Predator-induced female behavior in the absence of male incubation feeding: an experimental study

Parental care plasticity is critical to understanding the ecological and evolutionary influence of nest predation on life history strategies. In birds, incubation imposes a trade-off between the requirements of females (i.e., food) and egg requirements (i.e., heat and protection from predators). However, studies on this topic are rare and usually restricted to species where the male feeds the incubating female, relaxing her incubation costs. Males and females can reduce their activity at the nest to avoid detection by predators. However, females could follow two alternative antipredator strategies: to delay their return to the nest to avoid attracting attention from the potential predator or to return to the nest as soon as possible to enhance nest concealment. In this study, we manipulated the perceived risk of nest predation of incubating common blackbirds (Turdus merula), a species without incubation feeding, to study female behavioral changes induced by nest predation risk. We show experimentally that female blackbirds can reduce their nest visits in the situation with higher nest predation risk. In addition, we confirm that females significantly delay their return to the nest in the presence of a nest predator, contradicting the nest concealment hypothesis. However, our results could be interpreted as a passive antipredator response (to minimize clues given to predators) or as an active antipredator response (to search for predators to expel them from their territories).     

Hatching asynchrony is constrained by parental nest attendance during laying

Hatching asynchrony is widespread amongst animals, but no consensus has yet emerged as to why asynchronous hatching has evolved. It is generally thought to have adaptive benefits during the raising of dependent young. However, here, we considered an alternative view of hatching asynchrony in birds as a consequence of factors acting at the onset of incubation. We recorded parental nest attendance behaviour during laying using continuous records of nest temperature in herring gulls, Larus argentatus. We tested whether nest attendance during laying was related to individual factors (clutch size and diet) and whether it had consequences on fitness outcomes (hatching spread, incubation period, hatching success and chick survival). Low nest attendance was associated with small clutch size, and independent of clutch size, pairs on a more marine diet had lower nest attendance than pairs on a lower trophic level terrestrial diet, possibly due to higher foraging effort for marine food. Broods hatched more asynchronous where pairs had a lower nest attendance during laying or took longer to complete a clutch and where the last egg took longer to hatch. Low nest attendance was also related to shorter incubation periods, possibly representing a strategy of birds in poor condition to reduce the demand of incubation by reducing the length of incubation. We found that low nest attendance during laying and increasing hatching asynchrony had detrimental effects on hatching success for small eggs laid early in the laying sequence. Increasing hatching asynchrony also had a detrimental effect on the survival of the youngest sibling. In our study population, hatching asynchrony was influenced by a more complex set of factors than simply onset of incubation and appears to be constrained by circumstances at the onset of incubation rather than to be an adaptive strategy. Thus, factors acting both during offspring rearing and at the onset of incubation need to be considered for a better understanding of hatching asynchrony.     

Fattening strategies in dominance-structured greenfinch (Carduelis chloris) flocks in winter

Relationships between social status and levels of body reserves stored by members of greenfinch (Carduelis chloris) flocks in winter were investigated. In addition, the adjustment of reserves by birds of different rank to experimental changes in food predictability and to changed weather conditions were examined. Birds with low social status carried overall larger body reserves than high-ranked birds. The results of the experiments suggest that this was mainly because subordinates, due to their low priority to food access, perceived future foraging success as less predictable than dominants. In response to severe weather, which probably increased the risk of starvation for birds, dominants temporarily increased their reserves more than subordinates. This response also indicated that birds with low social status carried larger reserves than high-ranked birds, and demonstrated that dominants could increase food intake when there was a risk of energetic shortfall. The results suggest that fattening strategies in greenfinches depend on social status. In winter, subordinates may be forced to carry larger reserves than dominants to safeguard against social constraints in access to food during critical times. As body reserves may be costly to carry and acquire, this should reduce the probability of surviving the winter for subordinates compared to dominants. Received: 2 March 1995/Accepted after revision: 13 April 1996     

Adjusting the timing of hatching to changing environmental conditions has fitness costs in blue tits

