Effects of nest-box density on the behavior of Tree Swallows during nest building

ABSTRACT.   In territorial species, increased density is often linked to an increase in aggressive interactions, which may result in trade-offs between competitive behavior and nest construction. We examined the impact of nesting in areas of high-density versus low-density nest boxes on conspecific interactions and nest-building effort in a population of Tree Swallows ( Tachycineta bicolor ). We also examined whether expected differences in behavior related to variation in nest quality and reproductive success in high-density and low-density areas. No differences in either nest-building behavior or reproductive success were observed between areas of high-density and low-density boxes, but there was a tendency of more frequent behavioral interactions at high density. Similarly, there was a significant difference between pairs defending single nest boxes and those defending multiple nest boxes in the number of interactions with conspecifics. These results suggest that although there may be more competition with conspecifics for Tree Swallows nesting at high density, this does not appear to affect either nest quality or reproductive success.     

Influence of helping and breeding experience on reproductive performance in the Seychelles warbler: a translocation experiment

Reproductive success of the cooperative breeding Seychelleswarbler (Acrocephalus sechellensis) increases with age. Thisage effect is not due to differential survival or increasedreproductive effort, but to accumulated helping and breedingexperience. In their first year of breeding, reproductive performanceof inexperienced warblers with neither helping nor breedingexperience was significandy lower than that of warblers of thesame age with either previous helping or breeding experience.Reproductive performance was the same for primiparae with helpingexperience and for birds with breeding experience. Female primiparaewith helping experience or breeding experience built betternests and spent more time incubating than inexperienced females,which led to increased hatching success. Male primiparae withhelping experience or males with breeding experience guardedthe clutch better than inexperienced males, which led to reducedegg predation. Even-aged warblers with different previous experienceswere transferred to unoccupied islands, where birds startedbreeding immediately in high-quality territories. The experimentshowed that birds with helping experience produced their firstfledgling as fast as experienced breeders, and significandyfaster than inexperienced birds. Breeding performance did notimprove further with experience after the first successful breedingattempt. Only birds with previous breeding experience who pairedwith inexperienced birds, were likely to change mate. The otherpair combinations remained stable. Thus, primiparous birds withhelping experience have greater lifetime reproductive successthan inexperienced primiparae of the same age. This experimentshows that helping behavior has not only been selected for inthe context of promoting an individual's indirect fitness, butalso in the context of gaining helping experience which translatesinto improved reproductive success when a helper becomes a breeder.[Behav Ecol 7: 326-333 (1996)]     

Nest box orientation affects internal temperature and nest site selection by Tree Swallows

ABSTRACT.   Orientation of nests can influence nest microclimate, particularly temperature. However, few investigators have examined orientation preference and microclimate simultaneously. We examined the possible correlation between entrance orientation of artificial nest boxes used by Tree Swallows ( Tachycineta bicolor ) and the internal temperature of boxes. Tree Swallows showed a preference for east- and south-facing boxes, but only during the first half of the breeding season (before 1 June). During the second half of the breeding season (after 1 June), Swallows selected boxes based on availability. We found that east- and south-facing boxes were warmer than north- and west-facing boxes, but only during the first half of the breeding season when those boxes were preferred. Entrance orientation and box temperature were only correlated during the morning (06:00–12:00); the temperature of all boxes was similar during the afternoon. Our results suggest that Tree Swallows show a preference for nest boxes with a certain entrance orientation only when orientation influences microclimate, suggesting that warmer nest temperatures may provide fitness benefits.     

