Age-Related Changes in Signalling of Need by Nestling Tree Swallows (Tachycineta bicolor)

Despite a large literature on the ontogeny of behaviour, few studies have examined how the function of juvenile behaviour changes during development. One of the most widespread and important juvenile behaviours is begging, the display used by young animals to solicit food from their parents. Begging signals generally vary reliably with offspring need for food and have served as models for understanding the evolution of honest signalling. Little is known, however, about whether the relationship between begging and need varies over the period of rapid juvenile development. Here, we examine whether tree swallow, Tachycineta bicolor, begging calls consistently reflect hunger levels across the 20 d nestling period. We recorded begging calls at 5, 10 and 15 d post‐hatch, during an hour of food deprivation, and related call features to time without food (i.e. hunger) at each age. The overall correlation between call structure and hunger, as measured by canonical correlation, was consistent across ages. The particular features that correlated with hunger varied, however. Call rate and length increased with hunger at all ages, but call amplitude and frequency range increased with hunger at days 10 and 15 only. The results of our study suggest that begging calls consistently convey information about offspring hunger throughout the nestling period, with the number of call features encoding hunger increasing with nestling age. This change may enhance the ability of parents to assess offspring hunger levels by adding redundancy to the signal.     

Food abundance and clutch size of Tree Swallows Tachycineta bicolor

We studied Tree Swallows Tachycineta bicolor during eight breeding seasons (1977-84) at three sites that differed in food availability, which we measured as an index of the biomass of flying insects derived from daily catches in suspended nets. Mean clutch size differed significantly between sites (6.2, 5.8, and 5.6 eggs) and was related to average food abundance at the site. Within two sites, clutch sizes were significantly larger in years when food was more abundant during egg-laying ( r = 0.83 and 0.97 at the high and low food abundance sites, respectively; r = 0.63, not significant, at the medium abundance site). Both clutch size and laying season food abundance were positively correlated with food abundance during the nestling stage at the 'high' site, but not at the other two sites. Our results support the hypothesis that female Tree Swallows respond proximately to abundant food during egg-laying by laying more eggs, but they do not elucidate the ultimate evolutionary factors involved in determining clutch size.     

The environmental and genetic determinants of chick telomere length in Tree Swallows (Tachycineta bicolor)

Conditions during early life can have dramatic effects on adult characteristics and fitness. However, we still know little about the mechanisms that mediate these relationships. Telomere shortening is one possibility. Telomeres are long sequences of DNA that protect the ends of chromosomes. They shorten naturally throughout an individual's life, and individuals with short telomeres tend to have poorer health and reduced survival. Given this connection between telomere length (TL) and fitness, natural selection should favor individuals that are able to retain longer telomeres for a greater portion of their lives. However, the ability of natural selection to act on TL depends on the extent to which genetic and environmental factors influence TL. In this study, we experimentally enlarged broods of Tree Swallows (Tachycineta bicolor) to test the effects of demanding early‐life conditions on TL, while simultaneously cross‐fostering chicks to estimate heritable genetic influences on TL. In addition, we estimated the effects of parental age and chick sex on chick TL. We found that TL is highly heritable in Tree Swallow chicks, and that the maternal genetic basis for TL is stronger than is the paternal genetic basis. In contrast, the experimental manipulation of brood size had only a weak effect on chick TL, suggesting that the role of environmental factors in influencing TL early in life is limited. There was no effect of chick sex or parental age on chick TL. While these results are consistent with those reported in some studies, they are in conflict with others. These disparate conclusions might be attributable to the inherent complexity of telomere dynamics playing out differently in different populations or to study‐specific variation in the age at which subjects were measured.     

Cavity size, clutch-size and the breeding ecology of Tree Swallows Tachycineta bicolor

Previous studies provide evidence that cavity size influences clutch-size and reproductive success in some hole-nesting birds, because overcrowding in cavities may cause brood mortality due to trampling or hyperthermia. We tested this hypothesis with two experiments at nestbox populations of Tree Swallows Tachycineta bicolor in southeastern Ontario. Female Tree Swallows showed a preference for nesting in large boxes over small ones in the first experiment, and they laid significantly smaller clutches in small nestboxes during both experiments. Differences in clutch–size between large and small nestboxes could not be attributed to other factors known to influence clutch-size in birds (e.g. parental quality, habitat quality, laying date). Reproductive success, however, did not differ between pairs using the two box types during either experiment, nor did it differ during within-clutch-size comparisons between box types. Some measures of nestling quality were significantly lower for broods in large boxes, but during most comparisons there were no differences. These results do not support the adaptive reason for why cavity size affects clutch-size. We suggest that broods in our experiments did not experience the microhabitat conditions necessary to induce the expected differences in brood mortality.     