After laying the first egg, a bird can, to a certain extent, adjust the hatching date of the brood to environmental conditions. However, costs of this adjustment have remained largely unexplored. We studied potential costs of hatching delay in a population of blue tits in southern Finland. We explored the factors underlying hatching delay and investigated the association between hatching delay, clutch hatchability and female body condition. Finally, we reciprocally cross-fostered a large number of broods irrespective of their experienced hatching delay to address possible downstream effects of hatching delay on developmental parameters in offspring. We found that hatching delay was associated with early laying dates and low mean temperatures during the egg-laying phase. Furthermore, we found evidence that delayed hatching negatively affected the breeding performance. Hatchability of the clutch was lowered and the breeding female was energetically impaired, resulting in smaller clutch sizes, lower female body mass at hatching and lowered survival of nestlings reared in nests that had experienced a long hatching delay. In addition, delayed hatching had a significant negative effect on the body mass of nestlings prior to fledging. However, ultimately we did not find evidence that delayed hatching affected survival of the breeding female nor recruitment of fledglings in the local breeding population. Our study demonstrates that environmental conditions during egg laying can have lasting effects throughout the breeding and nestling phase. Furthermore, our results emphasize the importance of energetic tradeoffs by breeding females during the early breeding phase to manage reproductive costs.     

Egg adoption can explain joint egg-laying in common eiders

Hypotheses regarding the evolution and maintenance of intraspecific nest parasitism were tested with data collected during a 3-year study of common eiders (Somateria mollissima) breeding near Churchill, Manitoba. The nest parasitism rate was highest (42.4% of nests) during the year with the highest nest density and the best environmental conditions, and lowest (20.2% of nests) in the year with the lowest nest density and the poorest environmental conditions. Over the nesting season, parasitic eggs were laid at the same time as normally laid eggs. Most parasitic eggs (>75%) were laid before the host female laid her third egg. The majority of the parasitic eggs were the first or second egg produced by the parasitic female. When a parasitic egg was laid before or on the same day as the host female initiated her clutch, the probability of her first egg being depredated before incubation was significantly lowered. First- and second-laid eggs suffered a high rate of predation probably because nesting females do not attend their clutch until their second or third egg is laid. Hypotheses that some females use intraspecific nest parasitism to parasitize the parental care of other females were inconsistent with these data. Egg adoption is a likely explanation for the prevalence of females incubating parasitic eggs in this population. Received: 30 September 1997 / Accepted after revision: 6 May 1998     

Maternal energy expenditure does not change with flight costs or food availability in the pied flycatcher (Ficedula hypoleuca): costs and benefits for nestlings

We manipulated parental work load without changing brood size in a population of pied flycatchers Ficedula hypoleuca by removing two primaries (7 and 9) from each wing of females, thus reducing wing area and increasing flight costs. At other nests, we offered supplementary food in the form of live mealworms (10–20 g daily from hatching) to reduce brood demand and thus parental foraging costs. Other nests were left as controls. The daily energy expenditure of females feeding 12-day-old nestlings was measured with doubly labelled water D₂ ¹⁸O. Females in both treatments expended the same amount of energy, fed at the same rate and had similar body masses to birds in the control group. No effect of treatment on male mass and feeding effort was detected. More nestlings, however, died in nests of handicapped females. Nestlings of handicapped females had significantly lower body mass and haematocrit values than nestlings in food-supplemented nests, with nestlings in control nests occupying an intermediate position. The effects of both treatments on nestling mass, haematocrit values and mortality rates were only noticeable in nests infested with mites. Maternal energy expenditure is apparently constrained and offspring pay the costs imposed by reduced provisioning rate or increased demand caused by ectoparasites, while receiving benefits when food supply improves. The presumption that avian reproductive costs derive from changes in a flexible energy output may not be met in many cases. Received: 24 October 1998 / Received in revised form: 15 March 1999 / Accepted: 26 April 1999     

Incubation start and egg size in relation to body reserves in the common eider

Avian incubation is often initiated before all eggs are laid. In altricial birds this has been proposed to facilitate brood reduction through asynchronous hatching. However, in precocial birds eggs normally hatch synchronously even if incubation has started before all eggs are laid. Patterns of incubation start may be the adaptive trait selected for both in altricial and precocial species. Several hypotheses have been proposed to explain the timing of incubation start in birds. Decreasing egg-size after incubation start may be adaptively related to an early incubation start, either to ensure synchronous hatching or to decrease fitness cost of late hatched eggs. We have measured individual body condition, egg size and start of incubation in common eider Somateria mollissima, a precocial sea-duck which does not feed during the incubation period. Females in poor body condition start to incubate earlier in the laying sequence than those in good body condition. Furthermore females in poor body condition lay smaller final eggs than females in good body condition. The laying of smaller eggs late in the sequence is therefore probably related to energetic or nutritional state. We propose that females in poor body condition start to incubate early to shorten the nest period in order to reduce their mass loss, but at the cost of reduced size and growth of the ducklings from the eggs laid after incubation start. Females in good body condition on the other hand postpone incubation start at the cost of a longer incubation period and a higher mass loss to the benefit of synchronized hatching and a higher survival of ducklings.     

