Polluted environment and cold weather induce laying gaps in great tit and pied flycatcher

We studied the occurrence of laying gaps in free-living populations of the pied flycatcher, Ficedula hypoleuca, and the great tit, Parus major, in a pollution gradient of a copper smelter in south-west Finland. Laying gaps were 2.8 times more common in F. hypoleuca than in P. major. The probability of laying gaps was highest in the heavily polluted zone and lowest in the unpolluted zone for both bird species. Cold weather at the time of laying increased the number of laying gaps in both species, but in P. major this effect was most pronounced in the heavily polluted environment. In the most heavily polluted environment the laying gaps were more likely to occur near the beginning of the laying sequence in both species. The laying gap probability increased with increasing laying date in P. major but not in F. hypoleuca. We suggest that the increased number of laying gaps in the polluted environment results from limited Ca availability and the interference of heavy metals with Ca metabolism in laying females.     

Growth and mortality of nestling great tits (Parus major) and pied flycatchers (Ficedula hypoleuca) in a heavy metal pollution gradient

We studied nestling growth, growth abnormalities, mortality and breeding success of two hole-nesting passerines, the great tit (Parus major) and the pied fly-catcher (Ficedula hypoleuca), at 14 study sites around a copper smelter complex in Harjavalta, south-west Finland in 1991–1993. The main pollutants in the area are heavy metals and sulphuric oxides. Exposure of birds to heavy metals was shown by measuring their faecal concentrations. Copper, nickel and lead contents of nestling faeces were high near the factory and decreased with distance away from the pollution source. F. hypoleuca nestlings suffered high mortality very close to the factory complex, but did relatively well at all other sites. Breeding success of P. major was below background levels up to 3–4 km from the pollution source and nestlings grew poorly close to the factory. Growth abnormalities of legs and wings in F. hypoleuca nestlings were significantly more common near the factory than farther away. In contrast, F. hypoleuca nestlings grew equally well at all distances. The poor breeding success of F. hypoleuca close to the factory complex is probably related to the high amount of heavy metals in its diet, and low availability of calcium-rich food items may enhance this effect. We suggest that the poor breeding success of P. major is related to habitat changes that have taken place around the factory. The different responses of these two bird species are probably due to their different diet. Our results show convincingly that species-specific differences in response should be carefully considered when planning schemes for air pollution monitoring.     

The effects of diet quality and quantity on plumage colour and growth of great tit Parus major nestlings: a food manipulation experiment along a pollution gradient

The yellow carotenoid-based plumage coloration of great tit Parus major nestlings is found to be paler in polluted and urban environments. Because carotenoid pigmentation is often considered to be a condition dependent trait in birds we wanted to find out whether food-limitation and poor nestling condition could explain the pale plumage colour in a polluted area. P. major nestlings were supplemented with variable diets along a well known heavy metal pollution gradient around a copper smelter: two food treatments with carotenoids, one food treatment with little carotenoid and one unsupplemented control. Our field experiment showed that nestlings in the polluted area grew better with carotenoid rich diets, while such effect was not found in the unpolluted area. Nestlings showed higher plasma carotenoid (lutein) levels and higher plumage carotenoid chroma values in the unpolluted area than in the polluted area. However, plasma lutein levels or plumage colour were not associated with heavy metal levels in nestling faeces (a proxy for dietary exposure). Our results provide only weak evidence for carotenoid-based colouration to be condition-dependent in great tit nestlings as we found a positive relationship between body mass and carotenoid chroma in the non-supplemented control group only. The positive relationship between body mass and plumage colour intensity is more likely to be produced by the fact that good availability of caterpillars, an important food source for P. major, also means a good availability of carotenoids to nestlings. Our results suggest that main reason for pale nestling plumage in the polluted area is lower quality invertebrate food, and not nutrition-related oxidative stress.     