Laying order, hatching asynchrony and nestling body mass in Tree Swallows Tachycineta bicolor

We studied the reproductive biology of a box-nesting population of Tree Swallows Tachycineta bicolor in southeastern Wisconsin, USA. We were interested particularly in the relationship between laying order and hatching order and the extent to which each was a predictor of nestling body mass. We found that laying order was a significant predictor of hatching order. Laying and hatching order were related to nestling mass at 4 days of age and to a lesser extent at 12 days of age. In addition, we investigated the effects of natural variation in hatching asynchrony. Hatching asynchrony was positively related to the range of nestling body masses within a brood at days 4 and 12. The probability that brood reduction occurred was also positively related to the degree of hatching asynchrony, though this effect was significant only at day 4. Our results suggest that laying order and hatching order have their greatest effects on nestling Tree Swallows early in the nestling period.     

Relative importance off environmental variables in determining the growth off nestling Tree Swallows Tachycineta bicolor

Unpredictable, short-term changes in environmental conditions associated with weather patterns have severe impacts on the reproductive success of Tree Swallows Tachycineta bicolor , as well as other aerial insectivores. We examined the effects of day-to-day environmental variation on the growth of nestling Tree Swallows. Using path analysis, we compared the relative importance of changes in food availability, ambient temperature and wind speed on changes in mass and feather length over four years at our study site near Ithaca, New York. Changes in environmental conditions explained a significant amount of the variation in nestling growth, with daily high temperature having the greatest influence for nestlings younger than five days. Both food supply and temperature had significant direct effects on growth of nestlings older than nine days, with temperature also having a significant indirect effect through its influence on food availability. For both young and old chicks, mass gains were smaller for chicks hatched later in the season. Environmental conditions also explained a significant amount of the linear growth of the ninth primary feather in one of two years, with food supply showing the most consistent positive effect. Environmental conditions several days prior to the measurements significantly influenced feather growth, indicating that the effects of adverse conditions are spread out over several days of feather growth, but this effect was not seen for change in mass.     

Nestling tree swallows (Tachycineta bicolor) alter begging behaviour in response to odour of familiar adults,but not their nests

Communication using chemical cues is important for many taxa, including birds, but the use of olfaction for intraspecific communication has been investigated only recently in passerines and is understudied in nestlings. To address this knowledge gap, we explored whether nestling tree swallows (Tachycineta bicolor) would recognize and respond to chemical cues of conspecifics, specifically testing begging responses to familiar and unfamiliar nest and adult odours. For the nest odour experiment, nests were treated with either orange essential oil or distilled water to create scented and unscented (control) odour environments, respectively. For the adult odour experiment, adults attending the nestlings were considered “familiar adults” and adults attending a different brood in the population were considered “unfamiliar adults.” We found that begging responses of nestlings did not differ in response to orange oil odour or water, but nestlings begged significantly longer and more intensely in response to odours of a familiar than an unfamiliar adult, regardless of adult sex. This provides evidence that tree swallows use chemical cues to alter their behaviour and opens up many exciting avenues of future research.     

Differences in size between first and replacement clutches match the seasonal decline in single clutches in Tree Swallows Tachycineta bicolor

The seasonal decline in clutch size in birds can be a response to the environmentally conditioned decrease in prospects for offspring or a consequence of a lower physical ability of late‐breeding females. To find out which of the explanations apply in Tree Swallows Tachycineta bicolor, we assessed whether replacement clutch size in this species is affected by an individual female's ability to lay a certain number of eggs. To do this, we measured the decline in clutch size as a function of laying date between first and replacement clutches in individuals that re‐nested following natural failure, and compared this with the rate of decline in clutch size with laying date for Tree Swallows that laid only a single clutch in that season. Additionally, we assessed whether the clutch size and the rate of its seasonal decline varied across years. We accounted for the truncated and under‐dispersed nature of clutch size data by using a Bayesian approach in the analysis. We found little variation in the rate of clutch size decline across years at our breeding site. Accounting for this seasonal decline in clutch size, mean clutch size was similar between single‐time breeding females and those that laid replacement clutches, implying that the number of eggs laid on the second attempt by female Tree Swallows is determined by laying date, rather than by the female's physical ability to produce a clutch of a certain size.     

