FACTORS GOVERNING FEEDING RATE, FOOD REQUIREMENT AND BROOD SIZE OF NESTLING GREAT TITS PARUS MAJOR

SUMMARY Observations were made on feeding rates and food-consumption of nestling Great Tits Parus major mainly in Larch plantations at lake Yamanaka, Japan. Feeding frequencies were recorded by an automatic recorder. There were marked differences between early and late broods; the feeding frequencies were twice as great in early than in late broods of the same size. No clear tendency was observed in the variations of feeding frequencies in relation to brood size. There was, however, a clear inverse relationship between the frequencies and the average size of food brought to the nests. The males' share in terms of feeding frequencies is described. These figures, however, did not follow the males' contribution in terms of weight of food, which was nearly always higher than the females'. It is pointed out that feeding frequencies are far too variable to be used as a true index of food consumption by nestlings, and are not reliable. Attempts were made to measure the weight of food; the method is described. The average weight of food brought by males was lighter in early than in later broods. The total weight of food was estimated. The trend of daily food consumption per chick was similar to that of the chick's growth curve. It was found that up to about the tenth day of the nestling period daily food-intake per chick increased linearly as body weight increased. At some nests, rate of defaecation was observed. This was at first low, but it increased steeply on the third day, with a steady increase thereafter. By comparing the rates of food intake, faeces output, and weight increment of a chick, it was found that only 20–30% of digested matter (the difference between food-intake and faeces-output was used up daily (for body temperature regulation various external effort, etc.). The factors responsible for this high efficiency of growth in nestlings are discussed. There was a clear inverse relationship between the total weight of food brought per chick per day and the brood size. This is largely because the heat-loss is greater in small than in large broods, so that a chick from a small brood in fact needs more energy to maintain its body temperature after a certain age than one from a large brood. This is discussed in detail. Factors which caused variations in size of food are discussed in relation to feeding frequencies. It is pointed out that, because of the inverse relationship between energy requirement by each chick and brood size, the total food requirement by a brood as a whole did not vary directly in proportion to the brood size. An estimation showed that a b/3 still required about 75% of the total food required by a b/8. A smaller brood is less advantageous than expected to parents feeding nestlings when they encounter adverse conditions, e.g. food shortage in the habitat, or a lack of help by their mates, etc. On the other hand, it is suggested that once they have left the nest, the food-demand by a brood of fledglings the parents have to feed, so that, in the fledging period, in times of food shortage it would certainly be advantageous to have fewer young. It is suggested that, although fledglings may consume three to four times as much food as nestlings, the parents, in providing this food, would not work proportionately harder, since the parents' efficiency of providing food could be higher in feeding the fledglings, which always follow the parents as they are hunting, than in feeding the nestlings to which food has to be brought. On this basis, the adaptive significance of the length of the nestling period in nidicolous species is discussed in relation to clutch size, brood size and food requirement.     

Costs and benefits of parental care in eastern kingbirds

We manipulated brood sizes of eastern kingbirds (Tyrannus tyrannus)to measure the costs and benefits of parental care and to testwhether kingbirds showed evidence of individual optimizationof reproductive effort. We found that the number of feedingtrips (trips/h) increased and that per capita feeding rates(trips/nestling/h) declined as brood size increased. The declinein per capita feeding rates was mostly due to high feeding rateto broods of one: parents made roughly equal number of tripsto feed each nestling in broods of two to five. Nonetheless,nestling mass declined with brood size, probably because largebroods were fed more small prey. Nestling condition (mass adjustedfor structural size) differed only between broods of one andfive. After controlling for effects of brood size, feeding rateshad no supplementary influence on either nestling size or condition,but productivity and feeding rate were positively and significantlyrelated. Adult male condition did not vary with brood size,manipulated brood size, or total feeding rate, but declinedas the pair's per capita feeding rates increased. In addition,males that returned to breed were in better condition beforeleaving for migration than those that failed to return. Femalecondition tended to decline, and the probability of returningto breed dropped when broods were enlarged. However, femalecondition was independent of the probability of returning. Ourresults show that high feeding rates were costly, but that theycarried benefits (greater productivity). Some evidence for individualoptimization of reproductive effort existed: variability innestling and adult female condition were better explained bychanges in brood size than by the actual number of young inthe nest. However, most evidence supported the alternative thatincreased brood size was equally costly for all birds     

