Nestling Growth Rates Research Articles

Shared parental care is costly for nestlings of common cuckoos and their great reed warbler hosts

Obligate avian brood parasitism typically involves one of 2 strategies: parasite chicks are either 1) virulent and evict all other eggs and nest mates to be raised alone or 2) more tolerant and share foster parental care with host chicks for some or the entirety of the nestling period. We studied the consequences of experimentally forced mixed broods of age-matched one common cuckoo (Cuculus canorus) and 2 great reed warbler (Acrocephalus arundinaceus) chicks. In these broods, both cuckoo and host chicks grew slower than did either individual cuckoos or great reed warblers in broods of 1 parasite or 3 host chicks, respectively. Video records showed that in mixed broods, cuckoo chicks received feedings less frequently than the 33% predicted by chance at 4 days of age but parental food allocations increased to chance levels at 8 days of age. The consistent patterns of lower growth rates arose even though chicks in broods of 1 parasite and 2 hosts received the largest prey items per feeding. In addition, several other measures of parental provisioning also did not predict species and brood-specific differences in nestling growth rates across the different treatments. However, variation in begging displays and its specific costs on host and parasite chicks in the different nest treatments were not quantified in this study. We conclude that young of nest mate-evictor common cuckoos benefit from the sole occupancy of host nests in part owing to an initial competitive disadvantage for parental care in broods with age-matched great reed warbler chicks.

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

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

Melanin-based colorations signal strategies to cope with poor and rich environments

One hypothesis for the maintenance of genetic variation states that alternative genotypes are adapted to different environmental conditions (i.e., genotype-by-environment interaction G×E) that vary in space and time. Although G×E has been demonstrated for morphological traits, little evidence has been given whether these G×E are associated with traits used as signal in mate choice. In three wild bird species, we investigated whether the degree of melanin-based coloration, a heritable trait, covaries with nestling growth rate in rich and poor environments. Variation in the degree of reddish-brown phaeomelanism is pronounced in the barn owl (Tyto alba) and tawny owl (Strix aluco), and variation in black eumelanism in the barn owl and Alpine swift (Apus melba). Melanin-based coloration has been shown to be a criterion in mate choice in the barn owl. We cross-fostered hatchlings to test whether nestlings sired by parents displaying melanin-based colorations to different extent exhibit alternative growth trajectories when raised by foster parents in poor (experimentally enlarged broods) and rich (experimentally reduced broods) environments. With respect to phaeomelanism, barn owl and tawny owl offspring sired by redder parents grew more rapidly in body mass only in experimentally reduced broods. With respect to eumelanism, Alpine swift offspring of darker fathers grew their wings more rapidly only in experimentally enlarged broods, a difference that was not detected in reduced broods. These interactions between parental melanism and offspring growth rate indicate that individuals display substantial plasticity in response to the rearing environment which is associated with the degree of melanism: at least with respect to nestling growth, phaeomelanic and eumelanic individuals are best adapted to rich and poor environments, respectively. It now remains to be investigated why eumelanism and phaeomelanism have a different signaling function and what the lifelong consequences of these melanism-dependent allocation strategies are. This is important to fully appraise the role played by environmental heterogeneity in maintaining variation in the degree of melanin-based coloration.

Does testosterone mediate the trade-off between nestling begging and growth in the canary ( Serinus canaria)?

Nestling birds solicit food from their parents with vigorous begging displays, involving posturing, jostling and calling. In some species, such as canaries, begging is especially costly because it causes a trade off against nestling growth. Fitness costs of begging like this are predicted by evolutionary theory because they function to resolve conflicts of interest within the family over the provision of parental investment. However, the mechanism that links these costs with nestling behaviour remains unclear. In the present study, we determine if the relationships between nestling androgen levels, nestling begging intensities and nestling growth rates are consistent with the hypothesis that testosterone is responsible for the trade-off between begging and growth. We test this idea with a correlational study, using fecal androgens as a non-invasive method for assaying nestling androgen levels. Our results show that fecal androgen levels are positively correlated with nestling begging intensity, and reveal marked family differences in each trait. Furthermore, changes in fecal androgen levels between 5 and 8 days after hatching are positively associated with changes in nestling begging intensity, and negatively associated with nestling growth during this time. Although these correlational results support our predictions, we suggest that that experimental manipulations are now required to test the direct or indirect role of testosterone in mediating the trade-off between begging and growth.

