Effect of Feathers as Nest Insulation on Incubation Behavior and Reproductive Performance of Tree Swallows (\u3cem\u3eTachycineta bicolor\u3c/em\u3e)

Many species of birds line their nests with feathers, presumably because of the insulative qualities of feathers and because feathers may act as a barrier between nest parasites and nestlings. In 1993, we experimentally examined the role of feathers as nest insulation on the incubation behavior, nestling growth, and reproductive performance of Tree Swallows (Tachycineta bicolor) nesting in boxes in western Michigan. There were no significant differences between the incubation rhythms of females with experimental nests (i.e. no feathers) and females with control nests (i.e. with feathers). Nestlings that were reared in control nests had significantly longer right tarsi and right wing chords; their masses were significantly greater than nestlings reared in experimental nests. In addition, nested analyses of variance indicated that both female age class (i.e. second year, after second year, or after hatching year) and the brood within which a nestling was reared had significant effects on nestling growth until nestling day 12. Whether an individual nestling was infected with ectoparasites was independent of whether it was reared in an experimental or control nest. Nest insulation affected reproductive performance: females with experimental nests had significantly longer incubation periods and produced significantly fewer fledglings than did females with control nests. These results suggest that nest insulation may be an important factor influencing incubation behavior, nestling growth, and reproductive performance of Tree Swallows in western Michigan

Supplementary feeding increases nestling feather corticosterone early in the breeding season in house sparrows

Several studies on birds have proposed that a lack of invertebrate prey in urbanized areas could be the main cause for generally lower levels of breeding success compared to rural habitats. Previous work on house sparrows Passer domesticus found that supplemental feeding in urbanized areas increased breeding success but did not contribute to population growth. Here, we hypothesize that supplementary feeding allows house sparrows to achieve higher breeding success but at the cost of lower nestling quality. As abundant food supplies may permit both high-and low-quality nestlings to survive, we also predict that within-brood variation in proxies of nestling quality would be larger for supplemental food broods than for unfed broods. As proxies of nestling quality, we considered feather corticosterone (CORTf), body condition (scaled mass index, SMI), and tarsus-based fluctuating asymmetry (FA). Our hypothesis was only partially supported as we did not find an overall effect of food supplementation on FA or SMI. Rather, food supplementation affected nestling phenotype only early in the breeding season in terms of elevated CORTf levels and a tendency for more variable within-brood CORTf and FA. Early food supplemented nests therefore seemed to include at least some nestlings that faced increased stressors during development, possibly due to harsher environmental (e.g., related to food and temperature) conditions early in the breeding season that would increase sibling competition, especially in larger broods. The fact that CORTf was positively, rather than inversely, related to nestling SMI further suggests that factors influencing CORTf and SMI are likely operating over different periods or, alternatively, that nestlings in good nutritional condition also invest in high-quality feathers

No experimental evidence for local competition in the nestling phase as a driving force for density-dependent avian clutch size

1. In birds, local competition for food between pairs during the nestling phase may affect nestling growth and survival. A decrease in clutch size with an increase in breeding density could be an adaptive response to this competition. To investigate whether breeding density causally affected the clutch size of great tits (Parus major), we manipulated breeding density in three out of eight study plots by increasing nest-box densities. We expected clutch size in these plots to be reduced compared to that in control plots. 2. We analysed both the effects of variation in annual mean density (between-year comparisons) and experimental density (within-year comparison between plots) on clutch size variation, the occurrence of second broods and nestling growth. We examined within-female variation in clutch size to determine whether individual responses explain the variation over years. 3. Over the 11 years, population breeding density increased (from 0·33 to 0·50 pairs ha–1) while clutch size and the occurrence of second broods decreased (respectively from 10·0 to 8·5 eggs and from 0·39 to 0·05), consistent with a negative density-dependent effect for the whole population. Nestling growth showed a declining but nonsignificant trend over years. 4. The decline in population clutch size over years was primarily explained by changes occurring within individuals rather than selective disappearance of individuals laying large clutches. 5. Within years, breeding density differed significantly between manipulated plots (0·16 pairs ha–1 vs. 0·77 pairs ha–1) but clutch size, occurrence of second broods and nestling growth were not affected by the experimental treatment, resulting in a discrepancy between the effects of experimental and annual variation in density on reproduction. 6. We discuss two hypotheses that could explain this discrepancy: (i) the decline in breeding performance over time was not due to density, but resulted from other, unknown factors. (ii) Density did cause the decline in breeding performance, but this was not due to local competition in the nestling phase. Instead, we suggest that competition acting in a different phase (e.g. before egg laying or after fledgling) was responsible for the density effect on clutch size among years.