A comparative study of insect abundance and reproductive success of barn swallows Hirundo rustica in two urban habitats

Few studies have quantified the relative reproductive success of passerines in urban habitats. I studied food availability and reproductive success of barn swallows Hirundo rustica in two urban habitats during 2012–2015. Barn swallows breeding in the town center experienced lower insect densities than those in the town periphery. Lower food availability resulted in reduced feeding rates per capita, lower nestling body mass, longer nestling periods, longer inter‐clutch intervals, fewer first and second brood fledglings and a lower total number of fledglings produced during the breeding season in comparison to barn swallows breeding in the town periphery. I hypothesize that the lower intra‐specific competition for nest sites and fitness advantages linked to the solitary breeding in urban habitats balanced the apparent costs of reproduction in more urbanized habitats.     

Parental care behavior in the monogamous,sexually dimorphic Madagascar paradise flycatcher: sex differences and the effect of brood size

I studied the parental care behavior of the Madagascar paradise flycatcher Terpsiphone mutata in northwestern Madagascar. I especially focused on feeding, brooding and vigilance behaviors. Feeding rate did not differ between males and females, but females spent more time at the nest than males. Females dedicated their time to brooding, while males perched on the nest and were vigilant. Both parents changed the feeding rate in relation to brood size, so the feeding rate per nestling was not different among nests of different brood size. Duration of brooding by females increased with decreasing brood size, suggesting that the Royama effect, the pattern of lower feeding rate per nestling in larger broods, did not apply in this study. Males spent more time on vigilance than females. Anti-predator vigilance by males should be important for nestling survival given the high predation pressure typical of this population. In conclusion, males provide considerable parental care probably to minimize nestling starvation and to avoid nest predation. My results are not consistent with the general pattern of less parental effort by males in monogamous, sexually dimorphic species.     

Responses by Central‐Place Foragers to Manipulations of Brood Size: Parent Flickers Respond to Proximate Cues but do not Increase Work Rate

Manipulations of brood size measure the willingness or ability of parents to invest in offspring and different reproductive roles may lead to differences in feeding effort between the sexes. Parental investment in birds is usually assessed by quantifying feeding rates, but this provides an incomplete picture of parental effort because it fails to account for how parents collect food on the landscape. We studied northern flickers (Colaptes auratus), a woodpecker in which males provide the majority of parental care and used a repeated measures design and short‐term (24 h) brood enlargements (N = 35) and reductions (N = 27) to assess effects of treatment on feeding rates to nestlings and parental foraging behaviour. Parents of enlarged broods did not significantly increase feeding rate, resulting in a decline in nestling mass. Parents of reduced broods decreased their feeding rates by 84%, but increased per capita feeding rates, resulting in nestling mass gain. The variation in feeding rates to enlarged broods was not influenced by feather corticosterone, body condition, feather re‐growth rate or mass change between the incubation and nestling periods. Foraging pattern on the landscape remained the same during the enlarged treatment for both sexes. We conclude that flickers respond to proximate cues in brood demands, but do not increase feeding rates to enlarged broods, at least in the short term. A literature review suggested that this lack of response is atypical for short‐lived species. We hypothesize that parents in species with large home ranges and long nestling periods face energetic limitations that constrain their ability to respond to enlarged broods. We encourage future studies to assess foraging behaviour on the landscape to document important trade‐offs for parents such as predation risk and energy expenditure while feeding offspring.     