Effects of breeding density, synchrony, and experience on extrapair paternity in tree swallows

Breeding density, synchrony, and experience are expected toinfluence the frequency of extrapair paternity in birds. UsingDNA fingerprinting, we examined the effect of these factorson tree swallows nesting at relatively high (grids of nest-boxes)and low (solitary boxes at least 100 m from the nearest neighbor)densities and in relatively synchronous (Alberta) and asynchronous(Ontario) populations in Canada. The mean percentage of extrapairoffspring per nest did not differ significantly between birdsnesting in grids (43%, n = 22 families) and solitary boxes (57%,n = 12 families). Similarly, there was no significant differencein the mean percentage of extrapair offspring per nest betweenrelatively synchronous (60%, n = 12 Alberta families) and asynchronous(41%, n = 22 Ontario families) populations. We also found noconsistent pattern between extrapair paternity and breedingexperience among seven males and seven females examined overtwo to three breeding seasons. Female tree swallows can influencethe fertilization success of extrapair males by active selectionand rejection of copulation partners. We suggest that this abilitylimits the predicted effect of various ecological factors onthe frequency of extrapair paternity     

Effects of early experience on the reproductive performance of Atlantic salmon

To examine the link between early experience and subsequentreproductive performance, we experimentally manipulated theearly experience of Atlantic salmon (Salmo salar). Salmon ofa common genetic background were reared as juveniles eithernaturally in the river or artificially in a hatchery (sea ranched),depriving them of river experience, and then allowed to growto maturity naturally in the ocean. At maturity, the competitiveand reproductive abilities of these fish were compared in fourexperiments quantifying reproductive success. Although levelsof aggressive behavior were similar, the experience-deprived,sea-ranched males were involved in more prolonged aggressiveencounters and incurred greater wounding and mortality thanwild males. Furthermore, sea-ranched males were less able tomonopolize spawnings and as a result obtained 51% the reproductivesuccess of wild males across the experiments. This reproductiveinferiority varied directly with the male density and bias inthe sex ratio, reflecting the intensity of male breeding competition.A lower intensity of female than male competition was likelyresponsible for the lack of differences in breeding performancebetween sea-ranched and wild females. Sea-ranched females, however,produced smaller eggs than wild females, apparently in responseto their higher juvenile growth rate. Differences in migratorybehavior were also apparent, as sea-ranched males and femalesascended the River Imsa later than wild fish. Our results indicatethat early experience has implications for subsequent adultreproductive performance, affecting the development of specializedskills and traits important not only for early life, but alsolater life.     

Age-dependent reproductive costs and the role of breeding skills in the Collared flycatcher

This study addressed whether there are any age‐related differences in reproductive costs. Of especial interest was whether young individuals increased their reproductive effort, and thereby their reproductive cost, as much as older birds when brood size was enlarged. To address these questions, a brood‐size manipulation experiment with reciprocal cross‐fostering of nestlings of young and middle‐aged female Collared flycatchers, Ficedula albicollis, was performed on the Swedish island of Gotland. Nestlings’ body mass, tarsus length and survival were recorded to estimate the parental ability and parental effort of the experimental female birds. Female survival and clutch size were recorded in the following years to estimate reproductive costs. We found that middle‐aged female flycatchers coped better with enlarged broods than younger females or invested more in reproduction. In the following year, young female birds that had raised enlarged broods laid smaller clutches than the females from all the other experimental groups. This result shows that the young female birds pay higher reproductive costs than the middle‐aged females. Both young and middle‐aged female flycatchers seemed to increase their reproductive effort when brood size was increased. However, such an increase resulted in higher reproductive costs for the young females. The difference in reproductive costs between birds of different ages is most likely a result of insufficient breeding skills of the young individuals.     

Clutch and egg size of Tree Swallows along an elevational gradient

How avian reproductive behavior changes at high elevations, and whether changes are the result of adaptation or constraint remains unclear. We compared clutch and egg sizes in two populations of Tree Swallows (Tachycineta bicolor), one at an elevation of ~2500 m a.s.l. in Wyoming's Bighorn Mountains and the other at ~1350 m at the base of these mountains. Females at the high‐elevation site began laying 10 d later, on average, than females at the lower site. Females at the high site laid an average of 0.4 fewer eggs than those at the low site, a significant difference. Eggs were also slightly (2.3%), but significantly, smaller at the high site. Smaller clutches and eggs at the high site may have resulted in part from greater energetic constraints on females. Females at this site faced colder, windier weather during egg formation which would have increased thermoregulatory costs while simultaneously reducing the abundance of prey, i.e., flying insects. Laying a relatively small clutch at the high‐elevation site may also be adaptive, however. Having a smaller brood could help ensure there is an adequate supply of food for each offspring during bouts of inclement weather that are more severe at higher elevations. Also, if the delay in breeding inherent at high elevations reduces the survival prospects of the young, then producing fewer young could enhance a female's chances of breeding again, perhaps at a different location.     

