Environmental influences on the evolution of growth and developmental rates in passerines

Abstract.— The reasons why growth and developmental rates vary widely among species have remained unclear. Previous examinations of possible environmental influences on growth rates of birds yielded few correlations, leading to suggestions that young may be growing at maximum rates allowed within physiological constraints. However, estimations of growth rates can be confounded by variation in relative developmental stage at fledging. Here, we re-estimate growth rates to control for developmental stage. We used these data to examine the potential covariation of growth and development with environmental variation across a sample of 115 North American passerines. Contrary to previous results, we found that growth rates of altricial nestlings were strongly positively correlated to daily nest predation rates, even after controlling for adult body mass and phylogeny. In addition, nestlings of species under stronger predation pressure remained in the nest for a shorter period, and they left the nest at lower body mass relative to adult body mass. Thus, nestlings both grew faster and left the nest at an earlier developmental stage in species with higher risk of predation. Growth patterns were also related to food, clutch size, and latitude. These results support a view that growth and developmental rates of altricial nestlings are strongly influenced by the environmental conditions experienced by species, and they generally lend support to an adaptive view of interspecific variation in growth and developmental rates.     

Food habits and the survival and growth of nestlings in two sympatric kites (Milvus milvus and Milvus migrans)

A comparative study of the Red Kite and Black Kite was conducted in the Doana National Park in southwestern Spain. Although the diets of both kites were similar, the Red Kite captured significantly more prey items from scrub habitats close to nesting areas than from marsh habitats, which were further away. Habitat differences in prey abundance and distance from breeding areas suggest that the Red Kite depended on a less abundant but more stable food supply. Nestling mortality from starvation was highest for the last hatched nestlings in both species. The growth rate of Black Kite nestlings was markedly higher than that of Red Kite nestlings. Moreover, growth rates increased with clutch size in the Red Kite, whereas they were constant in the Black Kite. Differences between species in reproductive traits are discussed mainly with regard to their feeding ecology. The slower growth and greater effect of sibling asymmetry on mortality of the Red Kite are consistent with our interpretation that the Red Kite's food supply is less abundant than the Black Kite's.     

Illegal and Legal Parrot Trade Shows a Long-Term,Cross-Cultural Preference for the Most Attractive Species Increasing Their Risk of Extinction

Illegal trade constitutes a major threat for a variety of wildlife. A criminology framework has been recently applied to parrot poaching in Mexico, suggesting an opportunistic crime in which the most abundant and accessible species, and not the rare or highly priced species, were poached more often. We analyzed this information, together with additional long-term data (1981–2005) on both the legal and illegal trade of the 22 Mexican parrot species (n = 31,019 individuals), using multivariate statistics and hypothesis-testing approaches. Our results showed a selective capture of parrot species attending to their attractiveness. Parrot species widely differed in attractiveness to people (as reflected by their combined measures of body size, coloration, and ability to imitate human speech), and their attractiveness strongly correlated with their prices both in the Mexican and US markets. The most attractive and valuable species (amazons and macaws) were disproportionally caught attending to the number of years they were legally trapped. Similar patterns were found for parrots poached for the domestic Mexican market, for those smuggled to the USA, and for those legally exported before or after 1992, when the USA ban led parrot exports to be mostly directed to European countries. Finally, the long-term cross-cultural preference for the most attractive species has led them to be among the most threatened species today. Since current parrot poaching mostly responds to local demand, socio-ecological work is needed to reverse the long-standing pet-keeping tradition that may decimate the most desired species in Neotropical countries.     