Breeding time,food supply and fitness components of Blue Tits Parus caeruleus in Mediterranean habitats

We analysed the relationship between the timing of food availability and within-season variation of both reproductive success and nestling body size of Blue Tits Parus caeruleus in Mediterranean habitats. Synchronization between food supply and reproduction was expected to be positively related to fitness components. We measured deviation from maximum food supply using a parameter that we called “time-lag”, which quantifies the degree of synchronization between the date of maximum food requirements by the nestlings and the date of maximum caterpillar supply in the habitat. This parameter was expected to be related to reproductive success as measured by the number and body-condition of fledglings. The predictions were that time-lag should be negatively correlated with the proportion of nestlings raised to fledging and the size of the fledglings. These predictions have been tested in different types of habitat. The results demonstrate that caterpillar supply during a critical nestling period can have a strong influence on fitness components. As predicted, synchronization with caterpillar supply is positively related to the number and body size of fledglings. Since there is large between-habitat variation in the timing of food supply, the key issue seems to be the adjustment to local patterns of food availability.     

Intraseasonal effects of clutch manipulation on parental provisioning and residual reproductive value of eastern phoebes (Sayornis phoebe)

Summary First clutches of double-brooded eastern phoebes Sayornis phoebe were manipulated (up two eggs, down 2 eggs or no change) to test for intraseasonal reproductive tradeoffs and to test whether size of first brood influenced food delivery rates to nestlings and nestling quality in second broods.Considering all nests from both broods, rate of feeding nestlings increased linearly with brood size but nestling mass per nest decreased with increasing brood size. High nestling weights in small broods may have resulted from parents delivering better quality food, but we did not test this.Among treatment groups in first broods, nestlings from decreased broods weighed more than those in control or increased broods. Treatment did not influence the likelihood that second nests would be attempted after successful first nests nor did it alter the interval between nests. Nestlings of parents that renested weighed more than those of parents that did not, regardless of treatment, suggesting that post-fledging care may preclude renesting. Mass of individual females did not change between broods, regardless of brood size. Clutch sizes of second attempts were not affected by manipulations of first broods but increasing first broods reduced the number of nestlings parents were able to raise to day 11 in their second broods. However, manipulation of first broods did not affect mean nestling mass per nest of nestlings that survived to day 11.In phoebes, parents of small first broods are able to raise nestlings in better condition. We predict that in harsh years, parents of small first broods would be more likely to renest. Parents of enlarged first broods sacrificed quality of offspring in second broods, which seems a reasonable strategy if nestlings from second broods have lower reproductive value.     

Demographic influences of translocated individuals on a resident population of house sparrows

Translocation of individuals from source populations to augment small populations facing risk of extinction is an important conservation tool. Here we examine sex‐specific differences between resident and translocated house sparrows Passer domesticus in reproductive success and survival, and the contribution of translocated individuals to the growth of a local population. We found evidence for assortative mating based on origin revealed by fewer parentages between translocated males and resident females than expected, and the total number of fledglings produced by such pairs was lower. The reproductive success of translocated males was positively related to the size of the throat badge (a sexual ornament), such that only translocated males with a large badge size were as successful as resident males. However, offspring with parents of different origin had higher survival than offspring with parents of the same origin, which suggests hybrid vigour. The contribution of resident and translocated individuals to the stochastic component of the long‐run growth rate of the population was similar; neither the mean individual contributions in fitness nor the demographic variance differed between the two groups. Thus, this experiment shows that translocated individuals may have a similar demographic influence on the growth of local populations as resident individuals. Still, the intermixing of translocated and resident individuals was low, and fitness differed according to origin in relation to individual differences in a sexually selected trait. In addition, hybrid vigour with respect to offspring recruitment seemed to partially decrease the negative fitness consequences of the assortative mating based on origin.     