Sprague's pipit breeding biology and reproductive success in planted and native grasslands

Much of the native grasslands in agricultural regions have been converted to cropland or tilled and seeded with non‐native grasses for livestock production. Several grassland songbird species occupy planted grasslands, but occupancy or density may not be a reliable indicator of habitat quality. I studied the breeding biology of Sprague's pipit Anthus spragueii from 2004 to 2008 in Saskatchewan, Canada. My objective was to determine the extent to which the breeding biology, density and reproductive success of pipits varied in planted and native grasslands. Peak clutch initiation occurred in mid‐ to late‐May in planted and native grassland. Peak pipit density also occurred in May, but density drastically declined over the breeding season in planted grassland. Clutch size varied among years and declined over the breeding season, but was similar in planted (4.7  0.1 SE) and native grasslands (4.5  0.1 SE). Daily nest survival rates varied with age of the nest and date, but the relationships differed in the two habitats and was likely a result of lower nestling survival in planted grassland compared to native grassland. The number of young fledged per nest increased as the season progressed and tended to be greater in native (1.2  0.1 SE) than planted (0.9  0.2 SE) grasslands. Seasonal productivity was much greater in native grassland. Only three nests were initiated after May in planted grassland and all were unsuccessful, whereas pipit young fledged at higher rates from nests initiated in native grassland in June and July than planted grassland nests initiated in May. The number of fledged young from successful nests did not vary strongly with habitat, date or year. This research indicates that planted grasslands attract pipits at the beginning of the breeding season, but habitat suitability and reproductive success substantially declines as the breeding season progresses compared to that found in native grassland.     

Negligible effects of blood sampling on reproductive performance and return rates of Tree Swallows

Blood sampling is a frequently used method of collecting genetic and physiological data in natural populations, and understanding the possible impact of blood sampling on individuals and populations is important, both for the welfare of study organisms and to avoid introducing bias into analyses using bled individuals. Most studies of birds have revealed minimal negative effects of blood sampling. However, Brown and Brown (2009. Auk 126: 853–861) found that blood sampling reduced return rates of Cliff Swallows (Petrochelidon pyrrhonota), suggesting that these results are not always generalizable and that swallows (Hirundinidae) may be particularly sensitive to blood sampling. We examined the possible effects of blood sampling on the reproductive performance of female Tree Swallows (Tachycineta bicolor) and the return rates of both females and offspring in a population in New York state. To reduce the chances of Type II error, we tested 15 possible effects of blood sampling on reproductive performance and return rates using generalized linear mixed models. Overall, our results suggest that blood sampling had few negative effects on within‐year reproductive success or survival of adult females. The one exception was that bleeding nestlings had a negative effect on the number of young that fledged in broods of five or six nestlings. Bleeding did not negatively impact return rates of females or nestlings in our Tree Swallow population. Our results support those of other studies suggesting that blood sampling has few negative effects on within‐year reproductive success or survival of adult females, and reiterate the importance of systematically testing for possible effects with datasets compiled over multiple years because such effects may be impossible to detect via direct monitoring during a single breeding season.     

The application of reproductive technology to endangered species breeding programmes

Captive breeding plays an increasingly important role in species conservation, but special problems are encountered in achieving the ideal of a demographically stable but genetically diverse population. Breeding programmes involving co-operation among a number of centres are now being developed which will overcome some of these difficulties by identifying individual animals, genetic lineages or age cohorts from which to breed. Application of techniques such as artificial insemination, embryo transfer and semen collection and storage, as well as the monitoring of reproductive status will contribute to the success of such programmes. The usefulness of these procedures for various population problems is discussed and criteria for their appropriate implementation within breeding programmes is outlined.     

