case of GroEL. It is shown that the relatively weak positive intra-ring cooperativity found in the case of CCT may be due to heterogeneity in its subunit composition. Our results suggest that nested allosteric behavior may be common to chaperone double-ring systems.

Adenosine Triphosphate↗

Bayesian modeling of age-specific survival in bird nesting studies under irregular visits.

In this article, a Bayesian model for age-specific nest survival rates is presented to handle the irregular visit case. Both informative priors and noninformative priors are investigated. The reference prior under this model is derived, and, therefore, the hyperparameter specification problem is solved to some extent. The Bayesian method provides a more accurate estimate of the total survival rate than the standard Mayfield method, if the age-specific hazard rates are not constant. The Bayesian method also lets the biologist look for high- and low-survival rates during the whole nesting period. In practice, it is common for data of several types to be collected in a single study. That is, some nests may be aged, others are not. Some nests are visited regularly; others are visited irregularly. The Bayesian method accommodates any mix of these sampling techniques by assuming that the aging and visiting activities have no effect on the survival rate. The methods are illustrated by an analysis of the Missouri northern bobwhite data set.

Aging↗

Obligate brood parasites as selective agents for evolution of egg appearance in passerine birds.

Many passerine host species have counteracted the parasite egg mimicry in their coevolutionary arms race with the common cuckoo (Cuculus canorus) by evolving increased interclutch and reduced intraclutch variation in egg appearance. Such variations make it easier for hosts to recognize a foreign egg, reduce the possibility of making recognition errors, and reduce the ability of the cuckoo to mimic the eggs of a particular host. Here, we investigate if such clutch characteristics have evolved among North American passerines. We predict that due to the absence of brood parasites with egg mimicry on this continent, these passerines should (1) not show any relationship between rejection rates and intra- or interclutch variation, and (2) intraclutch variation should be lower and interclutch variation higher in European hosts exposed to cuckoo parasitism as compared to North American hosts parasitized by cowbirds. Here we present data that show support for most of these and other predictions, as well as when controlling statistically for effects of common descent. However, the effect of continent on intraclutch variation was less than predicted and we discuss a possible reason for this. All things considered, the results demonstrate that parasitism by a specialist brood parasite with egg mimicry is a powerful selective force regarding the evolution of egg characteristics in passerine birds.

Animals↗

Coevolution of an avian host and its parasitic cuckoo.

We use a quantitative genetic model to examine the coevolution of host and cuckoo egg characters (termed "size" as a proxy for general appearance), host discrimination, and host and cuckoo population dynamics. A host decides whether to discard an egg using a comparison of the sizes of the eggs in her nest, which changes as host and cuckoo eggs evolve. Specifically, we assume that the probability that she discards the largest egg in her nest depends on how much larger it is than the second largest egg. This decision rule (i.e., the acceptable difference in egg sizes) also evolves, changing both the chance of successful rejection of a cuckoo egg in parasitized nests and the chance of mistaken rejection of a host egg in both parasitized and unparasitized nests. We find a stable equilibrium for coexistence of the host and cuckoo where there is cuckoo egg mimicry, evolutionary displacement of the host egg away from the cuckoo egg phenotype, and host discrimination against unusual eggs. Both host discrimination and host egg displacement are fairly weak at the equilibrium. Cuckoo egg mimicry, although imperfect, usually evolves more extensively and quickly than the responses of the host. Our model provides evidence for both the evolutionary equilibrium and evolutionary lag hypotheses of host acceptance of parasitic eggs.

Animals↗

Nutrient transfer from sea to land: the case of gulls and cormorants in the Gulf of Maine.

1. The structure of communities is influenced by the transport of resources across ecosystem boundaries. Seabirds are capable of introducing large amounts of marine-derived nutrients to land, thereby modifying resource availability to terrestrial species. 2. In this study we investigated the hypothesis that variation in nesting densities of great black-backed gulls Larus marinus and double-crested cormorants Phalacrocorax auritus would modify the effect of these species on soil nutrients and plant species composition on offshore islands in the Gulf of Maine, USA. 3. Our results showed a significant positive correlation between nest density and concentrations of ammonia and nitrate in soils, but no significant relationship between nest density and phosphate. Ammonia and phosphate concentrations were good predictors of plant species composition; there were more annual forbs than perennial grasses in the abandoned cormorant colony compared with the gull colonies. Extremely high concentrations of ammonia in the highest density colony (active cormorant) may have been the main factor inhibiting plant germination at this site. All of the plant species in gull and cormorant colonies showed enriched delta(15)N signatures, indicating substantial input of marine-derived nitrogen from seabirds. 4. Our study demonstrated that gulls and cormorants are effective vectors for the transport of marine nutrients to terrestrial ecosystems. However, transported nutrients occurred in particularly high concentrations in areas with nesting cormorants. Nesting densities and species-specific variation in resource transport should be considered when predicting the effects of seabirds and other biogenic vectors of allochthonous resources.

Ammonia↗

Density effects on life-history traits in a wild population of the great tit Parus major: analyses of long-term data with GIS techniques.