A guide to nestling development and aging in altricial passerines

Nestling growth and development studies have been a topic of interest for a greater part of the last century (Sutton 1935, Walkinshaw 1948) and continue to be of interest today. This is not surprising since studies on nestling growth can provide a wealth of biological information that has larger implications for avian management and conservation. Despite this history of studying nestling development, basic information is still limited or absent for many species. Many questions remain unanswered, and contradictory conclusions are often found in the literature (Starck and Ricklefs 1998a). Therefore, much information on aging and development can still be gained from studying the development patterns of similar species and from comparative studies, across avian orders (Minea et al. 1982, Saunders and Hansen 1989, Carsson and Hörnfeldt 1993). Additionally, nestling growth studies can yield insight into the effects of different nesting strategies on productivity (O’Connor 1978), as well as the impacts of parental effort and environmental variables on fitness (Ross 1980, Ricklefs and Peters 1981, Magrath 1991). Since low reproductive success may play a significant role in the declines of many North American passerines (Sherry and Holmes 1992, Ballard et al. 2003), a better understanding of the factors that influence reproductive success is a vital component of avian conservation (Martin 1992). Data on nestling aging can be used to improve nest survival estimates (Dinsmore 2002, Nur et al. 2004), providing information that can be used to more precisely age nests (Pinkowski 1975, Podlesack and Blem 2002), (Jones and Geupel 2007). Indeed, the relatively short time period young spend developing in the nest is a critical part of a bird’s life cycle and a nestling’s developmental path can affect its survival to independence, its survival as an adult, and its future reproductive success

Maximal Displacement for Bridges of Random Walks in a Random Environment

It is well known that the distribution of simple random walks on $\bf{Z}$ conditioned on returning to the origin after $2n$ steps does not depend on $p= P(S_1 = 1)$, the probability of moving to the right. Moreover, conditioned on $\{S_{2n}=0\}$ the maximal displacement $\max_{k\leq 2n} |S_k|$ converges in distribution when scaled by $\sqrt{n}$ (diffusive scaling). We consider the analogous problem for transient random walks in random environments on $\bf{Z}$. We show that under the quenched law $P_\omega$ (conditioned on the environment $\omega$), the maximal displacement of the random walk when conditioned to return to the origin at time $2n$ is no longer necessarily of the order $\sqrt{n}$. If the environment is nestling (both positive and negative local drifts exist) then the maximal displacement conditioned on returning to the origin at time $2n$ is of order $n^{\kappa/(\kappa+1)}$, where the constant $\kappa>0$ depends on the law on environment. On the other hand, if the environment is marginally nestling or non-nestling (only non-negative local drifts) then the maximal displacement conditioned on returning to the origin at time $2n$ is at least $n^{1-\varepsilon}$ and at most $n/(\ln n)^{2-\varepsilon}$ for any $\varepsilon>0$. As a consequence of our proofs, we obtain precise rates of decay for $P_\omega(X_{2n}=0)$. In particular, for certain non-nestling environments we show that $P_\omega(X_{2n}=0) = \exp\{-Cn -C'n/(\ln n)^2 + o(n/(\ln n)^2) \}$ with explicit constants $C,C'>0$.Comment: Revised version, 19 pages, 1 figure To appear in: AIHP Prob. & Sta

Prothonotary warbler nestling growth and condition inresponse to variation in aquatic and terrestrial preyavailability

Aquatic prey subsidies entering terrestrial habitats are well documented, but little is known about the degree to which these resources provide fitness benefits to riparian consumers. Riparian species take advantage of seasonal pulses of both terrestrial and aquatic prey, although aquatic resources are often over-looked in studies of how diet influences the reproductive ecology of these organisms. Ideally, the timing of resource pulses should occur at the time of highest reproductive demand. This study investigates the availability of aquatic(mayfly) and terrestrial (caterpillar) prey resources as well as the nestling diet of the prothonotary warbler (Protonotaria citrea) at two sites along the lower James River in Virginia during the 2014 breeding season. We found large differences in availability of prey items between the two sites, with one having significantly higher mayfly availability. Nestling diet was generally reflective of prey availability, and nestlings had faster mean growth rates at the site with higher aquatic prey availability. Terrestrial prey were fed more readily at the site with lower aquatic prey availability, and at this site, nestlings fed mayflies had higher mean growth rates than nestlings fed only terrestrial prey. Our results suggest that aquatic subsidies are an important resource for nestling birds and are crucial to understanding the breeding ecology of riparian species

