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Effects of carotenoid availability during laying on reproduction in the blue tit.

Carotenoids are antioxidant pigments involved in several physiological processes and signalling in animals that cannot synthesise them and therefore must acquire them from food. We experimentally investigated the effects of carotenoid availability in the diet during egg laying on antioxidant deposition in egg yolk and the related effects on nestling condition, female body condition and parental investment in the blue tit (Parus caeruleus). Carotenoid supplementation of egg-laying females resulted in a significant increase in carotenoid concentration in egg yolk, but not in vitamin E or A concentration. There was no relationship between yellow plumage colour of adult females and carotenoid deposition in eggs, and no differential effect of feeding treatment depending on female colour. Nestlings from eggs laid by carotenoid supplemented females had longer tarsi, had faster development of the immune system as reflected by leukocyte concentration in blood, and grew brighter yellow feathers than nestlings from control females. However, nestlings from the two groups did not differ significantly in body mass, plasma antioxidants or plumage colour hue. At the time of chick rearing, carotenoid-fed females had increased plasma vitamin E levels compared to controls. However, females from the two treatment groups did not differ significantly in body condition or feeding rate. These results suggest that carotenoid availability is limiting during egg laying, and that females may have to balance the benefits of investing in egg quality against the potential costs of impairing their own future antioxidant protection. In addition, there may be considerable variation in carotenoid availability not only across seasons, but also among different stages of the breeding season.

Animals↗

Experimental removal of the male parent negatively affects growth and immunocompetence in nestling great tits.

Given the available empirical evidence on the benefits and costs associated with immune defence, a role for the immune system in the trade-off between current and future reproduction has been predicted. This hypothesis was studied in a free-living population of great tits (Parus major) by examining the effects of male removal on the immunocompetence, body condition, and recapture probability in the widowed females and their nestlings. Furthermore, we investigated whether growth and its relation to immunocompetence were affected in the nestlings. For a short-lived species such as the great tit, one could predict that widowed females will compensate for the lack of any male assistance in feeding of their chicks and that they consequently might jeopardize their own health. However, we did not find any negative effects of male removal on body mass or condition, nor on humoral immunocompetence against sheep red blood cells in the widowed females by the end of the feeding period. In contrast, we observed significantly reduced body mass and size as well as a reduced T-lymphocyte cell-mediated immune response (expressed as the thickness of the swelling to a subcutaneous injection with phytohemagglutinin) in the experimental nestlings compared to the control nestlings. In addition, the experimental nestlings showed a tendency for a reduced chance to be found breeding the following year. Furthermore, our results showed that in the experimental nestlings, which suffered from unfavourable growth conditions, tarsus length was inversely related to cell-mediated immunocompetence, whereas in control nestlings this relationship was significantly positive. The relationship between cell-mediated immunity and body condition was found to be significantly positive in the experimental nestlings while in control nestlings there was no relationship between both variables. The latter finding suggests different priorities of investment in body condition between different growth conditions.

Animals↗

Cross-fostering reveals an effect of spleen size and nest temperatures on immune responses in nestling European starlings.

Immunocompetence may be a good measure of offspring quality, however, factors affecting variation in immune responses are not clear. Research suggests that immune function can vary due to differences in genetics, development conditions and individual quality. Here, I examined factors affecting variation in immune response among nestling European starlings through a split-nest cross-fostering brood manipulation that included two important covariates: spleen size and nest temperatures. Immunocompetence was assessed via a cell-mediated immune response to phytohaemagglutinin (PHA). This paper provides the first direct evidence that individuals with large spleens also mount strong immune responses. Exposure to PHA did not cause splenomegaly, as there was no difference in spleen size between control birds and those injected with PHA. Offspring immune function was affected by common origin and by rearing environment, though rearing environment appeared to exert its influence only through nest temperatures. A comparison of the immune performance of siblings reared in their home nest versus those reared in other nests revealed a strong effect of maternal quality. As the difference in natal clutch size increased, the magnitude of the difference in immune performance between home-reared nestlings versus out-reared nestlings increased. Overall, nestling immune function appears to be determined by the combination of genetic, maternal and environmental effects.

