The role of posturing and calling in the begging display of nestling birds

Nestling birds produce a multicomponent begging display that has visual (e.g. posturing) and vocal (e.g. call rate) elements. Most work on the function of the display has focused on each component separately. However, understanding the evolution of complex displays such as begging requires knowledge of how the components function collectively. The purpose of our study was to determine how postural intensity and calling rate together influence parental feeding decisions in tree swallows, Tachycineta bicolor. We compared how begging components responded to a manipulation in which pairs of nestlings were either free to approach the parent when it arrived to feed (unconfined treatment) or confined to the back of their nestbox by a Plexiglass partition (confined treatment). We found no significant differences in postural intensity between treatments, but calling rate was significantly higher in the confined treatment. In both treatments, postural intensity, but not calling rate, correlated with hunger. Both components positively and independently correlated with the likelihood of a nestling being fed, although the correlation with postural intensity was stronger. Previous work suggested that both posture and call rate advertised hunger in nestling tree swallows. Here, call rate was not associated with hunger, but rather was affected by nestling position. These results suggest that calling may serve an additional role in helping nestlings in disadvantaged positions attract parental attention. The results also suggest that calling may have a complex relationship with hunger, position and nestmates.     

Sex-specific effects of yolk-androgens on growth of nestling American kestrels

Maternally derived androgen hormones concentrate in avian egg yolks as the yolks grow on the female’s ovary, possibly forming a basis for important maternal effects in birds. In the American kestrel (Falco sparverius), experimental elevation of yolk androgens in the first-laid egg of a clutch (a-egg) to the concentrations found naturally in a clutch’s later-laid eggs reduces the growth rate of a-egg nestlings compared to controls. These findings, together with discoveries from other species that the effects of yolk androgens on growth of female nestlings may differ from their effects on growth of male nestlings, raise the hypothesis that natural changes in yolk-androgen concentrations with laying order are ultimately due to a difference between the sexes in their yolk-androgen sensitivity and between early- and late-laid eggs in their sex ratio. By re-analyzing previously published data and adding to the analysis data from previously unanalyzed blood samples used for sex determination, we investigated possible sex-specific effects of yolk-androgens in the context of a potential sex-biased laying order in free-living American kestrels. We used a multi-level, mixed model with a Gompertz function to analyze growth of nestlings hatching from a-eggs that were control-treated or in which we experimentally elevated yolk-androgen concentrations shortly after laying to the higher concentrations naturally found in later-laid eggs. We discovered that male nestlings were more susceptible than female nestlings to growth inhibition by yolk-androgen elevation but did not find a bias in sex ratio with respect to laying order. Together, these findings do not support the above hypothesis. However, they are consistent with the hypothesis that sex differences in yolk-androgen sensitivity enable mothers to economically tune reproductive effort to an individual offspring’s reproductive value, which can vary more for one sex than the other.     

Paternal care in the European starling,Sturnus vulgaris: nestling provisioning

The extent to which male birds in polygynous species with biparental care assist in nestling feeding often varies considerably between nests of different mating status. Both how much polygynous males assist and how they divide their effort between nests may have a profound effect on the evolution of mating systems. In this study we investigated how males in the facultatively polygynous European starling Sturnus vulgaris invested in their different nests. The amount of male assistance affected the quality of the offspring. Polygynous males invested as much as monogamous males, but divided their effort asymmetrically between nests, predominantly feeding nestlings of first-mated (primary) females. Although females partly compensated for loss of male assistance, total feeding frequency was lower at primary females’ nests than at monogamous females nests. Secondary females received even less assistance with nestling rearing, and the extent to which males assisted decreased with the length of the interval between the hatching of the primary and secondary clutches. These results are contrasted with those from a Belgian populations of starlings with a much more protracted breeding season and thus greater opportunities for males to attract additional mates during the nestling rearing period. The results show that both the “defence of male parental investment model” and the “asynchronous settlement model” have explanatory power, but that their validity depends on the potential length of the breeding season. Received: 21 July 1995/Accepted after revision: 13 July 1996     