Providing chicks with extra food lowers male but not female provisioning in the House Sparrow Passer domesticus

We assessed whether adult House Sparrows Passer domesticus adjusted their provisioning in response to an experimental increase in the nutritional condition of their nestlings. When we supplemented chicks directly with additional food, male parents, but not female parents, reduced their provisioning. The results for males, but not females, run contrary to a previous experiment in this species. In addition, female provisioning was positively associated with both brood size and the age of the brood. In contrast, whereas male provisioning was positively associated with brood size, males did not increase provisioning as their chicks grew older. Males, but not females, exhibited repeatability in their provisioning. Food supplementation had a larger positive effect upon nestling survival in smaller broods than in larger broods. Overall, there appear to be fundamental differences between males and females in how decisions regarding the level of parental investment in the current brood are made.     

The importance of food quantity and quality for reproductive performance in alpine water pipits (Anthus spinoletta)

Studies relating reproduction to food availability are usually restricted to food quantity, but ignore food quality and the effects of habitat structure on obtaining the food. This is particularly true for insectivorous birds. In this study we relate measures of reproductive success, time of reproduction and nestling size of water pipits (Anthus spinoletta) to biomass, taxonomic composition and nutritional content of available food, and to vegetation structure and distance to feeding sites. Clutch size was positively correlated with the proportion of grass at the feeding sites, which facilitates foraging. This suggests that water pipits adapt their clutch size to environmental conditions. Also, pipits started breeding earlier and produced more fledglings when abundant food and a large proportion of grass were available, probably because these conditions allow the birds to gain more energy in less time. The number of fledglings was positively correlated with the energy content of available food. No significant relationships were found between feeding conditions and nestling size or the time that nestlings took to fledge. This suggests that water pipits do not invest more in individual nestlings when food conditions are favourable but rather start breeding earlier and produce more young. Taxonomic composition and nutritional content of prey were not correlated with any of the reproductive parameters, indicating that profitability rather than quality of food affects reproductive success. Received: 31 May 1996/Accepted: 12 August 1996     

Importance of feeding ecology to the reproductive success of Blackbirds Turdus merula nesting in rural habitats

The feeding ecology of Blackbirds Turdus merula breeding in contiguous woodland and farmland habitats was studied over three years. The aim of the study was to investigate how reproductive success was influenced by nestling diet and the provisioning rates of parents feeding nestlings. Parental provisioning rates increased with brood size, and consequently individual nestlings were no lighter in larger broods. None of the environmental factors measured had strong effects on parental provisioning rate. The nestling diet was dominated by caterpillars and earthworms, the former occurring in a short period in the middle of the breeding season. The availability of earthworms was higher in woodland and was dependent on rainfall in farmland. Nestling mass and provisioning rates were marginally higher under predominantly earthworm diets. Nestling mass increased with rainfall in farmland only, and was higher in farmland than in woodland or woodland-edge, although it is doubtful whether this result is of any significance for fledgling survival. Overall, Blackbirds were able to provision their nestlings adequately throughout the breeding season across a range of conditions. There was no evidence to suggest that reproductive success was constrained by aspects of feeding ecology within the natural range of brood size.     

Siblicide, starvation and nestling growth in the laughing kookaburra

I examined the growth of surviving nestlings in broods of the cooperatively breeding laughing kookaburra Dacelo novaeguineae , which has complex patterns of brood reduction. Laughing kookaburras usually lay three eggs that hatch asynchronously. Brood reduction occurs in nearly half of all broods and always affects the youngest nestling. In most cases, the youngest nestling is killed within a few days of hatching by aggressive attacks from its older siblings. In a smaller proportion of nests, the youngest nestling dies from starvation, rather than physical attack, much later in the nestling period when nestling growth rates and adult feeding rates peak (about 20 days post-hatching). These mechanistically and temporally distinct episodes of brood reduction were associated with very different patterns of growth in the senior nestlings. Seniors that killed their youngest sibling reached higher asymptotic weights than seniors that did not commit siblicide. In contrast, if the youngest nestling was not killed by its older siblings, but later starved to death, the surviving seniors were skeletally smaller and had retarded feather development compared to seniors from other broods. These differences in nestling growth may have longer-term fitness consequences, because kookaburra fledging weight is positively associated with both juvenile survival and successful recruitment into the breeding population. Therefore, although parents of broods without mortality produce the highest number of fledglings and also the highest number of independent juveniles, if parents are unable to raise a full brood, early siblicide may represent the best brood reduction option. Early siblicide is at least associated with high quality young that have enhanced survival and recruitment prospects. In contrast, the poor growth of seniors in broods where the youngest nestling starved suggests that parents overestimated the size of the brood they could provision.     