Egg shell quality,clutch size and hatching success of the great tit (Parus major) and the pied flycatcher (Ficedula hypoleuca) in an air pollution gradient

Egg shell thickness, egg volume, clutch size and hatching success of Parus major and Ficedula hypoleuca were studied at 14 study sites around a copper smelter complex in Harjavalta, south-west Finland, in 1991–1993. In 1991–1992 unhatched eggs were collected to measure shell quality. F. hypoleuca was more susceptible to pollutants than P. major, the response of which was weaker in all aspects studied. Egg shells of F. hypoleuca were about 17% thinner and eggs were about 8% smaller in volume near the factory than at a distance of 10 km. The clutch size of F. hypoleuca was significantly smaller and hatching success markedly lower at a study site next to the factory complex than at all other sites. In P. major, variation in shell thickness and egg volume was not significantly related to the distance from the pollution source. Clutch size and hatching success of P. major did not significantly differ among study sites, although the trend in hatching success was in the same direction as in F. hypoleuca. Clutches of both species contained less shell material and both species had more nests without eggs near the factory than further away. The surface structure of the eggshells was studied by scanning electron microscope. Especially in F. hypoleuca, the egg shell surface was more rough and porous near the factory. The roles of Ca and heavy metals in shell thinning are discussed.     

The diet of urban Magpies Pica Pica

This paper describes the diet of Magpies in the Manchester urban environment, based on analysis of stomach contents, pellets and faecal material. The diet comprised a selection of typical grassland fauna, the majority of which were surface-dwelling invertebrates. Weevils, click beetles, ground beetles, flies, caterpillars and spiders were common. However, soil-dwelling animals such as leatherjackets and worms were estimated to form a significant proportion of the weight of food. Vertebrate remains, such as those of the field vole Microtus agrestis and the House Sparrow Passer domesticus, were present at a low frequency. The most common constituents of the nestling diet were invertebrate groups that attained their peak seasonal abundance during the nestling period, which suggests that food resources may be utilized in proportion to their availability. During the winter when invertebrate material is scarce, the diet is largely composed of plant material; Magpies may experience a food shortage at this time of year. The summer diet of the adult is similar to that of nestling Magpies; neither include many eggs or nestlings.     

Leaf fluctuating asymmetry of common plantain as an indicator of habitat quality

Abstract

Although developmental instability (DI), measured as fluctuating asymmetry (FA), is expected to be positively related to environmental stress and negatively to habitat quality, the pattern found here was the reverse. Developmental instability of leaf traits (leaf width and vein distances within a leaf) was estimated (using two indices of FA: FA₄ and σ_i ²) and compared between three populations of Plantago major L. (Plantaginaceae) from northern Serbia. Two of the populations are from chronically polluted areas (Karaburma & Zemun), while Crni Lug is from an unpolluted, natural area. Results obtained using both FA indices were the same; higher asymmetry levels in the unpolluted area than in the polluted sites, were found for both traits. Between the two polluted sites, FA values were significantly higher in Karaburma site for vein distances within a leaf. Concerning differences in FA₄ values between samples, in two cases, results are similar to those found for σ_i ² values, for vein distances within leaf. These are the first quantitative data on P. major indicating that (i) plants living in the stressful sites are more symmetrical and (ii) leaf FA for plant species with wide ecological distribution such as P. major should be considered as an ‘index of habitat quality.’     

Activity and heavy metal resistance of non-specific esterases in leaf beetle Chrysomela lapponica from polluted and unpolluted habitats

We compared the general activity and heavy metal resistance of non-specific esterases in two populations of the leaf beetle Chrysomela lapponica from habitats severely contaminated by heavy metals (mostly Ni and Cu) and two populations from unpolluted habitats. Concentrations of Ni and Cu in adult beetles from the most polluted site were 7.7 and 3.6 times higher that in beetles from unpolluted habitats. Larval esterases showed higher activity and lower susceptibility to heavy metals than esterases of adults. Larval esterase activity did not differ between populations from polluted and unpolluted sites, but adult beetles from polluted localities had lower esterase activity than beetles from unpolluted habitats. Both Cu and Ni sulfates in millimolar concentrations in vitro suppressed esterase activity of larvae from unpolluted habitats, but caused no negative effect on esterases of larvae from polluted sites. Similarly, inhibition of adult esterase activity by Ni was stronger in beetles from unpolluted localities than in beetles from polluted localities. This indicates that resistance of non-specific esterases to heavy metals is higher in leaf beetle populations from contaminated environment.     