Context-dependent effects of nestling growth trajectories on recruitment probability in the collared flycatcher

Nestling growth is known as an important determinant of fitness in altricial birds, but its evolutionary potential has been debated, and little is known about detailed patterns of current selection on growth. Relationships are often reported between nestling growth and attributes of nestlings and parents, but the interpretation of these depends on the advantages a given growth difference confers to the chicks. Increased growth may have positive, negative or context-dependent effects on offspring fitness, but these effects are largely unknown in natural populations. We measured growth trajectories of body mass in fostered broods of collared flycatchers (Ficedula albicollis) in 3 years of contrasting food conditions. We examined the growth of young and their recruitment probability to the breeding population in relation to year quality, hatching rank, sex, paternal age and paternal attractiveness. We also looked at the interactive effects of growth and intrinsic offspring attributes on recruitment probability. The predictors of nestling growth and those of recruitment did not agree. Moreover, the recruitment consequences of a given nestling growth rate were significantly influenced by nestling rank and paternal ornamentation. Differential recruitment effects of nestling growth in relation to parental traits and nestling attributes suggest that using growth as a generally applicable measure of nestling quality may not be justified. These findings also have implications for morphological evolution and the indicator value of sexual signals.     

The importance of nestling location for obtaining food in open cup-nests

Observations and experiments were carried out at 34 four-nestling nests of Arabian babblers (Turdoides squamiceps) and feeding events were analyzed according to hatching order and nestling location in the nest. Nestlings were fed in a negative correlation with hatching order. Nestling locations in relation to the provisioners were defined as Near, Far, Right, Left, and Center. Feeders fed closer nestlings more often than those further away, and straight ahead rather than sideways. In the open circular nest, the feeding adults' positions were unpredictable and equally distributed around the nest. Each peripheral position therefore served equally as Near, Far, Right and Left positions. The Center was independent of the arrival direction of the feeders and was advantageous relative to the peripheral positions. The importance of the central position was demonstrated by three experiments. (a) Food deprivation for 1 h caused the first-hatched nestling to occupy the center position and gain more feedings than its siblings. (b) Introduction of a Perspex barrier into the nest, thus eliminating the central position and preventing the nestlings from changing places, led to a loss of importance of hatching order and equal provisioning of the nestlings. (c) Fencing the nest so that entry was possible from only one permanent direction, resulted in the first-hatched nestling occupying the near position more often than its nestmates and obtaining 52% of the feedings provisioned to the whole brood. These findings imply that the architecture of the nest has an important role in food distribution among the nestlings, and contributes to reducing inequalities in the siblings' abilities to obtain food.     

Quantifying the global threat to native birds from predation by non-native birds on small islands

Although invasive non-native species can adversely affect biodiversity in many ways, predation of native species by non-native species on islands can be severely damaging. Results of numerous studies document non-native birds preying on birds on islands, but our understanding of the number and type of species affected has been limited by the lack of a global review of these impacts. I identified the non-native bird species that have been recorded preying on birds, the locations where this predation occurred, and the bird species affected. Because the impacts of non-native birds can be particularly severe on small islands, I then identified the islands <500 km² around the world that are occupied by predatory non-native birds. By taking into account their life-history traits and predation history, I also identified the near-threatened and threatened bird species on these islands that they may prey on. The results indicated that predation by non-native birds was primarily a concern for threatened bird conservation on small islands; almost all predation impacts (91%) on near-threatened and threatened birds were recorded on islands, and median island size was 106 km². I also found non-native bird predation was a poorly known and widespread potential threat to avian biodiversity; worldwide, 194 islands of <500 km² were occupied by predatory non-native birds, but information on their impacts was unavailable for most of these islands. On them, where the impacts of non-native species can be severe, non-native birds may be preying on approximately 6% of the world's near-threatened and threatened bird species. Four non-native bird species I identified have been successfully eradicated from islands. If they were eradicated from the small islands they occupy, 70% of the near-threatened and threatened bird species I identified would no longer be affected by nest predation by non-native birds on small islands.     