Feeding decisions of eastern bluebirds are situationally influenced by fledgling plumage color

The relative amount of resources that avian parents provide to individual offspring within a brood represents a strategy that can have large effects on reproductive success. We tested whether parental feeding decisions of eastern bluebirds Sialia sialis are influenced by offspring plumage color by presenting pairs of differently colored fledglings side by side and observing how they were provisioned by parents. After a control period, we manipulated blue plumage color so that one sibling in each trial became relatively dark and one became relatively bright. During neither the control nor the experimental periods did either parent consistently feed naturally brighter or experimentally brightened sons more than drab sons. Under specific circumstances, however, both parents directed a higher proportion of their feeding attempts to more brightly colored sons. Paternal feeding attempts to brighter offspring during both the control and experimental periods increased in relation to the brightness of these fledglings relative to their brothers. Maternal feeding decision, on the other hand, were influenced by numerous variables during control and experimental periods including the date of the trial, the difference in mass between fledglings, the feeding behavior of fathers during the trial, the relative investment by fathers during the nestling stage, and the amount of UV chroma in fledgling plumage. Taken together, these results suggest that equal provisioning of offspring is the strategy most commonly adopted by eastern bluebirds but more brightly colored offspring will be fed preferentially when resources for offspring are limited.     

Reproductive success and nestling diet in the Lesser Spotted Woodpecker ( Picoides minor ): the early bird gets the caterpillar

Populations of Lesser Spotted Woodpecker (Picoides minor) are decreasing in size, necessitating management strategies. However, data on the reproductive biology of this species are scarce. The aim of this study was to investigate the influence of timing of breeding and nestling diet on the reproductive success of the Lesser Spotted Woodpecker and thereby contribute to an understanding of the determinants influencing its reproductive success. During 6 study years between 1996–2003, we investigated various variables of reproductive success in a Lesser Spotted Woodpecker population in the Taunus low mountain range, Germany. We observed nestling feeding at breeding holes to assess the composition of nestling diet and how it changes during the season. Clutch size, number of fledglings as well as body mass of the nestlings declined with the start of egg laying. Pairs composed of individuals that had breed together in previous years started egg laying earlier than newly established pairs. Nestling diet consisted mainly of aphids, caterpillars, craneflies and wood-living larvae. The composition changed considerably within the breeding season in both early and late broods. However, early broods were provided with caterpillars more frequently, whereas late broods received wood-dwelling larvae more often. Our results suggest that Lesser Spotted Woodpeckers time their breeding so as to coincide with the early nestling stage with the highest availability of caterpillars, since this is the time of highest energy demand. Moreover, food availability declines during the breeding season, and the decline in reproductive performance seems to be an effect of this development.     

Brood provisioning rates and fledgling behavior of Cordilleran Flycatchers in southwestern Colorado