Environmental Pollution Affects the Plumage Color of Great Tit Nestlings through Carotenoid Availability

Birds need to acquire carotenoids for their feather pigmentation from their diet, which means that their plumage color may change as a consequence of human impact on their environment. For example, the carotenoid-based plumage coloration of Great tit, Parus major, nestlings is associated with the degree of environmental pollution. Breast feathers of birds in territories exposed to heavy metals are less yellow than those in unpolluted environments. Here we tested two hypotheses that could explain the observed pattern: (I) deficiency of carotenoids in diet, and (II) pollution-related changes in transfer of carotenoids to feathers. We manipulated dietary carotenoid levels of nestlings and measured the responses in plumage color and tissue concentrations. Our carotenoid supplementation produced the same response in tissue carotenoid concentrations and plumage color in polluted and unpolluted environments. Variation in heavy metal levels did not explain the variation in tissue (yolk, plasma, and feathers) carotenoid concentrations and was not related to plumage coloration. Instead, the variation in plumage yellowness was associated with the availability of carotenoid-rich caterpillars in territories. Our results support the hypothesis that the primary reason for pollution-related variation in plumage color is carotenoid deficiency in the diet.     

Breeding success and lutein availability in great tit (Parus major)

The relationship among temporal variation in the availability of carotenoid-rich food, tissue carotenoid levels and breeding success are poorly known. We studied how diet quality and quantity affect the carotenoid profile and fledging success of great tit (Parus major) nestlings along a pollution gradient. We found declining seasonal trend in lutein concentration of caterpillars, which may be the explanation for the declining trend in nestlings' lutein concentration of plasma with season, despite the increase in caterpillar biomass. This may be because the biomass of most lutein-rich caterpillars (autumnal moths) decreased and less lutein-rich caterpillars (sawflies) increased during the breeding season. The temporal difference in occurrence of different caterpillar species means that peak lutein availability does not coincide with peak caterpillar abundance. However the positive association between total larval biomass and the number of great tit fledglings may suggest that fledging success depends more on total caterpillar availability than on lutein concentration of caterpillars.     

Density effect on great tit (Parus major) clutch size intensifies in a polluted environment

Long-term data on a great tit (Parus major) population breeding in a metal-polluted zone around a copper–nickel smelter indicate that, against expectations, the clutch size of this species is decreasing even though metal emissions in the area have decreased considerably over the past two decades. Here, we document long-term population-level changes in the clutch size of P. major and explore if changes in population density, population numbers of competing species, timing of breeding, breeding habitat, or female age distribution can explain decreasing clutch sizes. Clutch size of P. major decreased by one egg in the polluted zone during the past 21 years, while there was no significant change in clutch size in the unpolluted reference zone over this time period. Density of P. major nests was similar in both environments but increased threefold during the study period in both areas (from 0.8 to 2.4 nest/ha). In the polluted zone, clutch size has decreased as a response to a considerable increase in population density, while a corresponding density change in the unpolluted zone did not have such an effect. The other factors studied did not explain the clutch size trend. Fledgling numbers in the polluted environment have been relatively low since the beginning of the study period, and they do not show a corresponding decrease to that noted for the clutch size over the same time period. Our study shows that responses of commonly measured life-history parameters to anthropogenic pollution depend on the structure of the breeding population. Interactions between pollution and intrinsic population characters should therefore be taken into account in environmental studies.     

On the advantage of Folsomia fimetarioides over Isotomiella minor (Collembola) in a metal polluted soil

Summary Food preference, metal avoidance behaviour and desiccation tolerance were investigated for two collembolan species, Folsomia fimetarioides and Isotomiella minor, in order to test some hypotheses for their different distributions in relation to a heavy metal gradient. F. fimetarioides showed a higher preference for metal tolerant fungi than I. minor when they were offered fungi of different metal tolerance. When they could choose between polluted and unpolluted fungi and substrate, F. fimetarioides significantly avoided the polluted source while I. minor did not. Both species were negatively affected by drought. I. minor was more tolerant than F. fimetarioides when exposed to drought in combination with metal pollution. High supply of preferred fungal species and a better ability to avoid heavy metals favour F. fimetarioides and determine its dominance over I. minor in polluted soils.     