Site- and sex-level differences in adult feeding behaviour and its consequences to offspring quality in tree swallows (Tachycineta bicolor) following brood-size manipulation

In species with biparental care, males tend to invest less in offspring than do females, likely because of differences in the costs and benefits associated with parental effort. Here I test for sex differences in the response of tree swallows ( Tachycineta bicolor (Vieillot, 1808)) to a brood-size manipulation at two locations differing in food resources, Alaska and New York. I tested sex and habitat differences in how swallows responded to changes in offspring demand. At both sites, both sexes increased effort when feeding enlarged broods, although Alaskan males increased feeding less than Alaskan females. Males decreased feeding effort more to reduced broods than females, but only in Alaska. Food abundance was higher in Alaska than in New York, and Alaskan tree swallows made more feeding visits than New York tree swallows. In New York, food availability did not predict feeding rate and there was no sex difference in the response to brood manipulation. In both sites, male feeding effort was linked with nestling residual body mass, while female feeding effort was correlated with nestling growth rate. This study demonstrates that male tree swallows differ from females by being the first to reduce feeding effort under certain conditions and that male and female feeding rate affects offspring quality differently.

Increased susceptibility to oxidative damage as a cost of accelerated somatic growth in zebra finches

1. Most animals do not grow at their maximal rate. This might appear puzzling because the early attainment of a large body size incurs several selective benefits, such as reduced risk of predation and earlier reproductive output. Several hypotheses have been suggested to explain this paradox. Among them, the cost due to high levels of oxidative stress, as the consequence of sustained metabolic activity during growth, has been put forward. 2. In this study, we wished to assess this cost in captive zebra finches (Taeniopygia guttata). In order to manipulate access to food and consequently early growth, hatchlings were randomly assigned to reduced or enlarged broods. Even though nestlings raised in enlarged broods were smaller when 20-days-old compared to nestlings raised in reduced broods, they grew faster during the following 20-day period. 3. When 60-days-old, we measured the resistance of red blood cells against a free radical attack and correlated these values to nestling growth rate. In agreement with the prediction, we found that red blood cell resistance to free radicals was negatively correlated with growth rate, nestlings that grew faster being those with the weakest capacity to resist a free radical attack. These results support the hypothesis that oxidative damage might constrain growth rate. Furthermore, the fact that the relationship was only established during the period of accelerated growth after the initial delay also suggests that compensatory growth can negatively affect the individual resistance to oxidative damage.

Effects of Distance to Edge and Edge Type on Nestling Growth and Nest Survival in the Wood Thrush

Abstract Many studies investigating edge effects on forest-nesting birds have focused on nest success. Fewer have examined edge effects on other components of fitness. Nestling growth rates have been positively correlated with food availability, which may differ at edges compared to the forest interior. However, previous work has not examined growth as a function of distance to edge and edge type. We investigated whether edge type, distance to edge, hatching date, brood size, brood parasitism, year, and site influenced growth rates of wings, tarsi, and mass of nestling Wood Thrushes (Hylocichla mustelina) using regression tree analyses. Simultaneously, we examined edge effects on daily nest survival. We conducted our study in southwestern Michigan in 2002 and 2003. We located 175 nests and measured nestlings from 61 nests. Contrary to expectations, tarsal and mass growth rates were more rapid near edges than in the forest interior and tarsal growth rates were more rapid near gradual edge types than near abrupt edge types. Wing chord growth rates were more rapid in the forest interior than at edges. Mass and tarsal growth rates were more rapid later in the breeding season, when brood sizes were smaller. We found no effect of edge type or distance to edge on nest survival. The influence of edge on nestling growth rate but not nest survival suggests nestling growth rates may indicate edge-interior and edge type differences in habitat quality, even when high regional fragmentation levels overwhelm potential edge-interior differences in nest survival.