The behavior of young songbirds after fledging is one of the least understood phases of the breeding cycle, although parental provisioning rates and movement of fledglings are key to understanding life history evolution. We studied Cordilleran Flycatchers (Empidonax occidentalis) at two sites in southwestern Colorado, USA, from 2012 to 2017. We banded and sexed breeding adults to determine the relative contributions of males and females to nestling and fledgling care, and attached radio‐transmitters to nestlings to facilitate observations of brood behavior after fledging. Females made 60% and 78% of total observed feedings of nestlings and fledglings, respectively. Parental provisioning rates increased with nestling age, and per‐nestling provisioning rates increased with brood size. Parental provisioning rates declined just before fledging, then increased just after fledging. Fledging times of individuals in broods were asynchronous and concentrated during the late afternoon and early evening. Males stopped caring for fledglings before females even though this species is single‐brooded, with some late‐season broods being abandoned by males. Broods spent the first three weeks after fledging within 400 m of nests, after which they began to disperse. Most aspects of the breeding biology of Cordilleran Flycatchers in our study, including the duration of nestling and fledging periods, female‐dominated provisioning, and movement patterns of fledglings, were similar to those of other Empidonax species. However, the times when young fledged were not concentrated in the morning as reported in most other songbirds, and this result warrants additional study of the timing of fledging in ecologically and taxonomically similar species. The increased per‐nestling provisioning rate with increasing brood size was unexpected, and additional study is needed to determine if this increase results from a trade‐off between adult annual survival and productivity favoring increased provisioning of young in larger broods, or from the existence of high‐quality individuals where larger clutches and higher provisioning rates are linked.     

Parental care trade‐offs in the inter‐brood phase in Barn Swallows Hirundo rustica

In seasonal environments with limited time and energy resources, double‐brooded birds face trade‐offs in the timing of their two reproductive attempts and in the effort allocated to the first and the second broods. In the Barn Swallow Hirundo rustica a long care period for the first brood enhances the survival of first‐brood chicks, but also delays the start of the second brood, which in turn reduces the survival prospects of second‐brood chicks. Probably as a response to this trade‐off, double‐brooded Barn Swallows reduce the period of post‐fledging care for first‐brood fledglings. By radiotracking whole families, we investigated the determinants of this behaviour and its consequences for the survival of the first‐brood fledglings. The end of the females’ investment in post‐fledging care of the first brood was related to the beginning of egg synthesis for the second clutch. With the start of egg synthesis, females significantly reduced provisioning rates to the first‐brood fledglings to less than one‐fifth of the previous rates, while the proportion of time they spent foraging remained high. Assuming that the females’ foraging success was constant, we conclude that their energy income was allocated to egg production rather than fledgling provision. Males did not compensate for the females’ reduced feeding rates. Thus the start of egg production for the second clutch had a marked effect on the quantity of food received by first‐brood fledglings. In parallel with the changes in parental behaviour and provisioning rates, we observed a marked drop in the daily survival rate of first‐brood chicks. These results support the hypothesis that females face a strong trade‐off in the allocation of energy to subsequent broods. Energy allocation to a second clutch involves a cost in terms of reduced provisioning, and as a result the survival of first‐brood chicks is compromised. This is probably outweighed by the improved success of an early second brood.     

Brood parasitism increases provisioning rate, and reduces offspring recruitment and adult return rates, in a cowbird host

Interspecific brood parasitism in birds presents a special problem for the host because the parasitic offspring exploit their foster parents, causing them to invest more energy in their current reproductive effort. Nestling brown-headed cowbirds (Molothrus ater) are a burden to relatively small hosts and may reduce fledgling quality and adult survival. We documented food-provisioning rates of one small host, the prothonotary warbler (Protonotaria citrea), at broods that were similar in age (containing nestlings 8–9 days old), but that varied in composition (number of warbler and cowbird nestlings) and mass, and measured the effect of brood parasitism on offspring recruitment and adult returns in the host. The rate of food provisioning increased with brood mass, and males and females contributed equally to feeding nestlings. Controlling for brood mass, the provisioning rate was higher for nests with cowbirds than those without. Recruitment of warbler fledglings from unparasitized nests was 1.6 and 3.7 times higher than that of fledglings from nests containing one or two cowbirds, respectively. Returns of double-brooded adult male and female warblers decreased with an increase in the number of cowbirds raised, but the decrease was more pronounced in males. Reduced returns of warbler adults and recruitment of warbler fledglings with increased cowbird parasitism was likely a result of reduced survival. Cowbird parasitism increased the warblers’ investment in current reproductive effort, while exerting additional costs to current reproduction and residual reproductive value. Our study provides the strongest evidence to date for negative effects of cowbird parasitism on recruitment of host fledglings and survival of host adults.     