Carry-over effect of captive breeding reduces reproductive fitness of wild-born descendants in the wild

Supplementation of wild populations with captive-bred organisms is a common practice for conservation of threatened wild populations. Yet it is largely unknown whether such programmes actually help population size recovery. While a negative genetic effect of captive breeding that decreases fitness of captive-bred organisms has been detected, there is no direct evidence for a carry-over effect of captive breeding in their wild-born descendants, which would drag down the fitness of the wild population in subsequent generations. In this study, we use genetic parentage assignments to reconstruct a pedigree and estimate reproductive fitness of the wild-born descendants of captive-bred parents in a supplemented population of steelhead trout (Oncorhynchus mykiss). The estimated fitness varied among years, but overall relative reproductive fitness was only 37 per cent in wild-born fish from two captive-bred parents and 87 per cent in those from one captive-bred and one wild parent (relative to those from two wild parents). Our results suggest a significant carry-over effect of captive breeding, which has negative influence on the size of the wild population in the generation after supplementation. In this population, the population fitness could have been 8 per cent higher if there was no carry-over effect during the study period.     

Humoral immunocompetence correlates with date of egg-laying and reflects work load in female tree swallows

Because quality differences between individuals affect fitness,much research has attempted, with limited success, to relatephysiological condition (e.g., body reserves), to differencesin life history between individuals. Recently, it has beensuggested that immunocompetence may reflect condition, andit thus may mediate variation in individual quality and reproductiveperformance and, ultimately, fitness. We measured humoral immunocompetence(HIC) by immunizing female tree swallows with a harmless antigenand measured the specific antibody responses in a novel enzyme-linked immunosorbent assay developed for passerine birds. HIC was stronglycorrelated with egg-laying date, an important determinant ofreproductive success in female tree swallows. We also investigatedthe effect of increased workload on HIC by manipulating femaleflight costs by clipping flight feathers. Clipped females hadlower HIC than nonclipped females. These data suggest that HICis a measure that may reflect phenotypic quality and also appearsto be sensitive to increased workload in female tree swallows.     

Cost of reproduction and the evolution of deferred breeding in the western gull

Survival patterns in a population of western gulls (Larus occidentalis)of known age of first breeding, a, indicate a cost of reproductionrelated to the age of initial breeding. Among both sexes, birdsthat commenced breeding at the earliest ages (3 years in malesand 4 years in females) had higher annual mortality than thosethat deferred breeding one or more years. In addition, females(but not males) evidenced a cumulative cost of reproduction:holding age constant, females with more annual breeding attemptsdemonstrated poorer survival. These patterns of -specific survivalwere statistically significant after controlling for interannualvariation in food availability and are not explained simplyby variation in the intrinsic quality of individuals. To assessthe effects of these sex-specific costs on fitness, we combinedthe observed survival patterns with data on prebreeding survivorshipand -specific reproductive success to estimate rates of populationgrowth and lifetime reproductive success for different agesat first reproduction. Males showed a clearly defined fitnessoptimum at = 4 years, which coincided with the modal for malesin the population. Females showed no clear optimum, except thatbreeding at age 4 was suboptimal, hence females benefited fromdeferring breeding to ages 5-7 years. Observed age of firstbreeding also showed no clear mode for females, with slightpeaks at ages 5 and 7. As a result, in both sexes, the fitnesssurface for corresponded well with observed frequencies of. We suggest that stabilizing selection has acted to shape thephenotypic distribution of in males but, due to trade-offsbetween survival and early reproduction, stabilizing selectionis weak or absent in females     

Extra‐pair paternity in a population of Chilean Swallows breeding at 54 degrees south