The relative role of age and experience in determining variation in body mass during the early breeding career of the Common Tern (<Emphasis Type="Italic">Sterna hirundo</Emphasis>)

Body mass is a consistent individual trait that characterises the state of adult birds and mammals and is positively related with long-term reproductive success. However, whether and to what extent body mass changes over lifetime in long-lived birds is poorly studied. In this paper, we investigate how individual body mass varies with age. Furthermore, we try to separate possible effects of age and experience on body mass. This study was conducted in a Common Tern colony on the German Wadden Sea coast. Transponders allowed registration of individuals throughout the breeding season and consecutive years with an antenna system combined with electronic balances for recording individual body-mass changes within and between years. Individual body masses of males and females were measured during three stages of the breeding season: at arrival, during incubation and chick rearing when mass was lowest in both sexes. Individual-based longitudinal analyses clearly showed that body mass during arrival, incubation and chick rearing increased up to an age of 5 or 6 years. First-time breeders had lower body mass than experienced breeders. Experience had stronger effects on incubation mass than age. Recruiting age also affected body-mass development of breeders: Three-year-old recruits showed a stronger increase in mass with experience than 4-year-old recruits. We assume that increasing experience enables birds to cope better with the physiological challenge of reproduction. To explain the general phenomenon of higher body mass in older birds, our results support the constraint hypothesis rather than either the selection or restraint hypothesis.     

Consequences of immune system aging in nature: a study of immunosenescence costs in free-living Tree Swallows

Immunosenescence, the aging of the immune system, is well documented in humans and laboratory models and is known to increase infection risk, morbidity, and mortality among the old. Immunosenescence patterns have recently been unveiled in various free-living populations, but their consequences in the wild have not been explored. We investigated the consequences of immunosenescence in free-living Tree Swallows Tachycineta bicolor through a field experiment simulating a bacterial infection (challenge with lipopolysaccharide, LPS) in females of different ages during the nestling rearing period. We assessed behavioral and physiological responses of females, as well as growth and quality of their offspring, to determine the costs associated with the simulated infection. Results of the experiment differed between the two years of study. In the first year, old females challenged with LPS lost more body mass and reduced their nest visitation rates more, and their offspring tended to grow slower compared to similarly challenged younger females. In contrast, in the second year, old females did not appear to suffer larger costs than younger ones. Interestingly, immunosenescence was only detected during the first year of the study, suggesting that it is the dysregulated immune function characteristic of immunosenescent individuals rather than age per se that can lead to higher costs of immune defense in old individuals. These results provide the first evidence of costs of immunosenescence in free-living animals and support the hypothesis that old, immunosenescent individuals pay higher costs than younger ones when faced with a challenge to their immune system. Our results also suggest that these costs are mediated by an exaggerated sickness behavior, as seen in laboratory models, and can be modulated by ecological factors such as weather conditions and food availability.     

Reproductive performance in lesser snow geese: are two parents essential?

Summary The roles of male and female lesser snow geese (Anser caerulescens) in offspring care are well documented, but we know little about flexibility in these roles and how essential they are for offspring survival. We asked whether uni-parental care was adequate, which sex was required at various stages of the reproductive cycle, and what the costs and consequences were of variable amounts of parental care. We found that two parents were important in acquiring nest sites and producing clutches. Widows losing mates during the latter part of laying or in early incubation experienced similar rates of success in hatching clutches to paired females, but males losing mates during incubation experienced total nest failure. Partial clutch loss, hatch loss, intraspecific nest parasitism, duration of incubation periods and gosling weights at hatch did not differ for pairs or widows. In 1983 at La Pérouse Bay, Manitoba, Canada, widows and pairs had similar incubation behaviour, but widows were harassed more and spent more time in full alert posture while on their nests. In 1952 and 1953 at Southhampton Island, Northwest Territories, widows were significantly lighter in body weight prior to hatch than paired females. Thus while widows can bring clutches to hatch successfully, the loss of mates may result in additional physiological costs. Although all possibilities have not been tested experimentally, it appears that costs to uni-parental care up to hatch are not significant. Data collected so far on consequences of uni-parental care provided during the post-hatch period are equivocal, as in one year survival of goslings from single parent broods seemed poor and in another it did not appeart to differ from pairs. The minimum amount of parental care required to raise snow goslings from hatch to recruitment has yet to be determined.     