Correlates of cell-mediated immunity in nestling house sparrows

Cell-mediated immunity is an important vertebrate defense against pathogens, but components of this response may vary in quality. Such variation could arise through the effects of ecology on optimal immunocompetence. We used injections of phytohaemagglutinin (PHA) to measure the factors influencing T-cell proliferation in nestling house sparrows (Passer domesticus). Bivariate analyses revealed positive associations with nestling mass and size, but no effect of ectoparasites. The response to PHA was, however, strongly affected by brood identity. A mixed model with brood identity as a random factor and nestling mass, size, number of ectoparasites, parental feeding rate, clutch size, brood size at hatching, and date uncovered significant positive correlations between PHA response and both nestling mass and the brood size at hatching. Because many of these variables are related hierarchically, we used path analysis to explore the relationships in more detail. We found that a nestling immune response was affected by several indirect paths. Brood size at hatch had both positive and negative paths, and date in the season had several indirect negative effects through its effect on brood size and nestling mass. The approach used and the results obtained offer some new ideas for incorporating immune responses into life history theory.     

Prudent investment in rearing nestlings in bull-headed shrikes

I investigated seasonal changes in the relationships between brood size, body mass of nestlings and body mass of parents of the bull-headed shrike, Lanius bucephalus, in Ishikari, northern Japan. When the broods were 12days old, the body mass of the heaviest nestling in a brood did not differ among brood sizes, or throughout the season. However, the body mass of the lightest nestlings in a brood was different among brood sizes. The body mass of the lightest nestling in five- and six-nestling broods decreased throughout the season. The lightest nestling in four-nestling broods, and the lightest and the second lightest nestlings in five-nestling broods, were significantly lighter than the heaviest nestling in broods of this size. It is likely that pairs with six nestlings at 12days old can feed at least five of these nestlings enough to ensure their survival . The standardized body mass of parents (SBM), which was defined as the body mass divided by the length of the tarsus, did not differ among brood sizes, or throughout the season. It is possible that the relationship between the constancy of the SBM and the seasonal decline in the body mass of nestlings indicates that bull-headed shrikes have a limit to their parental efforts.     

Does cowbird parasitism increase predation risk to American redstart nests?

Brood parasitism and nest predation are major causes of reproductive failure for many bird species nesting in fragmented landscapes. While brood parasites and predators may act independently, they could also interact if brood parasites increase the likelihood that predators detect nests. In this study, we examined the interaction between cowbird parasitism and nest predation in a 10 year study on 466 American redstart Setophaga ruticilla nests in central Alberta, Canada. We used advanced nest survival models to examine the support for three mechanisms that might lead to a positive correlation between brood parasitism and nest predation: 1) the presence of a cowbird nestling might increase the detection of the nest by predators, 2) nests with lower cover are more likely to be detected by both cowbirds and predators, and 3) cowbirds and predators may co-occur in landscapes of similar structure. Twelve percent of nests were parasitized and those nests had a 16–19% higher rate of failure due to predators compared to unparasitized nests. Daily nest predation rates increased during the nestling stage for both groups, but more strongly for parasitized nests. Loud begging by the cowbird nestling and/or higher parental feeding rates for the cowbird may have increased nest detectability to predators. Brood parasitism and nest predation were also positively related to forest cover, indicating landscape level effects were influential. Most nest predators were forest species and we suspect cowbirds responded positively to forest cover because of the increased abundance of songbird hosts. Nest-site features had less of an impact on nest predation or brood parasitism, although nests with higher overhead cover were less susceptible to predators. Our study shows how multiple mechanisms, particularly the behavioral effects of the brood parasite nestling and landscape structure, can lead to a positive relationship between nest predation and brood parasitism.