Androgen-dependent maternal effects on offspring fitness in zebra finches

There is accumulating evidence that maternal hormones may play a role in offspring sex adjustment, but little is known about the costs of such hormone-mediated mechanisms. Recent studies have reported sex-specific effects of hormones on offspring viability. Specifically, we previously found that elevating the plasma androgen level in mothers results in a male-biased offspring primary sex ratio, but it affects the viability of sons negatively and daughters positively in zebra finches (Taeniopygia guttata; Rutkowska and Cichon, Anim Behav, 71:1283–1288, 2006). In this study, we studied further fitness consequences of exposure to elevated yolk androgen levels in zebra finches. We measured growth rate and cellular immune response of nestlings that hatched from eggs laid by females injected with testosterone during egg laying and nestlings of unaffected control females. We found that sons of testosterone-treated females grew slower in comparison to sons of control females. The significant interaction between experimental group and offspring sex indicates that sons of testosterone-treated mothers suffered impaired immune responsiveness while daughters seemed to benefit from elevated androgen level in terms of enhanced immune responsiveness. We found no effects of androgens on offspring performance at adulthood—neither fecundity of females nor attractiveness of males was affected. We conclude that the benefits of biasing sex ratio towards males by increasing androgen level in the yolk may be limited due to negative effects on male offspring performance early in life.

EFFECTS OF DISTANCE TO EDGE AND EDGE TYPE ON NESTLING GROWTH AND NEST SURVIVAL IN THE WOOD THRUSH

Many studies investigating edge effects on forest-nesting birds have focused on nest success. Fewer have examined edge effects on other components of fitness. Nestling growth rates have been positively correlated with food availability, which may differ at edges compared to the forest interior. However, previous work has not examined growth as a function of distance to edge and edge type. We investigated whether edge type, distance to edge, hatching date, brood size, brood parasitism, year, and site influenced growth rates of wings, tarsi, and mass of nestling Wood Thrushes (Hylocichla mustelina) using regression tree analyses. Simultaneously, we examined edge effects on daily nest survival. We conducted our study in southwestern Michigan in 2002 and 2003. We located 175 nests and measured nestlings from 61 nests. Contrary to expectations, tarsal and mass growth rates were more rapid near edges than in the forest interior and tarsal growth rates were more rapid near gradual edge types than near abrupt edge types. Wing chord growth rates were more rapid in the forest interior than at edges. Mass and tarsal growth rates were more rapid later in the breeding season, when brood sizes were smaller. We found no effect of edge type or distance to edge on nest survival. The influence of edge on nestling growth rate but not nest survival suggests nestling growth rates may indicate edge-interior and edge type differences in habitat quality, even when high regional fragmentation levels overwhelm potential edge-interior differences in nest survival.

Delayed behavioral effects of postnatal exposure to corticosterone in the zebra finch ( Taeniopygia guttata)