Parental care of replacement clutches in common terns (Sterna hirundo)

In long-lived seabirds with low annual reproductive output, the renesting decision after breeding failure is critical, and the parents have to weigh benefits of replacement clutches against possible future reproductive costs. In this study, we investigated factors influencing renesting decisions in common terns (Sterna hirundo) and compared aspects of breeding biology and body mass between two breeding attempts by the same pairs in each of 4 years of heavy losses due to predation. Renesting birds were characterized by early laying dates and by a high age. Among early breeders, high egg mass reduced the probability of renesting. A long relaying interval coincided with low mass of replacement eggs in one year, and short intervals with high egg mass in another. Further, egg mass decreased and relaying intervals increased the later the predation events occurred. Evidence of high levels of parental care of replacement clutches came from body mass data: female mass increase prior to egg laying was higher in the second attempt than in the first, whereas male mass was lower during the second courtship period than during the first. Male mass also affected relaying intervals and mass of replacement eggs. We conclude that common terns expend high levels of parental care of replacement clutches. Intrinsic factors related to individual quality (age, body condition) seemed most important for renesting decisions and for the degree of parental care, but foraging conditions seemed to have modifying effects. Received: 13 August 1999 / Received in revised form: 5 February 2000 / Accepted: 13 March 2000     

Dispersal behaviour in a subsocial spider: group conflict and the effect of food availability

This study investigates proximate factors influencing dispersal behaviour in the subsocial spider Amaurobius ferox. Dispersal of spiderlings from the natal web occurred as a progressive sigmoidal function (mean duration of dispersal period=31 days), with considerable individual variability in developmental instar and body mass within the clutch at the time of dispersal. The spiderlings showed a significant decrease in group cohesion on the 6th day after their second moult, which corresponded to the beginning of the dispersal period. Mutual aggressiveness appeared when the spiderlings began to show predatory behaviour (4th day post-second moult) and increased over the course of the dispersal period. While lack of prey in the maternal nest accelerated dispersal behaviour, the addition of prey items lengthened the dispersal period in previously non-fed clutches. Individuals that dispersed were smaller than the remaining individuals when measured on the day 50% of the clutch had dispersed. Timing of the appearance of the developmental characters (second moult, predation activity, agonistic behaviour against siblings, reduction of group cohesion, dispersal) suggests that the dispersal trait might have evolved in consequence of these different functional behaviours. Received: 10 August 1999 / Received in revised form: 15 March 2000 / Accepted: 1 April 2000     

Compensating for the costs of polygyny in hen harriers Circus cyaneus

In polygynous species, the adults are faced with a dilemma during chick rearing. Males must decide how to distribute food between their females and food allocation patterns are often highly unequal. In turn, the females that receive less food from males have to decide how much time to invest in additional hunting. If they spend more time hunting, then they leave their young exposed to weather and predators. However, if they stay at the nest, they increase the risk of their chicks starving. One way that birds may compensate for reduced provisioning is by increasing the size of prey caught. We tested this hypothesis by comparing prey deliveries to nests of hen harriers, Circus cyaneus, with females of different breeding status. As expected, male harriers delivered less food items to the nests of polygynous females, and especially their secondary, or β females. However, both sexes were able to compensate by delivering larger items and there was no difference in the overall mass of food delivered to nests. Moreover, females spent a similar amount of time at the nest, irrespective of status, and there were no overall differences in breeding success. Our results show that polygynous female harriers can compensate for the costs of polygyny, but we suggest that their ability to do so will vary according to the abundance of both large prey and predators.     

Food and predators affect egg production in song sparrows

Although the possibility that food and predators may interact in limiting avian populations has long been recognized, there have been few attempts to test this experimentally in the field. We conducted a manipulative food addition experiment on the demography of Song Sparrows (Melospiza melodia) across sites that varied in predator abundance, near Victoria, British Columbia, Canada, over three consecutive breeding seasons. We previously showed that food and predators had interactive effects on annual reproductive success (young fledged per female). Here, we report the effects on egg production. Our results show that food limits the total number of eggs laid over the breeding season ("total egg production") and that interactive food and predator effects, including food effects on nest predation, determine how those eggs are "parceled out" into different nests. Food addition alone significantly affected total egg production, and there was no significant interannual variability in this result. At the same time, both food and predators affected the two determinants of total egg production: "clutch number" (total number of clutches laid) and average clutch size. Both clutch number and size were affected by a food x predator x year interaction. Clutch number was lower at low-predator locations because there was less nest predation and thus less renesting. Food addition also significantly reduced nest predation, but there was significant interannual variation in this effect. This interannual variation was responsible for the food x predator x year interactions because the larger the effect of food on nest predation in a given year, the smaller was the effect of food on clutch number; and the smaller the effect of food on clutch number, the larger was the effect of food on clutch size. Potential predator and year effects on total egg production were thus cancelled out by an inverse relationship between clutch number and clutch size. We suggest that combined food and predator effects on demography could be the norm in both birds and mammals.     

