Patricia Brekke

Sensitive males: inbreeding depression in an endangered bird

Attempts to conserve threatened species by establishing new populations via reintroduction are controversial. Theory predicts that genetic bottlenecks result in increased mating between relatives and inbreeding depression. However, few studies of wild sourced reintroductions have carefully examined these genetic consequences. Our study assesses inbreeding and inbreeding depression in a free-living reintroduced population of an endangered New Zealand bird, the hihi (Notiomystis cincta). Using molecular sexing and marker-based inbreeding coefficients estimated from 19 autosomal microsatellite loci, we show that (i) inbreeding depresses offspring survival, (ii) male embryos are more inbred on average than female embryos, (iii) the effect of inbreeding depression is male-biased and (iv) this population has a substantial genetic load. Male susceptibility to inbreeding during embryo and nestling development may be due to size dimorphism, resulting in faster growth rates and more stressful development for male embryos and nestlings compared with females. This work highlights the effects of inbreeding at early life-history stages and the repercussions for the long-term population viability of threatened species.

High genetic diversity in the remnant island population of hihi and the genetic consequences of re-introduction

The maintenance of genetic diversity is thought to be fundamental for the conservation of threatened species. It is therefore important to understand how genetic diversity is affected by the re‐introduction of threatened species. We use establishment history and genetic data from the remnant and re‐introduced populations of a New Zealand endemic bird, the hihi Notiomystis cincta, to understand genetic diversity loss and quantify the genetic effects of re‐introduction. Our data do not support any recent bottleneck events in the remnant population. Furthermore, all genetic diversity measures indicate the remnant hihi population has retained high levels of genetic diversity relative to other New Zealand avifauna with similar histories of decline. Genetic diversity (NA, alleles per locus, allelic richness, FIS and HS) did not significantly decrease in new hihi populations founded through re‐introduction when compared to their source populations, except in the Kapiti Island population (allelic richness and HS) which had very slow post‐re‐introduction population growth. The Ne/Nc ratio in the remnant population was high, but decreased in first‐level re‐introductions, which together with significant genetic differentiation between populations (FST & Fisher’s exact tests) suggest that extant populations are diverging as a result of founder effects and drift. Importantly, simulations of future allele loss predict that the number of alleles lost will be higher in populations with a slow population growth, fewer founding individuals and with nonrandom mating. Interestingly, this species has very high levels of extra‐pair paternity which may reduce reproductive variance by allowing social and floater males to reproduce a life history trait that together with a large remnant population size may help maintain higher levels of genetic diversity than expected.

ESTIMATION OF PARAMETERS OF INBREEDING AND GENETIC DRIFT IN POPULATIONS WITH OVERLAPPING GENERATIONS

Many long‐lived plant and animal species have nondiscrete overlapping generations. Although numerous models have been developed to predict the effective sizes (Ne) of populations with overlapping generations, they are extremely difficult to apply to natural populations because of the large array of unknown and elusive life‐table parameters involved. Unfortunately, little work has been done to estimate the Ne of populations with overlapping generations from marker data, in sharp contrast to the situation of populations with discrete generations for which quite a few estimators are available. In this study, we propose an estimator (EPA, estimator by parentage assignments) of the current Ne of populations with overlapping generations, using the sex, age, and multilocus genotype information of a single sample of individuals taken at random from the population. Simulations show that EPA provides unbiased and accurate estimates of Ne under realistic sampling and genotyping effort. Additionally, it yields estimates of other interesting parameters such as generation interval, the variances and covariances of lifetime family size, effective number of breeders of each age class, and life‐table variables. Data from wild populations of baboons and hihi (stitchbird) were analyzed by EPA to demonstrate the use of the estimator in practical sampling and genotyping situations.

Maternally invested carotenoids compensate costly ectoparasitism in the hihi

Dietary ingested carotenoid biomolecules have been linked to both improved health and immunity in nestling birds. Here, we test whether maternally invested egg carotenoids can offset the cost of parasitism in developing nestling hihi (Notiomystis cincta) from the bloodsucking mite (Ornithonyssus bursa). Our results reveal clear negative effects of parasitism on nestlings, and that maternally derived carotenoids compensate this cost, resulting in growth parameters and ultimate mass achieved being similar to nonparasitized young. Our results offer an unique example of a direct positive relationship between enhanced maternal investment of carotenoids and an ability to cope with a specific and costly parasite in young birds. As O. bursa infestations reduce population viability in hihi, our findings also highlight the importance of key nutritional resources for endangered bird populations to better cope with common parasite infestations.

Characterization of microsatellite loci In The hihi Notiomystis cincta (Notiomystidae, Aves)

We have characterized 20 polymorphic microsatellite loci in the hihi Notiomystis cincta. Loci were identified by testing loci originally isolated in other avian species and by isolating new microsatellites from a hihi genomic library. These loci were characterized in unrelated hihi from a single population on Tiritiri Matangi Island (n = 98). Each locus displayed between two and 10 alleles, and observed heterozygosities ranged between 0.29 and 0.91. Nineteen of the 20 polymorphic loci could be assigned a chromosome location in the zebra finch, Taeniopygia guttata genome based on sequence homology.