Habitat quality, nestling diet, and provisioning behaviour in great tits Parus major

Variation in early nutrition is often a strong predictor of offspring condition and fitness. In the case of woodland passerine birds, nestling diet is determined by the spatiotemporal distribution of prey items such as caterpillars during the nestling period, and is usually quantified as differences in provisioning behaviour between habitats. However, the habitat level does not account for variation between individual territories, the level at which competition and selection are assumed to operate. Here we use nestbox cameras and Radio Frequency Identification technology (RFID) to simultaneously assess variation in both nestling diet (components) and provisioning rates (quantity) among a sample (n=22) of different quality great tit Parus major territories selected from a larger breeding population (n=310 fledged broods) in a single year. Caterpillars were by far the most numerous item provisioned to nestlings (mean=75% of prey items), as expected given the known importance of this food source for this species. Broods raised close to an oak tree, or far from the woodland edge, were provisioned the highest proportion of caterpillars. Provisioning rates declined seasonally and there was a weak association between low provisioning rates and caterpillar rich diets. During the first week of the nestling stage, nestling condition was unrelated to the proportion of caterpillars in the diet, provisioning rates and oak proximity. Condition at fledging was slightly improved in broods fed a higher proportion of caterpillars in the diet and in broods raised close to an oak tree. However, in our data early breeding was the only predictor of recruitment success, although power was low for this test. Analyses of long‐term data (41 years) from the same population confirmed a relationship between oak proximity and fledgling mass, but not recruitment success. Our results suggest that territory level environmental variation can affect offspring condition, probably through observed changes in nestling diet, but that such variation does not necessarily produce discernable effects on offspring fitness.     

Seasonal variation in diet quality: antioxidants,invertebrates and blue tits Cyanistes caeruleus

Breeding success is often dictated by the degree to which parents can synchronize the maximum food requirements of offspring to the peak in abundance of invertebrate prey. Less studied is how the nutritional quality of individual diet items impacts on breeding. In the present study, we assessed the abundance and antioxidant concentrations of arboreal arthropods from oak woodland and provisioning behaviour of the blue tit Cyanistes caeruleus. Dietary antioxidants are important during development because they defend against oxidative stress. Operophtera caterpillars, Erannis caterpillars, and spiders contained significantly different levels of individual carotenoids and α‐tocopherol. Concentrations of lutein and β‐carotene in Operophtera caterpillars did not vary seasonally, although concentrations of zeaxanthin declined and α‐tocopherol increased with date. Blue tit broods hatched later in the season received significantly fewer caterpillars and more spiders per chick compared to earlier broods. Reflecting changes in prey composition, blue tit nestling plasma showed decreases in zeaxanthin and increases in α‐tocopherol with date. Thus, processes that shift the timing of breeding in birds and/or prey composition are likely to alter antioxidant intake and thus potentially influence the oxidative stress status of animals. The data obtained in the present study suggest a mechanism by which environmental change as a result of human activities could influence the health and fitness of individuals in natural populations. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 708–717.     

Concentration and ecological risk of heavy metal in street dusts of Eslamshahr,Iran

This study was done to evaluate heavy metal concentrations in street dust samples, to compare measured concentrations in samples to background concentrations in order to make evaluations for pollution indices, and to describe the quality of street dust in the studied area in relation to pollution. A total of 30 cumulative samples were collected from the streets of Eslamshahr City. Concentrations of heavy metals were determined using an Inductively Coupled Plasma-Optical Emission Spectrometry (ICP-OES). Results determined mean concentrations (mg/kg) of the heavy metals Cd, Cr, Cu, Ni, Pb, and Zn, in collected samples of street dust as 0.34, 35.1, 239, 42.4, 71.3, and 573, respectively. I_geo values for Cd and Cr, Cu, Ni, Pb, and Zn showed level of moderately polluted, unpolluted, moderately to strongly polluted, unpolluted, moderately polluted and moderately to strongly polluted, respectively. The pattern of total metal concentrations in the studied area was ranked as follows: Zn and Cu>Pb>Cd>Ni>Cr. The highest values for the monomial potential ecological risk (E_r) were observed for Cd (114). The mean level of RI for the studied soil samples was 192 (91.3–244), which is classed as presenting a strong potential ecological risk.     