Animals↗

Habitat selection by dispersing yellow-headed blackbirds: evidence of prospecting and the use of public information.

In migratory birds individuals may prospect for potential breeding sites months before they attempt to breed and should use the cues most predictive of future reproductive success when selecting a breeding site. However, what cues individuals use when prospecting and which cues are used in selecting a breeding site are unknown for most species. I investigated whether yellow-headed blackbirds (Xanthocephalus xanthocephalus) prospect for future breeding sites and whether they select breeding habitats based on food availability, male or female density, or the average number of young produced per female in the previous year. Although it is often assumed that migratory birds prospect for potential breeding sites at the end of the breeding season, I investigated this by recording all visits to sites early and late in the breeding season. I found that males and females who visited sites other than the site at which they bred were more likely to disperse than individuals only observed at the site where they bred, and that males and females were more likely to prospect late in the breeding season. Both food availability and density in year(x) were not predictive of the number of young per female in year(x+1); however, the number of young produced per female at a site in year(x) was predictive of the number of young per female in year(x+1). As expected, dispersers used the most informative cue, the number of young per female and moved to sites with a relatively high number of young per female. This study suggests that individuals prospect for potential breeding sites late in the breeding season when they can use information gathered from the reproductive success of other individuals (i.e., public information) to select a breeding site.

Animal Migration↗

Serum immunoglobulin G levels are positively related to reproductive performance in a long-lived seabird, the common tern (Sterna hirundo).

The evolution of longevity requires a low risk of mortality from extrinsic factors, relative to intrinsic factors, so that individuals that differentially invest in physiological self-maintenance and minimize their annual reproductive costs will maximize lifetime fitness through a prolonged reproductive lifespan. The trade-off between reproductive effort and self-maintenance, as measured by immune function, has been well documented in short-lived birds, but is difficult to demonstrate in long-lived birds. To assess self-maintenance in a long-lived seabird, we measured serum protein levels, including immunoglobulin G (IgG = IgY), in 30 breeding pairs of common terns (Sterna hirundo) and their first-hatched (A) chicks. Most parents were of known age from banding as hatchlings; our sample was selected to contrast young breeders (6-9 years) with very old birds (17-23 years). Body-mass of the parents declined by 5% during the chick-rearing period, while serum protein levels were stable. Serum IgG levels were higher in parents of offspring with faster growth rates, while IgG levels were lower in parents whose broods were reduced by starvation. A-chicks in broods of two had higher IgG levels than singleton chicks. Albumin levels were not related to reproductive performance. Thus, despite adequate statistical power, we could find no evidence for a trade-off between reproduction and self-maintenance in common terns, even in old age. The results are consistent with life-history predictions for long-lived vertebrates, in which selection favors sustained self-maintenance across the reproductive lifespan. The positive relationships between IgG levels and reproductive performance indicate that IgG can be used as an index of parental "quality."

Aging↗

Combined food and predator effects on songbird nest survival and annual reproductive success: results from a bi-factorial experiment.

Food and predators have traditionally been viewed as mutually exclusive alternatives when considering factors affecting animal populations. This has led to long controversies such as whether annual reproductive success in songbirds is primarily a function of food-restricted production or predator-induced loss. Recent studies on both birds and mammals suggest many of these controversies may be resolved by considering the combined effects of food and predators. We conducted a 2x2 manipulative food addition plus natural predator reduction experiment on song sparrows (Melospiza melodia) over three consecutive breeding seasons. Food and predators together affected partial clutch or brood loss, nest survival (total clutch or brood loss) and annual reproductive success. When combined, our two treatments reduced partial losses by more than expected if the effects of food and predators were independent and additive. Food and predators also interacted in their effects on nest survival since food addition significantly reduced the rate of nest predation. While annual reproductive success was highly correlated with nest predation (r2=0.71) the strength of this relationship was reinforced by the indirect effects of food addition on nest predation. A stepwise multiple regression showed that the residual variation in annual reproductive success was explained by food effects on the total number of eggs laid over the season and the combined effects of food and predators on partial losses noted above. We conclude that annual reproductive success in song sparrows is a function of both food-restricted production and predator-induced loss and indirect food and predator effects on both clutch and brood loss. We highlight the parallels between our results and those from a comparable bi-factorial experiment on mammals because we suspect combined food and predator effects are likely the norm in both birds and mammals.