Evaluating the coastal environment for marine birds

The marine environment is being increasingly exploited by fisheries and the oil and gas industry. Conservationists urgently need the ability to identify the processes that determine patterns of abundance of marine species. We describe a preliminary Geographic Information System (GIS) in which spatial data on environmental variables (seabird colony locations, sea depth and seabed sediments) are integrated with realistic energy constraints faced by marine birds during the breeding season. A simple foraging model predicts the spatial variation in the quality of given locations as potential feeding sites under different feeding conditions and stages of the breeding cycle. We show how the approach can be used to help managers identify key marine areas and assess the impacts of environmental change or damage.     

Measuring the impact of the pet trade on Indonesian birds

The trade in wild animals involves one‐third of the world's bird species and thousands of other vertebrate species. Although a few species are imperiled as a result of the wildlife trade, the lack of field studies makes it difficult to gauge how serious a threat it is to biodiversity. We used data on changes in bird abundances across space and time and information from trapper interviews to evaluate the effects of trapping wild birds for the pet trade in Sumatra, Indonesia. To analyze changes in bird abundance over time, we used data gathered over 14 years of repeated bird surveys in a 900‐ha forest in southern Sumatra. In northern Sumatra, we surveyed birds along a gradient of trapping accessibility, from the edge of roads to 5 km into the forest interior. We interviewed 49 bird trappers in northern Sumatra to learn which species they targeted and how far they went into the forest to trap. We used prices from Sumatran bird markets as a proxy for demand and, therefore, trapping pressure. Market price was a significant predictor of species declines over time in southern Sumatra (e.g., given a market price increase of approximately $50, the log change in abundance per year decreased by 0.06 on average). This result indicates a link between the market‐based pet trade and community‐wide species declines. In northern Sumatra, price and change in abundance were not related to remoteness (distance from the nearest road). However, based on our field surveys, high‐value species were rare or absent across this region. The median maximum distance trappers went into the forest each day was 5.0 km. This suggests that trapping has depleted bird populations across our remoteness gradient. We found that less than half of Sumatra's remaining forests are >5 km from a major road. Our results suggest that trapping for the pet trade threatens birds in Sumatra. Given the popularity of pet birds across Southeast Asia, additional studies are urgently needed to determine the extent and magnitude of the threat posed by the pet trade.     

Sibling aggression, hatching asynchrony, and nestling mortality in the black kite (Milvus migrans)

In siblicidal species, hatching asynchrony could act to reduce sibling rivalry or promote the death of last-hatched chicks. The pattern of hatching asynchrony was experimentally altered in the black kite Milvus migrans. Hatching asynchrony in control broods was intermediate between those of experimentally synchronised and asynchronised broods. Sibling aggression and wounds on the chicks were more commonly observed early in the nestling period and in synchronous nests. Serious injuries were observed on last-hatched chicks in asynchronous nests, as were observations of intimidated or crushed chicks. Sibling aggression was related to food abundance, but some chicks died at an early age in nests with abundant food (cainism). Cainism was more commonly found in asynchronous nests. For species with facultative siblicide, moderate hatching asynchrony could be a compromise between reducing sibling rivalry and avoiding large size differences between sibs that would result in cainism. Female black kites preferentially fed the smallest chicks and exhibited behaviours to reduce sibling aggression, contrary to observations in other siblicidal species. In a highly opportunistic forager such as the black kite, a strategy may exist to protract the life of all the chicks in the brood, waiting for unpredictable situations of food overabundance. This would induce the appearance of a parent-offspring conflict over brood reduction, reflected in the existence of a possible anticipated response by some of the chicks (cainism) and in the appearance of special behaviours by the parents to selectively feed smaller chicks or reduce sibling aggression. In this facultatively siblicidal species, cainism does not seem to be the final stage of an evolutionary trend favouring the raising of high-quality chicks, but a manifestation of a parent-offspring conflict over brood size. Received: 9 March 1998 / Accepted after revision: 8 August 1998     