Geographic patterns of variation in life‐history traits have puzzled researchers for decades. However, the widely accepted idea that mating systems exhibit a tropical–temperate latitudinal trend, with extra‐pair mating systems being the norm among temperate species and genetic monogamy the norm among tropical species, is supported by sparse data, particularly for birds breeding in the tropics and even more so for birds that breed in the southern hemisphere temperate zone. Our objective was to examine the genetic mating system of Chilean Swallows (Tachycineta meyeni) breeding at 54°S in Tierra del Fuego, Argentina. From 2006 to 2009, we examined the paternity of young in 52 broods. Contrary to predictions based on their congeners that breed at equivalent latitudes in the northern hemisphere, Chilean Swallows in our study had low rates of extra‐pair paternity (EPP), with 13.5% of nests (N = 52) having at least one extra‐pair young and 6.8% of all nestlings (N = 161) fathered by extra‐pair males. These rates are also lower than those reported for species of Tachycineta swallows that breed at tropical latitudes. We found no support for a tropical–south temperate latitudinal cline in EPP rates. The highly unpredictable weather of the island of Tierra del Fuego might be influencing parental investment at this site; small clutch sizes and low EPP rates are expected if biparental attention is crucial for chick survival and reproductive success for these aerial insectivores. We argue that the sparse sampling of mating systems in birds worldwide may have contributed to a misapprehension of a global pattern. More studies of tropical and south‐temperate systems are needed to build on theories based on a wider set of taxa.     

Demography of male reproductive queues in cooperatively breeding superb fairy-wrens Malurus cyaneus

1. Subordinate helpers in cooperative societies may gain both immediate and future benefits, including paternity and territorial inheritance. However, if such opportunities correlate with rank in the queue, it is unclear why such queues should be stable. 2. In cooperatively breeding superb fairy-wrens Malurus cyaneus, only males are generally philopatric, and form stable hierarchical queues for the dominant position. 3. Male opportunities for reproduction are influenced both by their dominance status within the group, and their relatedness to the breeding female. For young queuing subordinates, the breeding female is typically their mother. Because of incest avoidance, reproduction is possible only through extra-group mating, even if the dominant position is achieved while the mother is still on the territory. If the mother dies while the helper is still a subordinate, he can seek matings both outside the group, and with the unrelated replacement female within the group. Finally, males can achieve the dominant position and pair with an unrelated female by inheritance, dispersal to a neighbouring vacancy, or by forming a liaison with an immigrant subordinate female that causes fission of the natal territory. 4. On average males spent more time living with unrelated females than with their mother. Subordinate males gained no survival advantages when living with their mother rather than an unrelated female, contrary to the prediction that parents facilitate the survival of their offspring. 5. Dominants and subordinates also had similar survival. Mortality accelerated over time, probably because older males invest more in extra-group courtship display. 6. Fairy-wren queues are likely to be stable because older birds are superior, and because extra-pair mating provides direct benefits to subordinates.     

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.     

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.     

Climatic effects on the breeding phenology and reproductive success of an arctic-nesting goose species

Climate warming is pronounced in the Arctic and migratory birds are expected to be among the most affected species. We examined the effects of local and regional climatic variations on the breeding phenology and reproductive success of greater snow geese ( Chen caerulescens atlantica ), a migratory species nesting in the Canadian Arctic. We used a long-term dataset based on the monitoring of 5447 nests and the measurements of 19 234 goslings over 16 years (1989–2004) on Bylot Island. About 50% of variation in the reproductive phenology of individuals was explained by spring climatic factors. High mean temperatures and, to a lesser extent, low snow cover in spring were associated with an increase in nest density and early egg-laying and hatching dates. High temperature in spring and high early summer rainfall were positively related to nesting success. These effects may result from a reduction in egg predation rate when the density of nesting geese is high and when increased water availability allows females to stay close to their nest during incubation recesses. Summer brood loss and production of young at the end of the summer increased when values of the summer Arctic Oscillation (AO) index were either very positive (low temperatures) or very negative (high temperatures), indicating that these components of the breeding success were most influenced by the regional summer climate. Gosling mass and size near fledging were reduced in years with high spring temperatures. This effect is likely due to a reduced availability of high quality food in years with early spring, either due to food depletion resulting from high brood density or a mismatch between hatching date of goslings and the timing of the peak of plant quality. Our analysis suggests that climate warming should advance the reproductive phenology of geese, but that high spring temperatures and extreme values of the summer AO index may decrease their reproductive success up to fledging.