Animals↗

Spatial heterogeneity in predator activity, nest survivorship, and nest-site selection in two forest thrushes.

The ability of prey to find and use predator-free space has far-reaching consequences for their persistence and interactions with their predators. We tested whether nest survivorship of the ground-nesting veery (Catharus fuscescens) and shrub-nesting wood thrush (Hylocichla mustelina) was related to the local absence of a major nest predator, the white-footed mouse (Peromyscus leucopus). Mouse-free space was defined by trap stations that failed to trap a mouse during the avian breeding season (approximately May through July). In addition, mouse activity was quantified at individual trap stations based on the number of captures during the same period (six 2.25-ha trapping grids, each containing 121 trap stations trapped repeatedly throughout the summer between 1998 and 2002.) Annual mouse-free space was correlated with other measures of mouse activity based on trapping data. Both mouse-free space and activity metrics were significantly related to annual rates of nest predation (i.e., nest daily mortality rate) in veery but not wood thrush. Likewise, mouse-free space and mouse activity within the nest neighborhood (approximately 30x30 m2 surrounding each nest) was significantly related to nest survivorship in veery but not wood thrush. More trap stations had consistently greater (hotspots) and lesser (coldspots) mouse activity than expected by chance, and veeries were significantly more likely to nest near stations that had below the grid-average trapping success. Our study thus documented significant spatial variability in predator activity and its relationship to nest predation and nest-site selection in a ground-nesting songbird.

Animals↗

Reproductive tactics under severe egg predation: an eider's dilemma.

Parental defence against predators may increase offspring survival but entail other costs. Egg predation is frequent early in the laying sequence of the common eider, which differs in this and in several other ways from most other waterfowl. We test the hypothesis that permanent presence at the nest from the second or third egg is an adaptation for reducing egg predation in eiders. Two other alternative hypotheses for lower predation at later nest stages are early predation loss of the most vulnerable nests and seasonal decrease in predation risk. Analyses of predation rates at the one-egg and later stages refute these two alternatives. Early nest attendance by eider females is estimated to increase clutch survival by about 20% in four-egg and 35% in five-egg clutches, albeit probably at a cost of smaller clutch size.

Animals↗

The reproductive phenology of an Amazonian ant species reflects the seasonal availability of its nest sites.

In saturated tropical ant assemblages, reproductive success depends on queens locating and competing for scarce nest sites. Little is known about how this process shapes the life histories of tropical ants. Here I investigate the relationship between nest site availability and an important life history trait, reproductive phenology, in the common Amazonian ant species Allomerus octoarticulatus. A. octoarticulatus is a plant-ant that nests in the hollow, swollen stem domatia on Cordia nodosa. I provide evidence that nest sites are limiting for A. octoarticulatus. Most queens produced by A. octoarticulatus colonies died before locating suitable host plants, and most queens that located hosts died before founding colonies, probably from intraspecific competition among queens for control of host plants. I further show that the reproductive phenology of A. octoarticulatus closely matches the seasonal availability of its nest sites, domatia-bearing C. nodosa saplings. Both the production and flight of A. octoarticulatus reproductives, and the number of C. nodosa saplings available for colonization by ants, peaked from March to May. There was correlative evidence that A. octoarticulatus colonies use temperature as a cue to synchronize their reproduction to the availability of C. nodosa saplings: both the production of reproductives by ant colonies and the number of C. nodosa saplings available for colonization were correlated with temperature, and not with rainfall. All of these results suggest that nest site limitation constrains the reproductive phenology of A. octoarticulatus.

Adaptation, Physiological↗

Speckled eggs: water-loss and incubation behaviour in the great tit Parus major.