Polygyny and male parental care in Savannah sparrows: effects on female fitness

Summary To determine the effects of male mating status on female fitness, we compared the reproductive success, survival, and future fecundity of female Savannah sparrows (Passerculus sandwichensis) mated to monogamous vs. polygynous males in a 5-year study on Kent Island, New Brunswick, Canada. The proportion of males with more than one mate varied from 15 to 43% between years and sites. Polygynous and monogamous males fledged young of equal size in every year of the study. Females who shared paternal care with other females laid as many eggs per clutch and clutches per season as monogamously mated females. In most years polygynously mated females showed no delay in laying a second clutch, and they suffered no reduction in fecundity the following year. Recruitment of a female's offspring into the breeding population was generally independent of her mating status. Fitness costs of being mated to a polygynous male were only apparent in one year of the study, during which females mated to polygynous males had higher over-winter mortality than those mated to monogamous males. That same year, young raised by polygynous males were only one-third as likely to survive to reproductive maturity (as inferred by returns) as those raised by monogamous males. A male's mating status had no effect on his own survivorship. A male's mating status did not necessarily reflect his contributions to raising nestlings, which may partially explain why monogamously and polygynously mated females had equal fitness. At 35 nests the proportion of food deliveries brought by individual males varied from 0 to 75%; on average, males brought fewer than 30% of all food deliveries. Yet parental care by polygynous males was no less than that of monogamous males, at least at the nests of their primary females. Secondary females tended to receive less male assistance during the nestling stage, but their reproductive success was indistinguishable from that of primary females. Females feeding young without male assistance made as many food deliveries/h as did pairs in which males brought at least 30% of all food deliveries. Unassisted females did not suffer diminished fledging success or produce smaller fledglings. The benefits of polygyny for male Savannah sparrows are clear: polygynous males recruit more surviving offspring into the breeding population than monogamous males. The fitness of females, on the other hand, appears to be unaffected by whether their mate was monogamous or polygynous except in occasional years. Polygyny may be maintained in this population by the constraints of a female-biased sex ratio, the inability of females to predict a male's paternal care based on his morphology or behavior, the poor correlation between a male's mating status and his assistance at the nest, and inconsistent natural selection against mating with a polygynous male. Correspondence to: N.T. Wheelwright     

Do males and females differ in the feeding of large and small siblings? An experiment with the bluethroat

Males and females have been reported to differ in their feeding of large and small siblings in several species of birds. According to recent hypotheses, this phenomenon may be related to a sexual conflict over avian hatching patterns. We designed an experiment to test for the existence of such a sex difference by manipulating nestling size hierarchies of the bluethroat (Luscinia s. svecica) in two directions; half the broods were “asynchronized” to yield large size-differences within broods and the other half were “synchronized” to yield small size-differences. In all broods, nestlings were categorized as being either large or small according to body mass. We recorded male and female food distribution by video early (day 4 after hatching) and late (day 8) in the nestling period. Males and females did not differ in their distribution of food among different-sized nestlings. With large size-differences, both males and females fed large nestlings nearly twice as often as small ones. In contrast, when the size-differences were small, food was more evenly distributed among nestlings. Early in the nestling period, males fed more nestlings during each feeding visit than did females. Our finding that male and female bluethroats do not differ in the feeding of large and small siblings is in contrast to most previous studies. Variation in costs and benefits to males and females from feeding different-sized nestlings, and restrictions to parental choice due to nestling interactions, may explain interspecific variation. Received: 27 June 1997 / Accepted after revision: 26 January 1998     

Weight comparisons and litter size manipulation in Columbian ground squirrels (Spermophilus columbianus) show evidence of costs of reproduction