Vitamin E supplementation enhances growth and condition of nestling barn swallows (Hirundo rustica)

To optimize fitness, organisms may have to trade the number and quality of individual offspring against their own condition and survival. Limiting micronutrients such as antioxidants may be crucial to this trade-off. We investigated whether vitamin E, a major antioxidant in the diet of vertebrates, is limiting to barn swallow (Hirundo rustica) nestlings. We manipulated brood size to alter the intensity of sib–sib competition and supplemented nestlings with two different physiological doses of vitamin E while establishing a control group. Treatment effects were measured on body mass and size, feather growth, T cell-mediated immune response and hematocrit. Supplementation with vitamin E at intermediate physiological doses improved nestling mass and condition and feather growth, whereas higher physiological doses did not enhance offspring quality compared to a control treatment. The positive effects of vitamin E on body mass and condition were only detectable from days 6 to 10 when maximum growth rate is attained. Experimental enlargement of broods reduced body mass and size and T cell-mediated immune response only during the late nestling period. The effect of vitamin E supplementation did not depend on brood size manipulation, as revealed by the nonsignificant statistical interaction. This result contradicts the hypothesis that availability of vitamin E depends on intrabrood competition and instead suggests that it depends on concentration of vitamin E in the insect prey of swallows. Thus, antioxidants may be available in limited amounts to barn swallow nestlings and such limitation affects growth. In addition, present results confirm that barn swallow parents trade progeny number against growth and immunity of individual offspring.     

Escape take-off strategies in birds: the significance of protective cover

Avian escape strategy is highly dependent on the ability to fly, and the success rate of raptor attacks is reduced if the prey gets fully airborne. Therefore, when escaping from predator attacks, the initial take-off is crucial and a rapid take-off, high velocity, and high angle of ascent maximize the chance of survival. However, due to the laws of gravity, birds face a trade-off between maximizing its linear acceleration and maximizing its rate of climb when taking off. The optimal policy between velocity and angle of ascent when a bird escapes from an attacking predator might depend on the detailed nature of the predator’s attack, the proximity to cover, and the presence of conspecifics. Many small birds rely on a quick dash to protective cover. In this study, we examine how the availability of protective cover affects take-off strategy in birds. Male great tits (Parus major) were subjected to a surprise attack by a model predator either in the presence or absence of protective cover. When attacked by the predator, great tits took off and perched in the cover when it was available. Birds subjected to the predator attack in the absence of cover took off in 10° higher angle of ascent and flew faster in the start of the take-off, compared to birds that took off in the presence of cover. Thus, this study for the first time shows that a bird do trade off speed with angle of ascent in an adaptive way, depending on the presence of protective cover.     

Wind selectivity of migratory flight departures in birds

Optimal migration theory predicts that birds minimizing the overall time of migration should adjust stopover duration with respect to the rate of fuel accumulation. Recent theoretical developments also take into account the wind situation and predict that there is a time window (a set of days) during which birds should depart when assisted by winds but will not do so if there are head winds. There is also a final day when birds will depart irrespective of wind conditions. Hence, the wind model of optimal migration theory predicts that birds should be sensitive to winds and that there should be a correlation between departures and winds blowing towards the intended migration direction. We tested this assumption by tracking the departures of radio-tagged passerines during autumn migration in southern Sweden. Our birds were moderately to very fat when released and therefore energetically ready for departure. There was a significant correlation between direction of departure and wind direction. We also found that during days when birds departed there was a significantly larger tail wind component than during days when birds were present but did not depart. Our results show that passerines do take the current wind situation into account when departing on migratory flights. We also briefly discuss possible clues that birds use when estimating wind direction and strength. The inclusion of wind is an important amendment to optimal migration theory of birds and should be explored further. Received: 1 March 1999 / Received in revised form: 4 October 1999 / Accepted: 16 October 1999     

Detecting pigments from colourful eggshells of extinct birds

The known chemical basis of diverse avian eggshell coloration is generated by the same two classes of tetrapyrrole pigments in most living birds. We aimed to extend the evolutionary scope of these patterns by detecting pigments from extinct birds’ eggs. In our samples biliverdin was successfully extracted from subfossil shell fragments of the blue-green egg-laying upland moa Megalapteryx didinus, while protoporphyrin was extracted from the beige eggs of two other extinct moa species. Our data on pigment detection from eggshells of other extant paleognath birds, together with published information on other modern lineages, confirm tetrapyrroles as ubiquitous and conserved pigments contributing to diverse eggshell colours throughout avian evolution.     