Tree swallows (Tachycineta bicolor) nesting on wetlands impacted by oil sands mining are highly parasitized by the bird blow fly Protocalliphora spp

Oil sands mining is steadily expanding in Alberta, Canada. Major companies are planning reclamation strategies for mine tailings, in which wetlands will be used for the bioremediation of water and sediments contaminated with polycyclic aromatic hydrocarbons and naphthenic acids during the extraction process. A series of experimental wetlands were built on companies' leases to assess the feasibility of this approach, and tree swallows (Tachycineta bicolor) were designated as upper trophic biological sentinels. From May to July 2004, prevalence and intensity of infestation with bird blow flies Protocalliphora spp. (Diptera: Calliphoridae) were measured in nests on oil sands reclaimed wetlands and compared with those on a reference site. Nestling growth and survival also were monitored. Prevalence of infestation was surprisingly high for a small cavity nester; 100% of the 38 nests examined were infested. Nests on wetlands containing oil sands waste materials harbored on average from 60% to 72% more blow fly larvae than those on the reference site. Nestlings on reclaimed sites suffered mean parasitic burdens about twice that of those on the reference site; and for comparable parasitic load, they exhibited greater pathologic effects (e.g., decreased body mass) than control nestlings. The heavy blow fly infestation on oil sands-impacted wetlands suggests that oil sands mining disturbs several components of the local ecosystem, including habitat characteristics, blow fly predators, and host resistance to parasites.     

Cavity use and breeding biology of endangered Bahama Swallows (Tachycineta cyaneoviridis): implications for conservation

Bird populations, especially on islands, have declined or gone extinct due to overhunting, habitat loss and fragmentation, and adverse effects of the introduction of non-native species. Bahama Swallows (Tachycineta cyaneoviridis), endangered secondary cavity nesters that breed on only three islands in the northern Bahamas, are an island species with a declining population, but the causes of this decline are unknown. During four breeding seasons on Great Abaco Island (2014–2017), we identified cavity-nesting resources used by breeding swallows in native pine forest and other habitats, and estimated phenology and breeding parameters from a subset of nests. Bahama Swallows nest in cavities in a variety of structures, but rely most on woodpecker-excavated cavities in pine snags and utility poles. Swallows nesting in cavities in pine snags had higher fledging success (92%) than those nesting in cavities in utility poles (50–62%), which were concentrated in non-pine habitat that may expose swallows to predation and increased competition for nest cavities from other species. The high reproductive success of Bahama Swallows nesting in the pine forest indicates that the decline in population cannot be attributed to poor productivity on southern Great Abaco. However, our results suggest that the dependence of Bahama Swallows on cavities excavated by Hairy Woodpeckers (Dryobates villosus) for nesting sites may be a factor in their decline, and highlight the potential importance of the protection and management of pine forests in future efforts to ensure the survival of Bahama Swallows.     

Temperature effects on food supply and chick mortality in tree swallows (Tachycineta bicolor)

Tree swallow (Tachycineta bicolor) breeding success in Ithaca, NY, USA, over the past quarter century has shown generally healthy fledgling production punctuated by years of high nestling mortality. This study tested the potential effects that temperature may have on the food supply and breeding success of swallows. Data from 17 years of daily insect samples were used to relate flying insect abundances to daily maximum temperatures and to define “cold snaps” as strings of consecutive days when the maximum temperatures did not exceed critical temperatures. The distributions of cold snaps and chick mortality events were investigated both through detailed reconstructions of the fates and fate dates of individual chicks, focused on the three breeding seasons of lowest fledging success, and with less detailed brood-level analyses of a larger 11-year dataset including years of more moderate mortality. Mark–recapture analyses of daily brood survival rate (DSR) reveal very strong support for the effects of cold temperatures on brood survival rates, and all the top models agree on a critical temperature of 18.5 °C for insect flight activity in Ithaca. The individual-level analyses, focused on years of higher mortality, favored a 3-day cold snap definition as the most predictive of DSR effects, whereas the larger-scale brood-level analyses revealed 1- and 2-day cold snaps as having the most significant effects on DSR. Regardless, all analyses reveal that, in an age of generally warmer climates, the largest effect of weather on swallow fledgling production is from cold temperatures.     