Seasonal insectivory by Black‐headed Trogons,a tropical dry forest frugivore

ABSTRACT Fruit‐eating birds are important seed dispersers in tropical forests, but little is known about the extent to which they rely on insects or how their diets vary seasonally. We used field observations of focal adults to quantify the diets of adult and nestling Black‐headed Trogons (Trogon melanocephalus) at nine nests in a lowland dry forest in Costa Rica. From May 2004 to August 2004, we documented 540 food deliveries to nests and 1080 food items consumed by adults. Adult and nestling trogons were largely insectivorous, feeding mainly on moth caterpillars (Lepidoptera). Fruit accounted for only 10.5% of items consumed by adults and 2.2% of items delivered to nestlings (6.1% and 0.6% of estimated dry mass, respectively). Adult and nestling diets differed significantly in both composition and prey size, with adults consuming more fruit and fewer large insects (Phasmatodea and Mantodea) than nestlings and eating more types of arthropods and fruit. Although both adults and nestlings relied heavily on moth larvae, adults preferentially consumed small caterpillars and delivered large ones to their nestlings. In addition, the proportion of large caterpillars delivered to nests remained constant throughout the nestling period, whereas the proportion of large caterpillars eaten by adults declined significantly with nestling age. Overall, arthropods delivered to nests averaged 70% heavier than those consumed by adults (estimated dry mass). Our results suggest that Black‐headed Trogons time reproduction to coincide with arthropod rather than fruit abundance, a pattern that may be more common among omnivorous forest birds than previously recognized.     

NESTLING DIETS OF PRAIRIE PASSERINE BIRDS AT MATADOR, SASKATCHEWAN, CANADA

The nestling diets of seven species of grassland passerines were studied from gullet samples collected in 1969–1971 at Matador, Saskatchewan, Canada. Relative population densities on census plots on grazed and ungrazed prairie showed that Horned Lark Eremophila alpestris, McCown's Longspur Calcarius mccownii and Chestnut-collared Longspur C. ornatus preferred grazed areas and Sprague's Pipit Anthus spragueii, Western Meadowlark, Sturnella neglecta, Savannah Sparrow Passerculus sandwichensis and Baird's Sparrow Ammodramus bairdii preferred ungrazed prairie. Fifteen invertebrate ordinal taxa, and seeds were represented to an important extent in combined gullet contents; but two taxa, Lepidoptera (77% larvae) and grasshoppers (all Acrididae), comprised the bulk of nestling diets. Four grasshopper species contributed 70–100% of grasshoppers eaten in June-August. Overlap at the family level was also high in several other important prey. Comparison with invertebrate populations of ungrazed prairie showed that the birds ignored some abundant taxa of very small invertebrates, and apparently preferred the two least abundant groups which offered large individual prey items. Average nestling food overlap indices for the season among all species except Savannah Sparrow and Baird's Sparrow, based on taxonomic composition, was 0.85. Overlap was moderate in May and June and became very high in July and August. The number of nestlings reached a peak in June and early July when nestling food overlap was much lower than later in the season. Horned Larks avoid competition by nesting earlier than do the other species, while McCown's Longspur, which depends on grasshoppers for nestling food more than do the other species, tends to breed later in the season than the other species. Finally, data suggest that nestling food may be superabundant.     

Obtaining accurate measurements of the size and volume of insects fed to nestlings from video recordings

Video recordings are commonly used to study the types, amount, and size of food items provided to nestling birds. However, the accuracy and repeatability of estimates of the size of food items from video recordings has not been examined. We assessed three aspects of the reliability of measuring prey size from video recordings of Great Tits (Parus major) provisioning nestlings. To test the accuracy of measurements of prey size (length and width) used to determine prey volume, we molded artificial plasticine caterpillars and compared their size and volume as determined using measurements of length and width on screenshots of video recordings (using the vertical diameter of nest‐box entrance holes as a size reference) to their actual size and volume. We also examined within‐ and among‐observer repeatability of measurements of the size and volume of actual prey items delivered to nestlings by adult Great Tits. We found that observers were able to accurately measure prey size and determine volume, with high agreement between the actual size and volume of plasticine caterpillars and the size and volume as determined from measurements made on screenshots from video recordings (r_ICC = 0.99). In addition, within‐ and among‐observer repeatability were also high (r_ICC = 0.98 and 0.93, respectively). Overall, our results suggest that the size of prey items delivered to nestlings by adults in video recordings can be accurately measured and those measurements, in turn, can be used to accurately determine the volume of those insect prey.     