Sexually selected dichromatism in the hihi Notiomystis cincta: multiple colours for multiple receivers

Why do some bird species show dramatic sexual dichromatism in their plumage? Sexual selection is the most common answer to this question. However, other competing explanations mean it is unwise to assume that all sexual dichromatism has evolved by this mechanism. Even if sexual selection is involved, further work is necessary to determine whether dichromatism results from competition amongst rival males, or by female choice for attractive traits, or both. Here, we test whether sexually dichromatic hihi (Notiomystis cincta) plumage is currently under sexual selection, with detailed behavioural and genetic analyses of a free‐living island population. Bateman gradients measured for males and females reveal the potential for sexual selection, whilst selection gradients, relating reproductive success to specific colourful traits, show that there is stabilizing selection on white ear tuft length in males. By correlating colourful male plumage with different components of reproductive success, we show that properties of yellow plumage are most likely a product of male–male competition, whilst properties of the black and white plumage are an outcome of both male–male competition and female choice. Male plumage therefore potentially signals to multiple receivers (rival males and potential mates), and this may explain the multicoloured appearance of one of the most strikingly dichromatic species in New Zealand.

Determinants of male floating behaviour and floater reproduction in a threatened population of the hihi (Notiomystis cincta).

Floating males are usually thought of as nonbreeders. However, some floating individuals are able to reproduce through extra‐pair copulations. Floater reproductive success can impact breeders’ sex ratio, reproductive variance, multiple paternity and inbreeding, particularly in small populations. Changes in reproductive variance alter the rate of genetic drift and loss of genetic diversity. Therefore, genetic management of threatened species requires an understanding of floater reproduction and determinants of floating behaviour to effectively conserve species. Here, we used a pedigreed, free‐living population of the endangered New Zealand hihi (Notiomystis cincta) to assess variance in male reproductive success and test the genetic (inbreeding and heritability) and conditional (age and size) factors that influence floater behaviour and reproduction. Floater reproduction is common in this species. However, floater individuals have lower reproductive success and variance in reproductive success than territorial males (total and extra‐pair fledglings), so their relative impact on the populations reproductive performance is low. Whether an individual becomes a floater, and if so then how successful they are, is determined mainly by individual age (young and old) and to lesser extents male size (small) and inbreeding level (inbred). Floating males have a small, but important role in population reproduction and persistence of threatened populations.

The Evolutionary Biology, Ecology and Epidemiology of Coccidia of Passerine Birds

Coccidia are intracellular parasites of the phylum Apicomplexa that cause a range of pathologies collectively termed coccidiosis. Species of coccidia of commercial importance have been well studied, with the effect of other species on passerine birds receiving increasing attention. In this chapter, we review the literature on coccidia in passerines, with a particular focus on wild populations. The taxonomy and life cycle of passerine coccidia are covered, as is their impact on the health of passerines, their epidemiology and their role in parasite-mediated natural and sexual selection. Coccidia can pose a significant threat to the health of wild passerine populations, and high rates of mortality have been observed in some studies. We examine some of the genetic factors that influence host resistance to coccidia and discuss how these parasites may be important in relation to sexually selected traits. General patterns are beginning to emerge with regard to the epidemiology of the parasites, and the influence of different aspects of the hosts ecology on the prevalence and intensity of coccidia is being revealed. We examine these, as well exceptions, in addition to the phenomenon of diurnal oocyst shedding that can bias studies if not accounted for. Finally, we discuss potential future directions for research on coccidia in passerines and the importance of understanding parasite ecology in the management of threatened species.

Do mothers bias offspring sex ratios in carotenoid-rich environments?

If environmental or maternal factors favor the fitness of one sex over the other, theory predicts that mothers should produce more offspring of the sex most likely to benefit from prevailing conditions. For species where males depend on carotenoid-based colorful ornaments to secure territory or attract mates, carotenoid availability in the environment could be one such component: mothers experiencing high availability of carotenoids should produce more sons. Here, we test this hypothesis by providing carotenoids to a wild population of a sexually dimorphic passerine, the hihi (stitch bird: Notiomystis cincta). Access to carotenoids during early life influences the color of male hihi plumage, which improves territory acquisition as adults. Therefore, carotenoid availability when young may influence male fitness. However, we found no evidence of sex ratio bias in treated or untreated groups, either before or after hatching. First-laid eggs, where carotenoid concentrations are usually highest, were also unbiased. For hihi, access to carotenoids during egg laying does not appear to encourage mothers to alter sex ratios of offspring. Alternatively, the fitness of daughters may also benefit from increased carotenoids during development. Disentangling these alternatives requires further work.

Research data supporting “Do mothers bias offspring sex ratios in carotenoid-rich environments?”

Data from an experimental treatment of carotenoids to hihi (Notiomystis cincta) mothers, conducted on Tiritiri Matangi Island, Auckland, New Zealand in 2004 and 2005. Spreadsheet 1 presents Laying rank and sex of eggs laid. Spreadsheet 2 contains clutch sex ratios and information about maternal age.