Many small passerine birds worldwide lay white eggs speckled with red, brown and black protoporphyrin pigment spots (maculation). Unlike some patterns of avian eggshell pigmentation which clearly serve a crypsis or signalling function, the ubiquity of maculation among passerines suggests that its origins lie in another function, not specific to any particular ecological or behavioural group. Elsewhere, we have presented evidence that protoporphyrin pigments serve a structural function related to eggshell thickness and calcium availability: eggshell maculation in the great tit Parus major increases with decreasing soil calcium levels, pigments demarcate thinner areas of shell, and both the pigment intensity and distribution are related to shell thickness. Here we show that maculation also affects the rate of water loss from the egg during incubation (approximately Mass Loss per Day or MLD, which is critical to egg viability), but not that of unincubated eggs. We also demonstrate, both by observation and experiment, that the effect of female incubation behaviour on MLD compensates in some way for variation in egg characteristics, and that differences between females in the degree of such compensation are related to differences in clutch maculation. Our results suggest that, while a principal function of maculation in this species may be to strengthen the eggshell, it may also reduce eggshell permeability when large amounts of pigment are used, and that this necessitates a behavioural adjustment from the female during incubation. We discuss these findings and make further testable predictions from our model.

Animals↗

Do avian brood parasites eavesdrop on heterospecific sexual signals revealing host quality? A review of the evidence.

Eavesdropping can be defined as the extraction of information from the interactions between other individuals. It provides a relatively cheap way of gathering relevant information for fitness enhancement. Here, we propose that obligate avian brood parasites, which always lay their eggs in foreign nests of individuals of other species, may eavesdrop on their host sexual signals to locate nests of high quality individuals in which to lay their parasitic eggs. Sexual signal variation can honestly signal parental quality. Thus, by eavesdropping on sexual signals, parasites may select high quality foster parents for their own offspring. Such a use of sexual signals within host populations by brood parasites differs from signal exploitation theory that proposes that parasite only use signals to locate potential host independently from signaller quality. Here, we review the avian literature concerning host choice within a host species by obligate avian brood parasites and find evidence for host selection within individuals of a host species on the basis of cues potentially functioning as sexually selected traits, or at least revealing parental abilities. We have also found support for the existence of benefits linked to host selection by avian brood parasites. Finally, one study reported on the attenuation of a sexual ornament in host populations under strong pressure by brood parasites. Most of these findings have been interpreted as evidence for host selection by avian brood parasites based on the conspicuousness of sexual signals. We suggest, however, that these findings may in fact reveal eavesdropping on host signalling performance by brood parasites which would use the information extracted to choose the better individuals among conspecifics of a given host. This provides a new perspective for the study of host selection in obligate brood parasites, and raises interesting questions for the study of animal cognition that would deserve experimental studies.

Animals↗

Orangutan leaf-carrying for nest-building: toward unraveling cultural processes.

We report an empirical study on leaf-carrying, a newly discovered nest-building technique that involves collecting nest materials before reaching the nest site. We assessed whether leaf-carrying by rehabilitant orangutans on Kaja Island, Central Kalimantan, owes to cultural influences. Findings derive from ca 600 h observational data on nesting skills and nesting associations in Kaja's 42 resident rehabilitants, which yielded 355 nests and 125 leaf-carrying cases by 34 rehabilitants. Regional contrasts with 14 other communities (7 rehabilitant, 7 wild) indicated cultural influences on leaf-carrying on Kaja. Association data showed exceptional social learning opportunities for leaf-carrying on Kaja, with residents taking differential advantage of these opportunities as a function of development, experience, and social position. Juvenile males with basic nesting skills were most influenced by social input. Most (27) leaf-carriers had probably learned leaf-carrying when caged and 7 probably learned it on Kaja. Social priming was probably the main impetus to leaf-carrying on Kaja, by simply prompting observers to copy when leaf-carrying associates collected nesting materials, what they collected, and where they used their collected materials. Implications concern acquisition processes and ontogenetic schedules that orchestrate sets of features-needs or interests, cognitive abilities, social preferences-which enable cultural transmission.

Animals↗

A simple alternative to line transects of nests for estimating orangutan densities.