Growth of yellow-headed blackbird Xanthocephalus xanthocephalus nestlings in relation to maternal body condition, egg mass, and yolk carotenoids concentrations

Condition‐dependent resource allocation to eggs can affect offspring growth and survival, with potentially different effects on male and female offspring, particularly in sexually dimorphic species. We investigated the influence of maternal body condition (i.e., mass‐tarsus residuals) and two measures of female resource allocation (i.e., egg mass, yolk carotenoid concentrations) on nestling mass and growth rates in the polygynous and highly size dimorphic yellow‐headed blackbird Xanthocephalus xanthocephalus. Egg characteristics and carotenoid concentrations were obtained from the third‐laid egg of each clutch and were correlated with the mass and growth rates of the first two asynchronously hatched nestlings. Maternal body condition was associated with the growth of first‐hatched, but not second‐hatched nestlings. Specifically, females in better body condition produced larger and faster growing first‐hatched nestlings than females in poorer body condition. As predicted for a polygynous, size‐dimorphic species, females that fledged first‐hatched sons were in better body condition than females that fledged first‐hatched daughters. Associations between egg mass, yolk carotenoid content, and nestling growth were also specific to hatching‐order. Egg mass was positively correlated with the mass and growth rates of second‐hatched nestlings, and yolk concentrations of β‐carotene were positively correlated with second‐hatched nestling mass. Surprisingly, the relationship between yolk lutein and hatchling growth differed between the sexes. Females with high concentrations of yolk lutein produced larger and faster growing first‐hatched sons, but smaller first‐hatched daughters than females with lower lutein concentrations. Mass and growth rates did not differ between first‐ and second‐hatched nestlings of the same sex, despite asynchronous hatching in the species. Results from this study suggest that maternal body condition and the allocation of resources to eggs have carotenoid‐, sex‐, and/or hatch‐order‐specific effects on yellow‐headed blackbird nestlings.     

Heritability of Nestling Growth in Cross-Fostered European Starlings Sturnus Vulgaris

In altricial birds, growth rates and nestling morphology vary between broods. For natural selection to produce evolutionary change in these variables, there must exist heritable variation. Since nestling traits are not any longer present in parents, traditional offspring-parent regressions cannot estimate heritabilities of these. In this study, a partial cross-fostering experiment was performed, where nestlings of the European Starling (Sturnus vulgaris) were reciprocally exchanged between nests. The experiment demonstrated a significant heritability of nestling tarsus length and body mass, but not of the growth trajectories followed by individual nestlings. The heritability estimate for tarsus length obtained in the cross-fostering experiment using full-sib analysis was lower than those obtained by offspring-parent regressions. This is likely due to a genotype-by-environment effect on tarsus length, with nestlings destined to become large but in poor condition having a low probability of appearing as parents. The main reason for the low heritability of growth was probably the large within-brood variation in growth pattern due to the initial size hierarchy of nestlings. Nestlings demonstrated targeted growth, where small-sized nestlings that initially grew slower than their siblings, managed to catch up.     

Influence of stochastic processes and catastrophic events on the reproductive dynamics of the endangered Maroon‐fronted Parrot Rhynchopsitta terrisi

Stochastic and catastrophic events may strongly impact the dynamics of wild populations. Annual fluctuations in rainfall may affect parrot populations, but few studies address the impact of other stochastic or catastrophic events on their population dynamics. The Maroon‐fronted Parrot Rhynchopsitta terrisi is an endangered species that nests colonially in cavities and crevices in limestone cliffs. From 1995 to 2010, we quantified Parrot attendance at nesting colonies throughout its breeding range, and reproductive output of nesting Parrots from 1997 to 2010 at the two most important nesting colonies. There was significant variation among colonies in the number of cavities occupied by Parrots each year. Rainfall significantly influenced both the number of occupied cavities and productivity, which declined after very dry years. Natural unpredictable events such as hurricanes did not modify the nesting activity of Maroon‐fronted Parrots at breeding colonies. However, wildfires increased in dry years, negatively affecting attendance at breeding colonies. The Maroon‐fronted Parrot may overcome the impacts of climatic variability, natural stochastic processes, and human‐induced catastrophic events by using nesting colonies as a network of resources throughout the breeding range. Given the current trends in climate change, it is likely the species may suffer stronger and more frequent unpredictable catastrophic events, potentially putting at risk its survival in the long term.     