Early developmental conditions can significantly influence the growth and survival of many animal species. We studied the consequences of exposure to corticosterone (CORT), a stress hormone, during the nestling stage on two behavioral traits (neophobia, social dominance) measured when the birds had reached independence. Nestling zebra finches ( Taeniopygia guttata) were exposed twice daily to exogenous CORT via oral administration for a 12-day period up until fledging. Experimental CORT administration depressed nestling growth rates, confirming results previously obtained in this species. Our data on neophobic behavior revealed a significant interaction between sex and treatment, with CORT-dosed males showing reduced latencies to approach a novel object, while there was little effect of corticosterone treatment on female neophobia. There was no significant effect of age (30 or 50 days), however, there was a non-significant trend towards an interaction between treatment and age, with neophobia increasing with age in the CORT-dosed birds, but decreasing in controls. At 50 days of age previous exposure to corticosterone resulted in reduced success in competitions for a non-food-based resource (a perch) in both sexes. There were no effects of brood size on any behavioral traits measured here, but this may be due to the small range in brood size used. Our results show that elevated levels of stress hormones during postnatal development can have significant effects on important behavioral traits, i.e., neophobia and dominance. Moreover, they confirm the importance of rearing conditions in shaping adult phenotypes.

Avian growth and development rates and age‐specific mortality: the roles of nest predation and adult mortality

Previous studies have shown that avian growth and development covary with juvenile mortality. Juveniles of birds under strong nest predation pressure grow rapidly, have short incubation and nestling periods, and leave the nest at low body mass. Life-history theory predicts that parental investment increases with adult mortality rate. Thus, developmental traits that depend on the parental effort exerted (pre- and postnatal growth rate) should scale positively with adult mortality, in contrast to those that do not have a direct relationship with parental investment (timing of developmental events, e.g. nest leaving). I tested this prediction on a sample of 84 North American songbirds. Nestling growth rate scaled positively and incubation period duration negatively with annual adult mortality rates even when controlled for nest predation and other covariates, including phylogeny. On the contrary, neither the duration of the nestling period nor body mass at fledging showed any relationship. Proximate mechanisms generating the relationship of pre- and postnatal growth rates to adult mortality may include increased feeding, nest attentiveness during incubation and/or allocation of hormones, and deserve further attention.

Growth and Plumage Development of Nestling Long-Eared Owls

Abstract Abstract We report the growth rates and plumage development of Long-eared Owls (Asio otus) in two developmental stages from hatching to fledging, the nestling period (1–22 days) and prefledging period (23–37 days). We used the logistic growth equation to model nestling mass gain. Mass gain was most rapid during the nestling period. Average mass gain per day was highest between seven and 13 days of age. At 16 days of age nestlings displayed defined facial disks and complete second down. Nestlings left the nest at 22 days of age. Mass gain slowed considerably after nest departure and continued to slow up to fledging. Appearance of defensive and concealment behaviors occurred nearly simultaneously with departure from the nest. To our knowledge this is the first detailed report of nestling growth rates and plumage development for Long-eared Owls in North America.

Geographic Variation in the Trade-Off Between Nestling Growth Rate and Body Condition in the Tree Swallow

AbstractNestlings can exhibit considerable variation in developmental patterns both within and among locations due to differences in environmental conditions and parental investment. I investigated trade-offs between nestling growth rate and residual body mass (body condition) at three locations across the range of the Tree Swallow (Tachycineta bicolor). Nestlings at the northern extreme of the range in Alaska had slower growth rates, lower body mass, and higher residual body mass than nestlings in New York and Tennessee. High insect availability was correlated with increased growth rates of nestlings in New York and Tennessee, but not in Alaska. Conversely, nestlings in Alaska showed increased residual body mass with high insect availability, but nestlings in New York and Tennessee did not. The trade-off between growth rate and residual body mass varied among sites, with fast-growing nestlings in Tennessee maintaining a higher residual body mass than those in Alaska. These results suggest that factors affecting offspring growth and condition vary among sites, leading to geographical differences in offspring development trajectories.

