Immune-challenged house wren broods differ in the relative strengths of their responses among different axes of the immune system

Single components of the immune system are widely used to assess immune function in free-living vertebrates. However, as different immunological components are triggered by different types of threats and may be regulated independently, there is little reason to assume that they should respond similarly if challenged. We investigated whether three commonly assayed immune responses, cutaneous immune activity (phytohaemagglutinin assay), antibody response (tetanus toxoid immunization), and plasma bactericidal activity (Escherichia coli killing) are positively related in nestling house wrens (Troglodytes aedon). Multivariate analysis revealed significant differences in overall immune responsiveness among broods (i.e. nests), primarily attributable to differences in plasma bactericidal activity. Among broods, humoral immune response was negatively related to cutaneous immune activity and positively related to plasma bactericidal activity. We found no significant relationships among these measures of immunity among individual nestlings within broods. Our results suggest that different broods (i.e. families) invest differentially in the various branches of the immune system. Further study is needed to characterize the roles of maternal, genetic and environmental effects on the expression of this physiological bias.     

Observer variation in estimates of Meadow Pipit Anthus pratensis and Skylark Alauda arvensis abundance on moorland

Capsule Counts least susceptible to observer effects were those within 25 m of a transect and the total number along a transect, with observer effects greater for distance sampling.     

The usefulness of sensitivity analysis for predicting the effects of cat predation on the population dynamics of their avian prey

Sensitivity analyses of population projection matrix (PPM) models are often used to identify life-history perturbations that will most influence a population's future dynamics. Sensitivities are linear extrapolations of the relationship between a population's growth rate and perturbations to its demographic parameters. Their effectiveness depends on the validity of the assumption of linearity. Here we assess whether sensitivity analysis is an appropriate tool to investigate the effect of predation by cats on the population growth rates of their avian prey. We assess whether predation by cats leads to non-linear effects on population growth and compare population growth rates predicted by sensitivity analysis with those predicted by a non-linear simulation. For a two-stage, age-classified House Sparrow Passer domesticus PPM slight non-linearity arose when PPM elements were perturbed, but perturbation to the vital rates underlying the matrix elements had a linear impact on population growth rate. We found a similar effect with a slightly larger three-stage, age-classified PPM for a Winter Wren Troglodytes troglodytes population perturbed by cat predation. For some avian species, predation by cats may cause linear or only slightly nonlinear impacts on population growth rates. For these species, sensitivity analysis appears to be a useful conservation tool. However, further work on multiple perturbations to avian prey species with more complicated life histories and higher-dimension PPM models is required.     

Extra-pair young in house wren broods are more likely to be male than female

Sex-allocation theory predicts that females should preferentially produce offspring of the sex with greater fitness potential. In socially monogamous animal species, extra-pair mating often increases the variance in fitness of sons relative to daughters. Thus, in situations where offspring sired by a female''s extra-pair mate(s) will typically have greater fitness potential than offspring sired by the within-pair mate, sex-allocation theory predicts that females will bias the sex of offspring sired by extra-pair mates towards male. We examined the relationship between offspring sex and paternity over six breeding seasons in an Illinois population of the house wren (Troglodytes aedon), a cavity-nesting songbird. Out of the 2345 nestlings that had both sex and paternity assigned, 350 (15%) were sired by extra-pair males. The sex ratio of extra-pair offspring, 0.534, was significantly greater than the sex ratio of within-pair offspring, 0.492, representing an increase of 8.5 per cent in the proportion of sons produced. To our knowledge, this is the first confirmed report of female birds increasing their production of sons in association with extra-pair fertilization. Our results are consistent with the oft-mentioned hypothesis that females engage in extra-pair mating to increase offspring quality.     

Sexing Wrens Troglodytes troglodytes indigenus using morphological measurements and discriminant analysis

Within the breeding population of a deciduous woodland, male Wrens averaged slightly larger than females in wing length, foot length, head plus bill length and body mass. First-year and older birds were similar for three of these morphometric variables, but the first-year group were shorter-winged than the older birds (by 3%). Despite this, wing length still proved to be a useful discriminator of sex when the data were pooled for all ages (74% of males exceeding the maximum female value of 49.8 mm). A discriminant function utilizing wing length and head plus bill length correctly sexed 96% of individuals in the sample from which it was derived (n = 49 males, 36 females). This function was converted into a predictive equation for the probability of an unknown case being male, and a plot of wing length against head plus bill length along with probability contours which enabled the sexing of unknown cases without the need for calculations. Measurements of the morphological variables investigated were found to be highly repeatable for a single worker, with the exception of body mass which differed substantially between the sexes but which was only moderately repeatable. The methods provided should prove valuable for sexing British mainland Troglodytes troglodytes indigenus, and probably also Western European T. t. troglodytes, but should not be applied to the larger, offshore island races.     