Effects of habitat fragmentation on provisioning rates, diet and breeding success in two species of tit (great tit and blue tit)

The aim of this study was to examine the effects of forest fragmentation on the ability of parent birds to provide their young with an adequate food supply. To examine whether prey population densities of the great tit (Parus major L.) and the blue tit (P. caeruleus L.) vary between study areas in different forest size classes we compared provisioning rates and chick diet and related these parameters to breeding success. We filmed 217 nests over two breeding seasons and collected data on frass fall as a general estimate of caterpillar availability. Nests which were attended by none or one parent only during filming (n = 46) were excluded from the analyses. In both years and for both species feeding rates were highest in the smallest fragments and lowest in the large forest. There was also a suggestion that differences in feeding rates between areas vary between years. We found no consistent tendency for prey size to change with forest size, although both species brought slightly smaller prey items to the nest in the smallest forest fragments and feeding rates correlated negatively with prey size. Caterpillars were the main item fed to nestlings, in both species. We found no evidence to suggest that either frass fall or the proportion of caterpillars in the diet varied with forest size. There was also no correlation between mean frass fall and the total number of caterpillars brought to the nests, in either species. Breeding success, as measured by clutch size, brood size, fledging weight and fledging success, did not differ between the small fragments and the large forest, in either species. There was also no relationship between provisioning rate (as concerns volume of prey fed to nestlings and the quality of chick diet) and breeding success parameters. In conclusion, this study does not suggest suboptimal foraging or breeding conditions in small fragments compared to a nearby large forest, for either species. Received: 23 June 1997 / Accepted: 29 December 1997     

The Influence of Habitat,Season, and Detectability on Abundance Estimates across an Amazonian Parrot Assemblage

The Amazon basin holds very high parrot species richness but almost nothing is known of parrot population densities in the region or how these vary between species, habitats, sites, and seasons. Such data are becoming important as humans impact on increasing areas of the region. Seventeen parrot species were surveyed using a line transect distance sampling method over 3 yr in floodplain and terra firme forests at two sites in the Tambopata region of southeast Peru. Density estimates for most species were in the range of 3.3–7.8/km², with Brotogeris cyanoptera and Amazona farinosa reaching densities of 22 and 23/km² in floodplain forest during the dry season. Parrot densities were higher in floodplain forest than in terra firme forest at both sites. The parrot communities of terra firme forests were similar across sites and seasons, but those in floodplain forests differed widely across sites and across seasons. Upper canopy birds are notoriously difficult to survey. We introduce a procedure to correct for the likely violation of the assumption that all birds on the transect line are detected (distance sampling assumption g(0) = 1). We correct g(0) based on calling rates of birds using a cue‐counting technique. Multipliers for g(0) differ across species and site, but not season. This method yielded density estimates on average 22 percent higher (6–40% higher in individual species) than those from the standard method.     

Predicting co-distribution patterns of parrots and woody plants under global changes: The case of the Lilac-crowned Amazon and Neotropical dry forests

Global climate and land-use changes are the most significant causes of the current habitat loss and biodiversity crisis. Although there is information measuring these global changes, we lack a full understanding of how they impact community assemblies and species interactions across ecosystems. Herein, we assessed the potential distribution of eight key woody plant species associated with the habitat of the endangered Lilac-crowned Amazon (Amazon finschi) under global changes scenarios (2050′s and 2070′s), to answer the following questions: (1) how do predicted climate and land-use changes impact these species’ individual distributions and co-distribution patterns?; and (2) how effective is the existing Protected Area network for safeguarding the parrot species, the plant species, and their biological interactions? Our projections were consistent identifying the species that are most vulnerable to climate change. The distribution ranges of most of the species tended to decrease under future climates. These effects were strongly exacerbated when incorporating land-use changes into models. Even within existing protected areas, >50 % of the species’ remaining distribution and sites with the highest plant richness were predicted to be lost in the future under these combined scenarios. Currently, both individual species ranges and sites of highest richness of plants, shelter a high proportion (ca. 40 %) of the Lilac-crowned Amazon distribution. However, this spatial congruence could be reduced in the future, potentially disrupting the ecological associations among these taxa. We provide novel evidence for decision-makers to enhance conservation efforts to attain the long-term protection of this endangered Mexican endemic parrot and its habitat.     