1. Population density often has strong effects on the population dynamics and reproductive processes of territorial animals. However, most estimates of density-dependent effects use the number of breeding pairs per unit area in a given season and look for correlations across seasons, a technique that assigns the same density score to each breeding pair, irrespective of local spatial variation. 2. In this study, we employed GIS techniques to estimate individual breeding densities for great tits breeding in Wytham Woods UK, between 1965 and 1996. We then used linear mixed modelling to analyse the effect of density on reproductive processes. 3. The areas of Thiessen polygons formed around occupied nestboxes were used to approximate territory size (necessarily inverse of breeding density). There were significant, independent and positive relationships between clutch size, fledging mass and the number of offspring recruited to the population, and territory size (all P < 0.001), but no effect of territory size on lay-date or egg mass. 4. Thiessen polygons are contiguous and cover all of the available area. Therefore, at low nest densities territory polygons were excessively oversized. Using a novel procedure to address this limitation, territory sizes were systematically capped through a range of maxima, with the greatest effect in the models when territories were capped at 0.9-2.3 ha. This figure approximates to the maximum effective territory size in our population and is in close agreement with several field-based studies. This capping refinement also revealed a significant negative relationship between lay-date and territory size capped at 0.9 ha (P < 0.001). 5. These density-dependent effects were also detected when analyses were restricted to changes within individual females, suggesting that density effects do not merely result from either increased proportions of low-quality individuals, or increased occupation of poor sites, when population density is high. 6. Overall, these results suggest that, in the current population, great tits with territories smaller than c. 2 ha independently lay smaller and later clutches, have lighter fledglings, and recruit fewer offspring to the breeding population. These analyses thus suggest a pervasive and causal role of local population density in explaining individual reproductive processes.

Animals↗

Condition-dependent nesting in bluegill sunfish Lepomis macrochirus.

1. There are two prominent, nonmutually exclusive hypotheses to explain the timing of reproduction in animals: energetic constraint and adaptive behaviour. 2. We tested these hypotheses by quantifying the costs and benefits of nesting at different times in the season for male bluegill sunfish Lepomis macrochirus, a species with paternal care, in Lake Opinicon (Ontario, Canada). 3. The value of nesting at different times during the breeding season (RSb) was determined from spawning individuals as RSb = Pb x Cb x Sb x Ob, where Pb is the probability of spawning during each bout b, Cb is the expected brood size, Sb is the expected brood survivorship to 'swim-up', and Ob is the survivorship of free-swimming fry to age 1 year. 4. The results show that the value of nesting peaks during the middle of the season. However, nesting patterns varied with male condition and not all males nested at the peak. 5. Larger males, which were able to nest multiple times, first nested early in the season when overwinter survivorship of offspring and renesting opportunities later in the season were maximized. These males had the highest seasonal reproductive success. 6. Smaller males, which nested a single time, delayed nesting until the middle of the season when spawning opportunities and brood sizes were greatest. 7. These data suggest that both energetics and adaptive behaviour play roles in determining the timing of reproduction.

Adaptation, Physiological↗

Nest ornamentation by female spotless starlings in response to a male display: an experimental study.

1. The use of behavioural traits by females in signalling condition has been practically ignored in evolutionary theory. However, females may also exhibit ornaments and behavioural displays, although less elaborated than those of males. 2. In this study we suggest that the carrying of feathers by spotless starlings Sturnus unicolor Temminck females to decorate the nest represents an elaborated and costly behaviour that is displayed in response to a courtship male behaviour: the carrying of nest green plants. 3. By experimentally increasing the amount of green plants in the nests, to give the appearance that highly attractive males defended them, we induced females to increase their feather carrying rates. 4. The amount of feathers carried to the nest was correlated to female reproductive experience and laying date, two variables correlated with female body condition. These results suggests that this behaviour may work as an honest indicator of female quality. 5. We conclude that male carrying plants and female carrying feathers can be viewed as two sex-specific functionally related signalling behaviours involved in mutual courtship or status signalling.

Animals↗

Effects of parental effort on blood stress protein HSP60 and immunoglobulins in female blue tits: a brood size manipulation experiment.

1. Physiological stress in animals may impose a limit for investment in current reproduction in the wild. A brood manipulation experiment was conducted in a population of blue tits Cyanistes caeruleus to study the effect of parental effort on changes in two types of proteins related with stress: the blood stress protein HSP60 and the plasma immunoglobulins. 2. Levels of HSP60 were reduced across the experiment for females attending reduced broods, and females attending enlarged broods experienced a reduction of immunoglobulin levels. Moreover, the overall changes in the levels of both proteins were positively related. 3. By controlling for the change in immunoglobulin levels we found an increase in HSP60 for females in the enlarged treatment, presumably to offset deleterious effects derived from increased effort. 4. Maternal effort was able to partially compensate for the effect of treatment as nestlings did not differ in mass and levels of immunoglobulins and HSP60 among treatments. 5. Physiological stress as reflected in stress and immunoglobulin proteins may limit maternal effort in breeding blue tits.

Animals↗

Spring temperature, clutch initiation date and duck nest success: a test of the mismatch hypothesis.

1. Increases in average global temperature during the twentieth century have prompted calls for research on the effect of temperature variation on avian population dynamics. Particular attention has been paid to the hypothesis that increased temperatures may affect a species' ability to shift their breeding efforts to match the phenology of their prey, and thus result in reduced reproductive success (the 'mismatch hypothesis'). 2. We used data from a long-term study of breeding ducks to investigate how duck nest success varied with clutch initiation date, and to test whether spring temperature affected this relationship in a manner consistent with the mismatch hypothesis. We modelled five possible functional forms of how nest success might vary with clutch initiation date and spring temperature, and used an information-theoretic approach to determine which model best described the nesting outcomes of five dabbling duck species nesting in Saskatchewan, Canada. 3. Probability of nest success for the five species did not vary strongly with clutch initiation date, and we found evidence consistent with the mismatch hypothesis for one species, northern pintail Anas acuta, although weight of evidence was weak. 4. Overall nest success of all five species was positively associated with spring temperature. These results suggest that increasing spring temperature alone (within the range observed in this study) may not affect nest success in a manner that would result in lower populations of breeding ducks.

Animals↗