DEVELOPMENT OF VOCALISATIONS IN NESTLING AND FLEDGLING HOUSE WRENS IN NATURAL POPULATIONS

ABSTRACT

We conducted a field study of the ontogeny of vocal signals in the house wren Troglodytes aedon during the nestling and fledgling phases of life. We did spectrographic analyses and quantification of the developmental changes that occurred in the acoustic features of the vocalisations. Evidence of progress to adult-like vocal patterns was of two types. First, nestling calls changed into a harsh-sounding call that resembles the adult chatter call, functionally a warning call. Second, fledglings also uttered subsong, and these vocalisations were similar to notes typical of adult male song. When the vocalisations produced by developing young were broken down into their constituent vocal features, we found that the time course of development was not strictly linear. Instead of a unidirectional change through the course of nestling and fledgling life, the trajectories of the vocal features fluctuated through time and sometimes exhibited abrupt changes. These sudden shifts occurred during nestling life as well as at the time of fledging. We speculate on the possible causes of these abrupt transitions. Changes in acoustic features upon fledging appear to be linked to new social functions.     

Increased cuckoldry as a cost of breeding late for male house wrens (Troglodytes aedon)

One factor hypothesized to influence the reproductive behaviorof individuals is the degree to which reproductive effortsare synchronized with others in the population. We asked whetherthe timing of a pair's breeding cycle, relative to cycles ofpairs on neighboring territories, affected rates of extrapairmating over 2 years in a Wyoming population of house wrens (Troglodytes aedon). Extrapair young (identified using 5 microsatelliteloci) occurred in 74% of nests of 19 pairs whose cycles began later than cycles of one or more neighbors compared to only26% of nests of 27 pairs whose cycles began earlier than, orsimultaneously with, cycles of all neighbors. Extrapair offspringoccurred in 65% of 17 nests belonging to males who initiallysettled and began nesting early relative to neighbors but who were forced to renest late after we removed their first mates.Rates of cuckoldry were not significantly different for forced-lateand naturally late males. Our experimental approach controlledfor possible effects of male quality, clearly demonstratingan effect of timing of breeding on extrapair mating activity.     

Female house wrens (Troglodytes aedon) increase the size, but not immunocompetence, of their offspring through extra-pair mating

House wrens are typically socially monogamous, but frequently engage in extra-pair matings leading to multisired broods. Because females do not appear to acquire direct material benefits from their extra-pair mates, we tested the hypothesis that female house wrens derive indirect genetic benefits, such as enhanced immunocompetence (cutaneous immune activity, humoral immunity, and plasma bactericidal activity) and condition (size and haematoserological traits) for their offspring, by mating polyandrously. We predicted that extra-pair young (EPY) should show greater immune responsiveness and better body condition than their within-pair maternal half-siblings (WPY). Contrary to our prediction, WPY had higher cutaneous immune activity than their EPY brood-mates in two of three years, and EPY and WPY did not differ in measures of innate and humoral immunity. WPY also had higher albumin to γ-globulin ratios than EPY; however, they were not in better condition based on other measures. EPY had consistently longer tarsi (a measure of long-bone size) than their WPY half-siblings, suggesting that females engage in extra-pair copulations with larger males. The benefits of large structural size in the study population is unknown, but based on evidence from other passerines, we suggest that structural size may be an important fitness-related trait in house wrens. We conclude that our results are not consistent with the hypothesis that females gain immune-related benefits for their offspring by engaging in extra-pair matings. Further study of the fitness consequences of differences in tarsus length is needed to determine whether females acquire size-related benefits for their offspring from extra-pair mates.     

The effect of an extreme winter on per-capita growth rates in some resident bird populations: an example of r-selection?

Among the breeding birds of an English oakwood studied over a period of 27 years, five species averaged more than 10 pairs. Two of these populations, the robin and especially the wren, crashed after the extreme winter 1962–1963, whereas the populations of three other (group II) species did not crash. In stepwise multiple regressions between per-capita growth rate, r , and four winter temperatures, population size and year, population size was selected in all species. Winter temperatures were selected only in robin and wren. The residual variation of r which remained after the selected variables were used in a multiple regression equation was randomly distributed around zero in the three group II species. In the robin and wren, however, the residual values for the years following the cold winter were significantly higher compared to those in the years before it. This suggests that something changed either in the environment or in the birds. Robin residuals after the cold winter returned towards zero, whereas wren residuals remained at the new high level. I argue that these results can best be explained through assuming that catastrophic winter mortality was non-random and could have caused micro-evolutionary changes in life-history traits.