Diet and Geophagy Across a Western Amazonian Parrot Assemblage

We identified species‐ and community‐level dietary characteristics for a species‐rich Amazonian parrot assemblage to determine relationships among dietary metrics and use of geophagy sites. Previous studies suggest that soil is consumed at geophagy sites in this region mainly to supplement dietary sodium. We accumulated 1400 feeding records for 16 parrot species over 2 yr and found that seeds, flowers, and fruit pulp featured prominently in diets, while bark, insects, and lichen were consumed in small quantities. Food availability across 1819 trees was measured, and we found that flower availability was highest in the dry season and fruit production peaked in the wet season, but that phenology patterns of the 20 most commonly foraged plant species suggest no serious food bottlenecks. Partitioning of available food resources among the 13 most commonly encountered parrots is suggested by an ordination analysis (DCA), which placed the large macaws (Ara) with the Amazona parrots at the ‘primary forest’ end of a dietary resource axis and four smaller species at the ‘successional forest’ end of the axis. Parrot species associated with successional forest also consumed less plant species overall. Furthermore, these parrot species consuming successional forest resources had higher claylick visitation rates than those consuming primary forest resources suggesting they derive the greatest benefits from soil consumption.     

GROWTH RATES OF BIRDS IN THE HUMID NEW WORLD TROPICS

The growth curves of 40 species of lowland neotropical birds were fitted by logistic equation. The birds were mostly from Panama, Trinidad and Surinam. The growth constants of the fitted equations (asymptote A and growth rate K) were compared within and among species, and with previously published data on temperate species. Growth parameters of tropical passerines are about as variable within species as they are within temperate species. In both cases, variation in A and K between broods is greater than it is within broods. Panamanian birds breed during the dry-wet transition and conditions for growth apparently improve as rainfall increases. Asymptotes of growth curves are higher, and mortality within broods lower, as the breeding season progresses. Asynchronous hatching and the reduction of brood-size by selective starvation of young is a prominent phenomenon during the early part of the breeding season. Several instances are reported, however, of young persisting in nests with inadequate feeding and greatly subnormal weights. Slowed development under conditions of poor nutrition may be adaptive in the tropics if periods of low food availability are short and allow the possibility of recovery from undernourishment. As a group, neotropical lowland passerines (30 species) grow 23% more slowly than a sample of 51 temperate passerines. Variation of growth rates among these tropical species is similar to variation among temperate species, and it is related to adult body-size the length of the nestling period. Young of tropical and temperate species attain similar asymptotes, relative to adult body-weight, by the end of the nestling period. Hypotheses are advanced which might explain the slower growth rate of tropical species, and tested to the extent available data permits. (1) Because brood-size can be changed only by adding or removing whole young, changes in growth rate could provide finer adjustment of the energy requirements of the young to the feeding capacity of the parents. This model predicts different means and variances for growth rate within groups of species with different clutch-sizes, predictions not supported by available data. (2) Growth rate is shown to increase the maximum energy requirement of a nestling only if K exceeds some value determined by the energy requirement of the young, growth rate should vary in proportion to the level of basal maintenance metabolism. In a small sample of tropical species, rates of basal metabolism were 25% lower than in a comparable sample of temperate species. These data therefore support the hypothesis, although the cause of the lower metabolic rate of the tropical nestlings is not known. (3) Daily periods of hypothermia could reduce the energy requirement of the young and at the same time reduce their growth rate; but observations of body temperatures of tropical nestlings are contrary to this hypothesis. (4) The short day-length of tropical climates reduces the time during which young can assimilate energy relative to their energy expenditures. This model predicts that tropical nestlings would have less productive energy available, (consistent with their reduced growth rates), but it also predicts that arctic birds should grow faster than temperate species, which is not confirmed by available data. (5) The low nitrogen content of fruits may cause the slow growth of a few strictly frugivorous species (Oilbird and Bearded Bellbird), but among other tropical species growth rate is not correlated with the estimated proportion of fruit in the diet.     