Generation lengths of the world's birds and their implications for extinction risk

Birds have been comprehensively assessed on the International Union for Conservation of Nature (IUCN) Red List more times than any other taxonomic group. However, to date, generation lengths have not been systematically estimated to scale population trends when undertaking assessments, as required by the criteria of the IUCN Red List. We compiled information from major databases of published life-history and trait data for all birds and imputed missing life-history data as a function of species traits with generalized linear mixed models. Generation lengths were derived for all species, based on our modeled values of age at first breeding, maximum longevity, and annual adult survival. The resulting generation lengths varied from 1.42 to 27.87 years (median 2.99). Most species (61%) had generation lengths <3.33 years, meaning that the period of 3 generations—over which population declines are assessed under criterion A—was <10 years, which is the value used for IUCN Red List assessments of species with short generation times. For these species, our trait-informed estimates of generation length suggested that 10 years is a robust precautionary value for threat assessment. In other cases, however, for whole families, genera, or individual species, generation length had a substantial impact on their estimated extinction risk, resulting in higher extinction risk in long-lived species than in short-lived species. Although our approach effectively addressed data gaps, generation lengths for some species may have been underestimated due to a paucity of life-history data. Overall, our results will strengthen future extinction-risk assessments and augment key databases of avian life-history and trait data.     

Region-wide retreats from lower elevations of range-restricted birds across the Northern Andes

Local studies show upslope shifts in the distribution of tropical birds in response to warming temperatures. Unanswered is whether these upward shifts occur regionally across many species. We considered a nearly 2000-km length of the Northern Andes, where deforestation, temperature, and extreme weather events have increased during the past decades. Range-restricted bird species are particularly vulnerable to such events and occur in exceptionally high numbers in this region. Using abundant crowd-sourced data from the Cornell Lab of Ornithology database, eBird, and the Global Biodiversity Information Facility, we documented distributions of nearly 200 such species. We examined whether species shifted their elevational ranges over time by comparing observed versus expected occurrences below a low elevational threshold and above a high elevational threshold for 2 periods: before and after 2005. We predicted fewer observations at lower elevations (those below the threshold) and more at upper elevations (those above the threshold) after 2005. We also tested for deforestation effects at lower elevations within each species’ distribution ranges. We compared relative forest loss with the differences between observed and expected occurrences across the elevational range. Species’ retreats from lower elevations were ubiquitous and involved a 23–40% decline in prevalence at the lowest elevations. Increases at higher elevations were not consistent. The retreats occurred across a broad spectrum of species, from predominantly lowland to predominantly highland. Because deforestation showed no relationship with species retreats, we contend that a warming climate is the most parsimonious explanation for such shifts.     

The air speed of migrating birds and its relationship to the wind

Summary This paper deals with the flight speeds of migrating birds observed by radar over a Swiss Alpine pass. The distributions of air speeds for different classes of birds and the influence of the wind on air speeds were investigated. Our findings differ greatly from observations of bird migration over the North Atlantic Ocean and the North American continent. Our data reveal: (1) that the air speeds of the bulk of migrating birds were in the range of 8–18 m/s and that the amount of slow flyers (air speed below 5 m/s) was less than 5%; (2) that there was an obvious influence of the wind on air speeds and, moreover, the data showed a distinctly variable compensatory behavior among different bird classes; (3) that the wind component in the direction of the bird's heading was a better predictor for air speed than the wind direction. Although we do not speculate about possibilities and mechanisms of wind detection, a simple argument shows how birds could estimate wind directions if they do use the moving surface as a reference system.