GROWTH STRATEGIES OF PASSERINE BIRDS ARE RELATED TO BROOD PARASITISM BY THE BROWN-HEADED COWBIRD (MOLOTHRUS ATER)

Sibling competition was proposed as an important selective agent in the evolution of growth and development. Brood parasitism by the brown-headed cowbird (Molothrus ater) intensifies sibling competition in the nests of its hosts by increasing host chick mortality and exposing them to a genetically unrelated nestmate. Intranest sibling competition for resources supplied by parents is size dependent. Thus, it should select for high development rates and short nestling periods, which would alleviate negative impacts of brood parasitic chicks on host young. I tested these predictions on 134 North American passerines by comparative analyses. After controlling for covariates and phylogeny, I showed that high parasitism rate was associated with higher nestling growth rate, lower mass at fledging, and shorter nestling periods. These effects were most pronounced in species in which sibling competition is most intense (i.e., weighing over about 30 g). When species were categorized as nonhosts versus old hosts (parasitized for thousands of years) versus new hosts (parasitized the last 100-200 years), there was a clear effect of this parasitism category on growth strategies. Nestling growth rate was the most evolutionarily flexible trait, followed by mass at fledging and nestling period duration. Adjustments during incubation (incubation period length, egg volume) were less pronounced and generally disappeared after controlling for phylogeny. I show that sibling competition caused by brood parasites can have strong effects on the evolution of host growth strategies and that the evolution of developmental traits can take place very rapidly. Human alteration of habitats causing spread of brood parasites to new areas thus cascades into affecting the evolution of life-history traits in host species.

Reduced nestling growth of East African StonechatsSaxicola torquata axillarisin the presence of a predator

We investigated nestling growth of tropical East African StonechatsSaxicola torquata axillaristo evaluate the effects of nest predation, predator presence and food availability. We provided some Stonechat pairs with supplemental food, while others in a similar habitat served as a control. Concomitantly, we assessed the presence of Fiscal ShrikesLanius collarisin supplemental fed and unsupplemented territories. Fiscal Shrikes prey on adult Stonechats and nestlings. We found that nestling growth was considerably reduced in Stonechat pairs that shared their territory with a Shrike. This effect was greater in nestlings of pairs that did not receive supplemental food. The reduction in nestling growth rates was significantly correlated with a reduced rate of visiting by the parents. Behavioural observations further suggested that parents reduced their feeding visits to the nest presumably to minimize their own predation risk, rather than predation risk of their brood. Our experiments show that the lower reproductive investment in tropical Stonechats can be attributed to risk‐sensitive behaviour of the parents, especially when food is in limited supply.

Increased energy expenditure by a seabird in response to higher food abundance

Variability in forage fish abundance strongly affects seabird behavior and reproductive success, although details of this relationship are unclear. During 1997 and 1998, we measured (1) daily energy expenditure (DEE) of 80 parent black-legged kittiwakes Rissa tridactyla at 2 colonies in Prince William Sound, Alaska (North Icy Bay and Shoup Bay), (2) abundance of surface-schooling forage fishes within the foraging range of each colony, and (3) diet composition, energy delivery rates to nestlings, and reproductive success of kittiwakes at these same colonies. Female DEE was highest at North Icy Bay in 1998, while male DEE did not differ by colony year. Abundances of Pacific herring Clupea pallasi and sand lance Ammodytes hexapterus were highest near North Icy Bay in 1998 and nearly equal in density, although Age 1+ herring comprised the majority of the diet there. Energy delivery rates to nestlings, nestling growth rates, and productivity were also highest at North Icy Bay in 1998. We suggest that female kittiwakes responded to the increased abundance of Age 1+ herring near North Icy Bay in 1998 by increasing their DEE, which in turn positively affected reproductive success. Given that adult kittiwakes have been shown to suffer decreased survival as a response to increased energy expenditure during brood rearing, the positive correlation we observed between increased abundance of a high quality food source, parental effort, and productivity is consistent with maximizing lifetime reproductive success. The lack of a response in male DEE suggests that brood-rearing roles in kittiwakes differ between genders.