Brood size and environmental conditions sex-specifically affect nestling immune response in the European starling Sturnus vulgaris

In sexually size-dimorphic species, the larger sex can be more sensitive to stressful environmental conditions, often resulting in reduced growth and elevated mortality rates. Development of the immune system is regarded as highly resource dependent, and recent data suggest that nestling passerines experience a possible resource-based trade-off between growth and immunity. Given the hypothesized importance of maximizing growth for the larger sex, the corresponding immune system may also exhibit similar sensitivity to limited resources. To better understand how natural variation in brood size and resources might differentially affect growth and immune function in nestlings of a sexually size-dimorphic species, we examined the relationship between brood size and inter-sexual differences in cell-mediated immunity (CMI) and survival in European starling Sturnus vulgaris nestlings where males are larger in both mass and structural size. We hypothesized that male CMI response should be negatively impacted by increasing sibling competition (brood size), especially during periods of low resource availability. In a year of reduced parental provisioning rates and reduced chick growth rates, male offspring exhibited the predicted negative relationship, whereas female CMI response was unaffected. However, in a year of improved provisioning rates and chick growth, neither sex exhibited a negative relationship between immune response and brood size. Thus, natural variation in brood size can affect sex-specific immunity differently in offspring of a sexually size-dimorphic passerine. However, this relationship appears resource-dependent, suggesting that the hypothesized resource-based trade-off may be compensated for in years of adequate resource abundance.     

The dynamics of recovery and growth: how defoliation affects stored resources

Growth rate varies widely among species and the trade-off between growth rate and storage or maintenance traits is a principal axis of variation between species. Many plant species have substantial root stores, but very little is known about how growth rate modifies responses of these stores to defoliation and other stresses. Species with different growth rates are predicted to respond in distinct ways, because of variation in the pre-defoliation allocation to storage. Here, we quantified the dynamics of stored carbohydrates in seven species with varying growth rate, following defoliation in a pot experiment. For faster growing species, there was significant reduction in carbohydrate concentration following defoliation, followed by relatively fast recovery, whereas for slower growing species, carbohydrate concentration levels remained relatively invariant across treatments. Results for total carbohydrates mirrored those for concentration, but were not as significant. Our findings were consistent with the idea that faster growing species respond more rapidly than slower growers to defoliation, through changes in carbohydrate pool concentrations. Growth rate as an indicator of life-history and ecological strategy may therefore be key to understanding post-defoliation recovery and storage strategies.     

Is hatching asynchrony beneficial for the brood?

Many hypotheses have been proposed to explain why female birdsstart to incubate before clutch completion (IBCC). Some of thosesuggest that the resulting hatching asynchrony (HA) is adaptivebecause it increases the size hierarchy among offspring andin turn reduces nestling competition and energy demands duringthe peak feeding period. Others argue that IBCC is a good strategyin unpredictable environments. When food conditions deteriorate,the large size hierarchy quickly results in the death of thelast hatched nestlings, allowing the remaining ones to surviveand fledge in better condition. In comparison, under favorableconditions, all nestlings can fledge independent of hatchingorder. To test these hypotheses, we performed a brood size manipulationexperiment (as a simulation of good and bad years) in collaredflycatchers Ficedula albicollis and examined the effect of sizehierarchy on offspring and brood performance. We found thatchicks with an initial size disadvantage experienced reducedbody mass growth and had shorter feathers at fledging in bothreduced and enlarged broods. In enlarged broods, they also fledgedwith a smaller skeletal size. Although broods on average orparents could possibly still benefit from HA when food is scarce,this was not seen in the current study. Parental survival wasnot related to the size hierarchy in the broods, and the averagebody mass growth of the nestlings was slower in broods witha high initial size variance. We therefore conclude that HAand the resulting size hierarchy are probably detrimental forthe growth of nestlings in both good and bad years, at leastin species where nestling mortality does not occur early inlife.     