Reproductive effort reduces long-term immune function in breeding tree swallows (Tachycineta bicolor)

We examined whether strategies of reproductive allocation may reduce long-term immunocompetence through the effects of manipulated effort on secondary or acquired immunity. We tested whether increased reproductive effort leads to reduced immune function and survival by manipulating brood size in tree swallows (Tachycineta bicolor) and exposing breeding females to a primary and secondary exposure of sheep red blood cells to elicit a humoral immune response. Females raising enlarged broods produced fewer secondary antibodies than did females raising control or reduced broods. Most importantly, individuals with high secondary responses were more likely to survive to breed 3 years after brood manipulations, suggesting that differences in disease susceptibility may be caused by trade-offs in reproductive allocation. We also found that individual quality, measured by clutch initiation date, mediated the effects of brood manipulations, with higher-quality birds showing a greater ability to deal with increases in effort.     

Population decline in tree swallows ( Tachycineta bicolor ) linked to climate change and inclement weather on the breeding ground

Population decline and the threat of extinction are realities currently facing many species. Yet, in most cases, the detailed demographic data necessary to identify causes of population decline are unavailable. Using 43 years (1975−2017) of data from a box-nesting population of tree swallows (Tachycineta bicolor), we identified reduced survival of offspring as a probable demographic cause of population decline. Poor fledging success was associated with increased predation and poor weather conditions during early nestling development. Low juvenile survival and subsequent recruitment was linked to poor weather conditions during the post-fledging period and may also be linked to conditions on the wintering grounds. Regional weather conditions during critical stages of breeding (early nestling and post-fledging) have become progressively worse over the 43-year study period. None of the other factors linked to offspring survival have similarly deteriorated. Overall, our results suggest tree swallows should be added to the growing list of species challenged by climate change, and that other species of aerial insect specialists may face similar impacts of climate change.     

High levels of extra-pair paternity in an isolated, low-density, island population of tree swallows (Tachycineta bicolor)

Molecular genetic studies have suggested that apparently nonbreeding males ('floaters') may account for a significant proportion of extra-pair paternity (EPP) in avian populations. Attempts to determine the influence of breeding density on EPP are therefore confounded by the presence of a subpopulation of floaters whose numbers are difficult to estimate. To study EPP in a tree swallow (Tachycineta bicolor) population with few floaters, we chose a nestbox grid on an island with an excess of available breeding sites and very few floaters. We obtained DNA samples from 13 complete families and performed DNA profiling on them using four microsatellite loci. For comparison, we also obtained a sample of 58 extra-pair young (EPY) from a mainland population typed at five microsatellite loci. Paternity assignments among resident males in both populations were made using the microsatellite profiles and a likelihood-based statistical method. Of the 67 island nestlings typed, we found 37 (55%) nestlings from 11 (85%) different nests that were EPY. The proportion of nestlings that were EPY and the proportion of broods containing EPY did not differ significantly between island and mainland populations studied previously. There was no significant difference between island and mainland populations in the proportion of extra-pair paternities assigned among neighbouring resident males. Male breeding density does not appear to affect the ability of female tree swallows to obtain extra-pair fertilizations, at least over the range of densities studied so far. The rate of EPP has remained remarkably consistent over many years, studies and populations implying an important role of active female choice in determining EPP.     

Timing of copulations and the pattern of paternity in relation to laying order in tree swallows Tachycineta bicolor

It is well established that extra-pair young are present to varying degrees in nests of most avian species. However, the timing of extra-pair copulations during a female's fertile cycle is not well known, nor is how these copulations translate into the distribution of extra-pair young within the laying sequence. We determined the rates of successful within-pair copulations where at least one cloacal contact was achieved, and the number of extra-pair copulations/attempts in 17 pairs of tree swallows Tachycineta bicolor during the mornings encompassing the female's fertile period. We also examined nestling paternity with respect to laying order in eighteen mixed-paternity tree swallow broods. The rate of successful within-pair copulations peaked three days prior to a female laying her first egg, and decreased steadily throughout the laying period. We therefore predicted that extra-pair young would be more likely to hatch out of later-laid than earlier-laid eggs within a female's brood. We found no evidence of an increased frequency of extra-pair young with laying order. Instead, extra-pair young appear to be randomly distributed within the laying sequence of tree swallows.     