GEOGRAPHIC VARIATION IN THE TRADE-OFF BETWEEN NESTLING GROWTH RATE AND BODY CONDITION IN THE TREE SWALLOW

Abstract Nestlings can exhibit considerable variation in developmental patterns both within and among locations due to differences in environmental conditions and parental investment. I investigated trade-offs between nestling growth rate and residual body mass (body condition) at three locations across the range of the Tree Swallow (Tachycineta bicolor). Nestlings at the northern extreme of the range in Alaska had slower growth rates, lower body mass, and higher residual body mass than nestlings in New York and Tennessee. High insect availability was correlated with increased growth rates of nestlings in New York and Tennessee, but not in Alaska. Conversely, nestlings in Alaska showed increased residual body mass with high insect availability, but nestlings in New York and Tennessee did not. The trade-off between growth rate and residual body mass varied among sites, with fast-growing nestlings in Tennessee maintaining a higher residual body mass than those in Alaska. These results suggest that factors a...

GROWTH STRATEGIES OF PASSERINE BIRDS ARE RELATED TO BROOD PARASITISM BY THE BROWN-HEADED COWBIRD (MOLOTHRUS ATER)

Sibling competition was proposed as an important selective agent in the evolution of growth and development. Brood parasitism by the brown-headed cowbird (Molothrus ater) intensifies sibling competition in the nests of its hosts by increasing host chick mortality and exposing them to a genetically unrelated nestmate. Intranest sibling competition for resources supplied by parents is size dependent. Thus, it should select for high development rates and short nestling periods, which would alleviate negative impacts of brood parasitic chicks on host young. I tested these predictions on 134 North American passerines by comparative analyses. After controlling for covariates and phylogeny, I showed that high parasitism rate was associated with higher nestling growth rate, lower mass at fledging, and shorter nestling periods. These effects were most pronounced in species in which sibling competition is most intense (i.e., weighing over about 30 g). When species were categorized as nonhosts versus old hosts (parasitized for thousands of years) versus new hosts (parasitized the last 100–200 years), there was a clear effect of this parasitism category on growth strategies. Nestling growth rate was the most evolutionarily flexible trait, followed by mass at fledging and nestling period duration. Adjustments during incubation (incubation period length, egg volume) were less pronounced and generally disappeared after controlling for phylogeny. I show that sibling competition caused by brood parasites can have strong effects on the evolution of host growth strategies and that the evolution of developmental traits can take place very rapidly. Human alteration of habitats causing spread of brood parasites to new areas thus cascades into affecting the evolution of life-history traits in host species.

GROWTH AND PLUMAGE DEVELOPMENT OF NESTLING LONG-EARED OWLS

Abstract ABSTRACT We report the growth rates and plumage development of Long-eared Owls (Asio otus) in two developmental stages from hatching to fledging, the nestling period (1–22 days) and prefledging period (23–37 days). We used the logistic growth equation to model nestling mass gain. Mass gain was most rapid during the nestling period. Average mass gain per day was highest between seven and 13 days of age. At 16 days of age nestlings displayed defined facial disks and complete second down. Nestlings left the nest at 22 days of age. Mass gain slowed considerably after nest departure and continued to slow up to fledging. Appearance of defensive and concealment behaviors occurred nearly simultaneously with departure from the nest. To our knowledge this is the first detailed report of nestling growth rates and plumage development for Long-eared Owls in North America.

Effects of extra food on nestling growth and survival in red-backed voles (Clethrionomys gapperi)

Nestling mortality is typically high among small mammals, with food often considered an important limiting factor because of the increased energetic costs females incur during lactation. We provided female red-backed voles, Clethrionomys gapperi (Vigors, 1830), with sunflower seeds during lactation to test the hypothesis that food supply influences nestling growth and survival. Food supplementation did not increase nestling survival, but newly emerged young from fed mothers were significantly heavier than those from unfed mothers. High mortality/dispersal of postemergent young, particularly females, was associated with high densities of breeding females. We suggest that any survival advantage incurred from higher nestling growth rates may be mitigated by density-dependent mortality/ dispersal postemergence.