Azure‐winged Magpies Cyanopica cyanus trade off reproductive success and parental care by establishing a size hierarchy among nestlings

It is common in birds that the sizes of nestlings vary greatly when multiple young are produced in one nest. However, the methods used by parents to establish size hierarchy among nestlings and their effect on parental provisioning pattern may differ between species. In the Azure‐winged Magpie Cyanopica cyanus, we explored how and why parents controlled the sizes of nestlings. Asynchronous hatching was the main cause of size hierarchy within the brood, although the laying of larger eggs later in the laying sequence reduced this effect. Parents with asynchronous broods produced more eggs and fledged more nestlings than those with synchronous broods but their brood provisioning rates, food delivery per feeding bout and feeding efficiency did not differ. We performed a cross‐fostering experiment to synchronize some asynchronous broods. Provisioning rates of asynchronous broods were lower than those of synchronized broods, but the daily growth rates and fledging body mass of their nestlings were not different. Our findings indicate that parents of asynchronous broods can achieve higher reproductive success than those of synchronous broods based on the same parental care, and the same reproductive success as those of synchronized broods based on less parental care. It appears that parent birds can better trade off reproductive success and parental care by establishing a size hierarchy among nestlings.     

The effects of hatching asynchrony on growth and mortality patterns in Eurasian Hoopoe Upupa epops nestlings

Growth is a fundamental life history trait in all organisms and is closely related to individual fitness. In altricial birds, growth of many traits is restricted to the short period between hatching and fledging and strongly depends on the amount of food that parents deliver and the extent of hatching asynchrony. However, empirical studies of energy allocation to growth of different body size traits as a function of hatching asynchrony are scarce. We studied growth and mortality of Eurasian Hoopoe Upupa epops, a species with a long breeding season and high brood size variance, whose nestlings show pronounced hatching asynchrony, in order to test how hatching asynchrony affects different growth traits in the context of territory quality, season and brood size. The growth of five body traits (body mass, and lengths of tarsus, third primary, bill and longest crest feather) was investigated to understand how it was affected by brood size, hatching date and order, and territory quality. In total, 241 nestlings from 39 nests were measured every 4 days in 2014 in south‐western Switzerland. Brood size, hatching date and hatching order had the strongest influence on growth trajectories, although tarsus growth was only marginally affected by these variables. Nestlings that hatched earlier than their siblings were heavier and had longer third primaries, bills and crest feathers compared with later‐hatched siblings. In territories of high quality, hatching order differences disappeared for body mass growth, but persisted for lengths of third primary, bill and crest feathers. Brood size was inversely associated with third primary, bill and crest feather lengths, but positively associated with body mass. Nestling mortality was higher in later‐hatched nestlings and in broods that were raised in territories of lower quality. Our study shows that in nestlings, energy was allocated differentially between body traits and this allocation interacted with hatching order and territory quality. Rapid mass gain by nestlings was prioritized in order to increase competitive ability. Our results provide support for the brood reduction hypothesis as an explanation of hatching asynchrony in Hoopoes.     