The price of insulation: costs and benefits of feather delivery to nests for male tree swallows Tachycineta bicolor

Many species of birds line their nests with feathers, and it has been hypothesized that this functions to provide a thermally stable microenvironment for the development of eggs and nestlings. Feathers in the nest may also function as a mechanism for parasite control, providing a physical barrier that protects nestlings from ectoparasites. We tested these hypotheses by performing a feather removal and addition experiment in tree swallows Tachycineta bicolor, a species well‐known for lining their nests with feathers. While we found no evidence that quantity of feathers in nests influenced the ability of females to produce and incubate eggs, offspring in well‐feathered nests had longer flight feathers and were structurally larger just prior to fledging that those in nests with fewer feathers. Furthermore, we also demonstrated a positive correlation between feathers and the abundance of larval blow flies Protocalliphora spp. in nests, a result opposite to that predicted by the anti‐parasite hypothesis. While our study provides strong support for the insulation hypothesis, we also discuss the possibility that devoting time to feather gathering may result in males losing paternity in their nests, although manipulative studies will be necessary to fully evaluate this idea.     

Warm temperatures lead to early onset of incubation, shorter incubation periods and greater hatching asynchrony in tree swallows Tachycineta bicolor at the extremes of their range

The onset of incubation varies in birds, with many species beginning incubation prior to clutch completion. Here we examine whether early onset is more likely to occur during high temperatures, a critical prediction of the egg-viability hypothesis, which suggest that birds begin incubation prior to clutch completion in order to maintain egg-viability. We examined onset of incubation in tree swallows Tachycineta bicolor at two locations at the extremes of their breeding range, Alaska and Tennessee. A majority of individuals (68%) began incubation prior to clutch completion. While females in Tennessee were more likely to begin incubation early, there was no difference between sites when differences in temperatures inside nestboxes were controlled in analyses. Rather, early onset of incubation was predicted by the proportion of daily temperatures above physiological zero during laying, a critical prediction of the egg viability hypothesis. Both warm weather and early onset led to shorter incubation periods and increased levels of hatching asynchrony. We found no effect of timing of nesting, female body condition index or clutch size on the probability of beginning incubation prior to clutch completion. Our results are consistent with the egg viability hypothesis, not consistent with a threshold clutch size rule, and do not support the hurry-up hypothesis, that individuals breeding later in the season would begin incubation early to reduce the time spent nesting. Overall, our results suggest that broad scale geographic differences in incubation behaviour may be explained by individual-level responses to environmental conditions.     

Date and parental quality effects in the seasonal decline in reproductive performance of the Tree Swallow Tachycineta bicolor: interpreting results in light of potential experimental bias

In many bird species early breeders have higher reproductive performance than late breeders from the same population. This could be caused by a reduction in environmental factors related to date per se (Date Hypothesis), or because poorer performers nest later (Parent Quality Hypothesis). We manipulated hatch date of Tree Swallows Tachycineta bicolor by switching clutches with different lay dates, generating broods with advanced or delayed timing, and assessed the impact of the experiment on nestling mass. The Date Hypothesis better explained the decline in nestling mass in the first half of the season, while the Parent Quality Hypothesis was supported in the second half. We also found that female mass loss was unintentionally reduced in advanced females and suggest that such impacts of the experiment on parent quality, or correlations between nestlings and their actual parents via heritability or maternal effects, could bias hatch-date manipulation experiments towards supporting the Date Hypothesis. Differential costs of incubation, either due to naturally low temperatures early in the season, or due to the unintentional manipulation of female incubation costs, appear to have driven support for the Date Hypothesis early in the season.     

The Mitogenome Relationships and Phylogeography of Barn Swallows (Hirundo rustica)

The barn swallow (Hirundo rustica) poses a number of fascinating scientific questions, including the taxonomic status of postulated subspecies. Here, we obtained and assessed the sequence variation of 411 complete mitogenomes, mainly from the European H. r. rustica, but other subspecies as well. In almost every case, we observed subspecies-specific haplogroups, which we employed together with estimated radiation times to postulate a model for the geographical and temporal worldwide spread of the species. The female barn swallow carrying the Hirundo rustica ancestral mitogenome left Africa (or its vicinity) around 280 thousand years ago (kya), and her descendants expanded first into Eurasia and then, at least 51 kya, into the Americas, from where a relatively recent (<20 kya) back migration to Asia took place. The exception to the haplogroup subspecies specificity is represented by the sedentary Levantine H. r. transitiva that extensively shares haplogroup A with the migratory European H. r. rustica and, to a lesser extent, haplogroup B with the Egyptian H. r. savignii. Our data indicate that rustica and transitiva most likely derive from a sedentary Levantine population source that split at the end of the Younger Dryas (YD) (11.7 kya). Since then, however, transitiva received genetic inputs from and admixed with both the closely related rustica and the adjacent savignii. Demographic analyses confirm this species’ strong link with climate fluctuations and human activities making it an excellent indicator for monitoring and assessing the impact of current global changes on wildlife.