What makes a host profitable? Parasites balance host nutritive resources against immunity

Numerous host qualities can modulate parasite fitness, and among these, host nutritive resources and immunity are of prime importance. Indeed, parasite fitness increases with the amount of nutritive resources extracted from the host body and decreases with host immune response. To maximize fitness, parasites have therefore to balance these two host components. Yet, because host nutritive resources and immunity both increase with host body condition, it is unclear whether parasites perform better on hosts in prime, intermediate, or poor condition. We investigated blood meal size and survival of the ectoparasitic louse fly Crataerina melbae in relation to body condition and cutaneous immune response of their Alpine swift (Apus melba) nestling hosts. Louse flies took a smaller blood meal and lived a shorter period of time when feeding on nestlings that were experimentally food deprived or had their cutaneous immune response boosted with methionine. Consistent with these results, louse fly survival was the highest when feeding on nonexperimental nestlings in intermediate body condition. Our findings emphasize that although hosts in poor condition had a reduced immunocompetence, parasites may have avoided them because individuals in poor condition did not provide adequate resources. These findings highlight the fact that giving host immunocompetence primary consideration can result in a biased appraisal of host-parasite interactions

Integrated behavioural and stable isotope data reveal altered diet linked to low breeding success in urban-dwelling blue tits (Cyanistes caeruleus)

Animals often show reduced reproductive success in urban compared to adjacent natural areas. The lower availability and quality of natural food in cities is suggested as one key limiting factor. However, only few studies have provided conclusive support by simultaneously assessing food availability, diet and fitness. We consolidate this evidence by taking a holistic approach, comparing blue tits breeding in forest, suburban and urban areas. We (a) assessed arthropod availability, (b) investigated parental provisioning behaviour, (c) inferred diet through stable isotope analysis, and (d) measured reproductive success. At the urban site, we found a significant reduction in caterpillar availability, the main food source of blue tits, and consequently urban tits fed their offspring with fewer caterpillars than forest and suburban birds. Stable isotope analysis confirmed that diet in the urban area was fundamentally different than in the other sites. Reproductive success was lower in both urban and suburban sites compared to the forest site, and was positively associated with volume of provisioned caterpillars. Our findings provide strong integrative evidence that urban blue tit nestlings are not receiving a suitable diet, and this may be an important limiting factor for urban populations of this and potentially many other species

Averaged large deviations for random walk in a random environment

In his 2003 paper, Varadhan proves the averaged large deviation principle for the mean velocity of a particle taking a nearest-neighbor random walk in a uniformly elliptic i.i.d. environment on $\mathbb{Z}^d$ with $d\geq1$, and gives a variational formula for the corresponding rate function $I_a$. Under Sznitman's transience condition (T), we show that $I_a$ is strictly convex and analytic on a non-empty open set $\mathcal{A}$, and that the true velocity of the particle is an element (resp. in the boundary) of $\mathcal{A}$ when the walk is non-nestling (resp. nestling). We then identify the unique minimizer of Varadhan's variational formula at any velocity in $\mathcal{A}$.Comment: 14 pages. In this revised version, I state and prove all of the results under Sznitman's (T) condition instead of Kalikow's condition. Also, I rewrote many parts of Section 1, streamlined some of the proofs in Section 2, fixed some typos, and improved the wording here and there. Accepted for publication in Annales de l'Institut Henri Poincar

Development of the Innate Immune Response in NestlingTree Swallows (Tachycineta bicolor)

The innate immune system provides an immediate, short term, first line of defense from pathogens; its appearance early in development in vertebrates is evidence of its critical importance. Even so, few studies have investigated the development of the immune response as juveniles transition into adults. Ultimately, the ability to respond to pathogens confers fitness benefits in terms of health, survival, and reproductive success, and it follows that functions such as rapid growth cannot be fully met simultaneously since energy is a limiting resource. As a result, defense mechanisms are compromised at an early age due to energy allocation to rapid growth; therefore, immunity should increase as individuals mature. I studied the development of innate immunity in nestling Tree Swallows using microbicidal assays which were conducted in vitro to assess the ability of the immune system to kill E. coli via lysis. This research may provide insight into patterns of disease susceptibility, which in turn influence evolutionary fitness and population dynamics