Nesting Habitat of the Lilac-crowned Parrot in a Modified Landscape in Mexico

Parrot populations are being increasingly pressured to occupy modified or fragmented landscapes, yet little is known of the habitat requirements of most species, particularly with regard to the effects on breeding habitat. We evaluated nesting habitat of the lilac-crowned parrot Amazona finschi in the modified landscape of coastal Michoacan in Mexico. We located 90 parrot nests in 12 tree species in Michoacan, with lilac-crowned parrots presenting a narrow niche-breadth of tree species used for nesting. Considering an additional 82 nest trees recorded for lilac-crowned parrots in Jalisco, we determined a 51 percent similarity in cavity resource use by parrots in the two dry forest regions. Overall, the predominant nest tree species with 76 percent of nests were Astronium graveolens , Piranhea mexicana , Brosimum alicastrum , and Tabebuia spp., all characteristic of semi-deciduous forest. Only 8 percent of nests occurred in trees characteristic of deciduous forest. Parrots utilized large trees with canopy level cavities as nest sites, and preferred conserved semi-deciduous forest for nesting, with fewer nests than expected in deciduous forest and transformed agricultural land. Nest areas in semi-deciduous forest occurred on significantly steeper terrain, as remnant semi-deciduous forest is restricted to steep ridges and canyons. Those parrot nests in modified habitats and forest patches were located near to continuous forest, with nest trees in open agricultural land being significantly closer to continuous forest than nests in disturbed forest patches. These results demonstrate the importance of conserved semi-deciduous forest as breeding habitat for the threatened, endemic lilac-crowned parrot, making wild populations of the species vulnerable to the high rate of transformation and fragmentation of tropical dry forest.     

Parent age differentially influences offspring size over the course of development in Laysan albatross

Offspring growth and survival are predicted to be higher for older parents, due to a variety of mechanisms, such as increased breeding experience or greater investment favored by low residual reproductive value. Yet the extent to which parent age affects offspring viability is likely to vary between different aspects of growth and survival, perhaps being most pronounced at the most stressful stages of reproduction. We studied the link between parent age and nestling growth and survival in the Laysan albatross, a long-lived seabird with a mean first breeding age of 8 years. Offspring of older parents were more likely to survive to fledging. Among those that did fledge, nestlings of older parents grew more rapidly. However, parent age did not influence the eventual asymptotic size that nestlings reached before fledging: fast-growing nestlings of older parents reached 90% of asymptotic size roughly 1 week sooner, but slow-growing nestlings of younger parents eventually caught up in size before fledging. Older parents bred c . 2 days earlier than younger parents, but hatch date did not explain observed variation in offspring success. The extent to which parent age accounted for variation in size of individual nestlings was not constant but peaked near the midpoint of development. This could reflect a time period when demands on parents reveal age-based differences in parental quality. Overall, growth and survival of offspring increased with parent age in this species, even though the late age of first breeding potentially provides a 7-year period for birds to hone their foraging skills or for selection to eliminate low-quality individuals.     

三种鹭异步孵化与雏鸟生长的比较研究

根据1996～1999年的野外工作和实验饲养,研究了池鹭、白鹭、夜鹭3种鹭的繁殖、雏鸟生长和异步孵化对雏鸟生长的影响.3种鹭于4月上、中旬迁到常山、余杭鹭保护区,9月下旬陆续迁离.孵卵期池鹭为23.0 d(n=26),白鹭23.9 d(n=32),夜鹭25.3 d(n=34);孵化率池鹭为76.32%,自鹭86.96%,夜鹭95.45%,池鹭繁殖力为3.21只,白鹭3.38只,夜鹭3.50只.雏鸟体重增长与成体体形大小呈负相关.幼雏体重增长与孵化顺序相关.在生长早期(≤8 d)全部雏鸟正常发育,之后差异显著.第1和第2孵化雏鸟生长曲线相似,但明显高于第4雏鸟,表明较早孵化者获得食物能力较强,而迟孵出雏鸟有食物不足现象.该3种鹭孵育幼雏能力的最适度估计为3只左右.