I studied reproductive costs in the female Columbian ground squirrel (Spermophilus columbianus) using individually marked animals. I compared weight changes during the active season and over winter, and mortality for females that did and did not wean young. Females raising young were heavier at emergence in that spring than unsuccessful ones. Females that did not raise young gained more weight during summer, were heavier than successful females at the time of entry into hibernation, and were heavier emerging from hibernation the following spring. Over-winter mortality was higher for females that reared young compared to reproductively unsuccessful females. A food supplementation experiment showed that energy-rich food can accelerate individuals’ weight gain. Interactions between litter size, birth weight, weight at emergence from the natal burrow, survival of young to yearling age, and maternal fitness were also studied. Litter sizes were experimentally manipulated to evaluate how females cope with costs of rearing one additional young. Birth weight of juveniles was positively correlated with survival to emergence from the natal burrow and with survival to yearling age. Partial litter loss was higher in experimentally enlarged litters than in either experimentally decreased or control litters. Total litter loss, survival of adult females or the probability of weaning young the following year were not affected by the litter size manipulation. Females appear to adjust the size of their litter before birth, and to some extent during lactation, to their ability to wean young. Received: 20 January 2000 / Received in revised form: 12 March 2000 / Accepted: 18 March 2000     

Management of fat reserves in tufted titmice (Parus bicolor ): evidence against a trade-off with food hoards

Caching species can manage their energy supply by adjusting body fat, number of caches, or both. It has been hypothesized that because body fat has a higher fitness cost than caches, small food-hoarding birds respond to increased starvation risk by increasing the number of their caches rather than their fat load. This hypothesis predicts that when birds cannot cache they should compensate for the loss of external energy storage by (1) shifting the time of their daily body mass accumulation toward earlier in the day and (2) increasing the overall level of their fat reserves. During the winter of 1995–1996, we tested these predictions with a caching species, the tufted titmouse (Parus bicolor). Each of six experimental birds was fed a diet of uncachable sunflower seed powder for 6 days, preceded and followed by 6-day control periods during which they were fed cachable sunflower seeds. The daily pattern of body mass gain was unaffected by the opportunity to cache. Furthermore, when unable to cache, the birds did not increase either their mean daily body mass, body mass in the middle of the day, or evening body mass compared to the two control periods. These results argue against the hypothesis of a trade-off between fat reserves and food caches in tufted titmice, and suggest that fat reserves are managed independently of external food caches. Received: 9 April 1997 / Accepted after revision: 30 August 1997     

Risk-taking by female ducks: intra- and interspecific tests of nest defense theory

We tested several predictions of nest defense theory by observing variation in flushing distance and probability of nest abandonment within and between six species of waterfowl. In these species, only the females incubate eggs and attend offspring. First, we examined whether flushing distance by females varied in relation to clutch size, stage of incubation, and time of season, after controlling for the number of visits made to nests by observers. Revisits by observers appeared to affect flushing distance by females for reasons unrelated to the relative value of the current clutch. We found that as incubation progressed, females allowed observers to approach more closely before flushing from the nest. In some species, females with larger clutches allowed closer approaches to nests before flushing which was also consistent with nest defense theory. In contrast, time of season (Julian date) did not relate to flushing distance for any species. When species were compared, we found that species with moderate to high yearly mortality and high reproductive output per breeding attempt (e.g., northern shoveler and blue-winged teal) were less likely to abandon nesting attempts and exhibited riskier behavior (remained at nests when approached closely by observers) than species that had lower yearly mortality (e.g., mallard). Our results show that flushing distance and patterns of nest abandonment by female ducks conform to several predictions of nest defense theory.Correspondence to: M.R.L. Forbes     

Fitness costs of an immune response in the house martin (<Emphasis Type="Italic">Delichon urbica</Emphasis>)

Immune responses constitute a major way for hosts to defend themselves against parasites. Because hosts do not habitually produce strong responses all the time, immune responses might be costly to produce or maintain. We tested experimentally if the production of a response to a challenge with a novel antigen resulted in a cost in terms of fitness using the highly colonial house martin Delichon urbica as a model system. We injected adult breeding birds during laying of the first clutch with either Newcastle disease virus (NDV) or a control injection, and the clutch was subsequently removed to induce relaying. NDV stimulates the non-specific immune system, causing production of antibodies during a period of more than 2 weeks. Accordingly, we found an increase in leukocyte counts in experimental birds compared to controls. Experimental treatment reduced the frequency of re-laying, caused a delay in timing of relaying and a reduction in brood size. Quality of nestlings in terms of body size, body mass and T-cell-mediated immune response did not differ significantly between treatments. Overall, seasonal reproductive success differed significantly between treatments, showing that the production of an immune response by adult birds is costly in terms of future fecundity.