We conducted a validation of the line transect technique to estimate densities of orangutan (Pongo pygmaeus) nests in a Bornean swamp forest, and compared these results with density estimates based on nest counts in plots and on female home ranges. First, we examined the accuracy of the line transect method. We found that the densities based on a pass in both directions of two experienced pairs of observers was 27% below a combined sample based on transect walks by eight pairs of observers, suggesting that regular line-transect densities may seriously underestimate true densities. Second, we compared these results with those obtained by nest counts in 0.2-ha plots. This method produced an estimated 15.24 nests/ha, as compared to 10.0 and 10.9, respectively, by two experienced pairs of observers who walked a line transect in both directions. Third, we estimated orangutan densities based on female home range size and overlap and the proportion of females in the population, which produced a density of 4.25-4.5 individuals/km(2) . Converting nest densities into orangutan densities, using locally estimated parameters for nest production rate and proportion of nest builders in the population, we found that density estimates based on the line transect results of the most experienced pairs on a double pass were 2.82 and 3.08 orangutans/km(2), based on the combined line transect data are 4.04, and based on plot counts are 4.30. In this swamp forest, plot counts therefore give more accurate estimates than do line transects. We recommend that this new method be evaluated in other forest types as well.

Animals↗

Patterns of avian nest predators and a brood parasite among restored riparian habitats in agricultural watersheds.

In fragmented edge-dominated landscapes, nest predation and brood parasitism may reduce avian reproductive success and, ultimately, populations of some passerine species. In the fragmented agroecosystem of northwest Mississippi, placement of drop-pipe structures has been used as a restoration technique for abating gully erosion along stream banks. These actions have formed small herbaceous and woody habitat extensions into former agricultural lands. We quantified species relative abundances, species richness, and evenness of avian nest predators and a brood parasite within four categories of constructed habitat resulting from drop-pipe installation. Differences in the abundance of two nest predators, cotton mouse (Peromyscus gossypinus) and blue jay (Cyanocitta cristata), were observed among constructed habitats. However, relative abundances of other predators such as common grackle (Quiscalus quiscula), American crow (Corvus brachyrhynchos), and hispid cotton rat (Sigmodon hispidus), and the obligate brood parasite brown-headed cowbird (Molothrus ater) did not differ among four habitat categories. Although species richness, abundance, and evenness of potential nest predators were generally similar among the constructed habitats, predator species composition varied, suggesting that these habitats supported different predator communities. This difference is important because as each predator species is added to or deleted from the community, variation may occur in the framework of prey search methods, predator strategies, and potentially overall predation pressure. We suggest that land managers using drop-pipes as part of stream restoration projects allow for the development of the constructed habitat with the largest area and greatest vegetative structure.

Agriculture↗

Contaminant exposure of barn swallows nesting on Bayou d'Inde, Calcasieu Estuary, Louisiana, USA.

Current and historical point source discharges, storm water runoff, and accidental spills have contaminated the water, sediment, and biota within the Calcasieu Estuary in southwestern Louisiana. In 2003, barn swallow (Hirundo rustica) eggs and nestlings were collected beneath two bridges that cross Bayou d'Inde, the most contaminated waterway within the Calcasieu Estuary. Samples were also collected from a bridge over Bayou Teche, a reference site in south central Louisiana. Polychlorinated biphenyl (PCB) concentrations in barn swallow eggs and nestlings were significantly higher at the downstream site on Bayou d'Inde (2.8 micro g/g PCBs in eggs and 1.5 micro g/g PCBs in nestlings) than at the other two sites (< 0.2 micro g/g PCBs in eggs and nestlings at both sites). Ethoxyresorufin-O-dealkylase activity in nestling livers was significantly higher at the downstream site on Bayou d'Inde (50 pmol/min/mg) compared to the other two locations (24 pmol/min/mg, each), probably because of exposure to PCBs. Polychlorinated dibenzo-p-dioxin and polychlorinated dibenzofuran concentrations in eggs and polycyclic aromatic hydrocarbons in nestlings were at background concentrations at all sites. Trace element concentrations in barn swallow eggs and nestling livers were at background levels and did not differ among the three sites. A biomarker of DNA damage did not differ among sites.

Animals↗

Mercury and growth of tree swallows at Acadia National Park, and at Orono, Maine, USA.