Pollution related effects on immune function and stress in a free-living population of pied flycatcher Ficedula hypoleuca

We investigated whether exposure to heavy metal pollution affected the immune function of individuals in a free living population of a small insectivorous passerine bird, the pied flycatcher Ficedula hypoleuca . We measured humoral immune responses in two study areas: a polluted area in the vicinity of a copper smelter and a control area far from the smelter. Plasma corticosterone level and blood heterophil/lymphocyte ratio (H/L) were used as more general physiological measures of stress. The immune response of F. hypoleuca was not suppressed by pollution stress. In contrast, we found that F. hypoleuca males showed stronger humoral immune responses to a novel antigen (tetanus toxoid) in the polluted environment than in the unpolluted one. After the immunization of males, numbers of lymphocytes rose significantly more in the polluted area, leading to a smaller H/L ratio than in males from the control area. Females showed no pollution related effects on their immune responses. Corticosterone levels of males and nestlings were not related to pollution levels. Nestlings showed somewhat higher H/L ratios and lower fledging success in the polluted area, both factors indicating increased stress levels in a polluted area. Our results suggest that humoral immune response of male F. hypoleuca may be enhanced under moderate levels of heavy metal pollution. Enhanced immune function may, however, also be costly for birds and the higher humoral immune responses in polluted areas may thus have negative effects on the birds' breeding performance and survival.     

Food diversity and niche-overlap of sympatric tits (Great Tit,Parus major,Blue Tit,Cyanistes caeruleus and Coal Tit Periparus ater) in the Hyrcanian Plain forests

Abstract

This study was carried out between early summer 1998 and late summer 1999 in the Hyrcanian Plain forests, the southern Caspian Sea woodlands. In total, the gizzard contents of 241 specimens of the Great Tit (Parus major Linnaeus, 1758), Blue Tit (Cyanistes caeruleus Linnaeus, 1758), and Coal Tit (Periparus ater (Linnaeus, 1758)) were collected so that the percentages of grit, plants and animal materials could be estimated and the plant and invertebrate species consumed identified. The data were tested by a series of Mann-Whitney U, ANOVA and Kruskal-Wallis tests. To survey niche overlap of food materials between species and season, the symmetrical overlap index developed by Pianka (1973) was used. The most important invertebrates (maximum of percentage per season) in the diet of P. major were Coleoptera (36.5%) and Lepidoptera (33.6%), for C. caeruleus Coleoptera (34.5%) and Araneae (31.2%), and for Pe. ater Coleoptera (38.2%) and Araneae (35.6%). The most important plants consumed by P. major were Sorghum halepense (68.7%) and Ficus carica (65.6%), by C. caeruleus were F. carica (67.5%) and Alnus glutinosa (39.4%), and by Pe. ater were Lonicera spp. (82.9%). There was a greater degree of overlap and competition for animal food, but plant feeding overlap among the three tit species was low. Two species, P. major and C. caeruleus, showed high feeding overlap for animal items (0.92), whereas Pe. ater and C. caeruleus had no significant feeding overlap. The Coal Tit had a greater tendency to forage for vegetable matter and P. major had a greater tendency to forage for animal matter. Analysis of gizzard contents of P. major and Pe. ater showed that, despite the high degree of feeding overlap, the extent of plant materials consumed indicated no significant competition between the two species. Because Pe. ater is present mostly in autumn and winter, this study cannot establish any degree of competition and food-niche overlap during the breeding season between this and other tit species.     

Aspects of the breeding biology of the Chinspot Batis Batis molitor in Acacia savanna in Swaziland

Some aspects of the breeding ecology of the Chinspot Batis (Batis molitor) were studied in Mlawula Nature Reserve, north-eastern Swaziland. Nests were predominantly built in thorny bushes or trees. Eggs were laid between 20 September and 2 January. However there was a definite peak in November, during which the majority of eggs were laid. Nesting success in the Chinspot Batis was over 30%, while fecundity was 0.65 fledglings/pair/annum. Except for a single occasion, pairs did not double brood unless breeding failed or fledged chicks disappeared. Replacement nests, however, were the norm where a previous nest had failed. Adult batises were observed to feed predominantly on caterpillars and moths. Observations at the nest confirmed that batises fed predominantly moths and caterpillars to their nestlings. The rate at which nestlings were fed depended on their age, older chicks being fed more frequently than younger ones. The correlation between feeding rate and nestling age, and between feeding rate and fledgling age, was significant. In contrast to nestlings, younger fledglings were fed at a higher rate than older ones.