In 1997 and 1998 we weighed nestling tree swallows (Tachycineta bicolor) and measured selected body components at two colonies: Acadia National Park on Mt. Desert Island, and at Orono, ME. We used differences in mean growth variables among individual nestlings to evaluate differences between colonies, years, and amount of total mercury (THg) in carcasses and methyl mercury (MeHg) in feathers. We marked nestlings on the day hatched and measured body components every day in 1997 and every other day in 1998 until nestlings fledged. We calculated linear growth rates and asymptotic means as appropriate. In 1998, linear growth rate of weight was higher at Acadia than at Orono, but not different in 1997. We detected no mean differences in asymptotic mean weight of nestlings between colonies or years. In 1997, mean linear growth rates of the wing (chord), tail, tarsus, and mandible were higher at Acadia than at Orono. The amount of MeHg in feathers was associated with a lower linear growth rate of weight during early age (2-10 days), but asymptotic mean weight during days 11-16 was not different. No effect on linear growth of tail feathers or wing was associated with the amount of MeHg in feathers or THg in carcasses. Fledgling tree swallows that survive to migrate, however, will leave Maine with substantial concentrations of Hg in their tissues.

Animals↗

Mercury in tree swallow food, eggs, bodies, and feathers at Acadia National Park, Maine, and an EPA superfund site, Ayer, Massachusetts.

We monitored nest boxes during 1997-1999 at Acadia National Park, Mt. Desert Island, ME and at an old-field site in Orono, ME to determine mercury (Hg) uptake in tree swallow (Tachycineta bicolor) eggs, tissues, and food boluses. Also, in 1998-1999 we monitored nest boxes at Grove Pond and Plow Shop Pond at a U.S. Environmental Protection Agency Superfund site in Ayer, MA. We recorded breeding success at all locations. On average among locations, total mercury (THg) biomagnified 2 to 4-fold from food to eggs and 9 to 18-fold from food to feathers. These are minimum values because the proportion of transferable methyl mercury (MeHg) of the THg in insects varies (i.e., 35%-95% of THg) in food boluses. THg was highest in food boluses at Aunt Betty Pond at Acadia, whereas THg in eggs was highest at the Superfund site. A few eggs from nests at each of these locations exceeded the threshold (i.e., 800-1,000 ng/g, wet wt.) of embryotoxicity established for Hg. Hatching success was 88.9% to 100% among locations, but five eggs failed to hatch from 4 of the 11 clutches in which an egg exceeded this threshold. MeHg in feathers was highest in tree swallows at Aunt Betty Pond and the concentration of THg in bodies was related to the concentration in feathers. Transfer of an average of 80%-92% of the Hg in bodies to feathers may have enhanced nestling survival. Residues of Hg in tissues of tree swallows in the Northeast seem higher than those of the Midwest.

Animals↗

Nest liquid resources of several cavity nesting bees in the genus Centris and the identification of a preservative, levulinic acid.

Twig-nesting species of bees in the genus Centris including C. bicornuta, C. analis, C. vittata, and C. nitida, found in the dry forest of Guanacaste Province of Costa Rica, provision their nests with pollen and nectar, rather than pollen and oil as reported for other Centris species. The liquid contents of the nests of these four species were found to contain sugars including 66-75% fructose, 25-33% glucose, and a trace of sucrose. The sugar concentration averaged 47.2%, slightly higher than most flower nectars. No tri-, di-, or monoglycerides, the main components of the flower oil of Byrsonima crassifolia, were detected in the nest provisions. Although these four Centris species are also known to collect oil from B. crassifolia, the oil appears to be used for activities other than nest provisioning. The liquid nest contents did have a slight goat-like odor, suggesting the presence of short-chain fatty acids, and were found to contain a small amount (less than 1%) of three fatty acids. Two of these, butanoic and octanoic acid, were found in trace amounts and are responsible for the goat-like odor. A third was identified as levulinic acid, which made up about 99% of the nest fatty acid contents. This fatty acid had little odor, but may be important as a fungicidal agent. Attempts to determine the source of the fatty acids, were not successful.

Animals↗