Speciation, despite ongoing gene flow can be studied directly in nature in ring species that comprise two reproductively isolated populations connected by a chain or ring of intergrading populations. We applied three tiers of spatio-temporal analysis (phylogeny/historical biogeography, phylogeography and landscape/population genetics) to the data from mitochondrial and nuclear genomes of eastern Australian parrots of the Crimson Rosella Platycercus elegans complex to understand the history and present genetic structure of the ring they have long been considered to form. A ring speciation hypothesis does not explain the patterns we have observed in our data (e.g. multiple genetic discontinuities, discordance in genotypic and phenotypic assignments where terminal differentiates meet). However, we cannot reject that a continuous circular distribution has been involved in the group's history or indeed that one was formed through secondary contact at the 'ring's' east and west; however, we reject a simple ring-species hypothesis as traditionally applied, with secondary contact only at its east. We discuss alternative models involving historical allopatry of populations. We suggest that population expansion shown by population genetics parameters in one of these isolates was accompanied by geographical range expansion, secondary contact and hybridization on the eastern and western sides of the ring. Pleistocene landscape and sea-level and habitat changes then established the birds' current distributions and range disjunctions. Populations now show idiosyncratic patterns of selection and drift. We suggest that selection and drift now drive evolution in different populations within what has been considered the ring.

Pathogens have been hypothesized to play a major role in host diversity and speciation. Susceptibility of hybrid hosts to pathogens is thought to be a common phenomenon that could promote host population divergence and subsequently speciation. However, few studies have tested for pathogen infection across animal hybrid zones while testing for codivergence of the pathogens in the hybridizing host complex. Over 8 y, we studied natural infection by a rapidly evolving single-strand DNA virus, beak and feather diseases virus (BFDV), which infects parrots, exploiting a host-ring species complex (Platycercus elegans) in Australia. We found that host subspecies and their hybrids varied strikingly in both BFDV prevalence and load: both hybrid and phenotypically intermediate subspecies had lower prevalence and load compared with parental subspecies, while controlling for host age, sex, longitude and latitude, as well as temporal effects. We sequenced viral isolates throughout the range, which revealed patterns of genomic variation analogous to Mayr's ring-species hypothesis, to our knowledge for the first time in any host-pathogen system. Viral phylogeny, geographic location, intraspecific host density, and parrot community diversity and composition did not explain the differences in BFDV prevalence or load between subpopulations. Overall, our analyses suggest that functional host responses to infection, or force of infection, differ between subspecies and hybrids. Our findings highlight the role of host hybridization and clines in altering host-pathogen interactions, dynamics that can have important implications for models of speciation with gene flow, and offer insights into how pathogens may adapt to diverging host populations.psittacine circovirus | host-pathogen coevolution | heterosis | sympatric speciation | crimson rosella

The plumages of parrots provide some of the most striking colouration in nature. We summarise the diversity of mechanisms producing colour in parrots and the current evidence for the adaptive significance of variation in the colour of parrot plumages. Only recently have detailed studies begun to unravel the mechanisms of their colour-production and colourvision systems. Parrots produce much of their plumage colouration through a unique suite of pigments (psittacofulvins), or through a feather tissue nanostructure that results in coherent scattering of light, or a combination of the two (producing green). Psittacofulvins are found nowhere else in nature, and may even generate fluorescence in many parrot species. Compared with other avian taxa, the adaptive significance of parrot plumage colouration remains poorly understood, although some studies suggest that plumage colouration may form important sexual signals and may be used in mate-choice by several species. There is evidence to suggest that parrot colouration can be subject to both environmental and genetic control. We emphasise that parrots offer a distinctive and useful colouration system for further study. Further research is required to unravel how the dramatic colour patterns of parrots evolved, and what roles colour signals may play in the life histories of parrots.

The vast majority of bird species build a nest in which to breed. Some species build more than one nest, but the function of most multiple nest‐building remains unclear. Here we describe the unusual nest‐building behaviour of the Australian Reed Warbler Acrocephalus australis, and test experimentally the hypotheses that multiple nest‐building is related to individual condition or territory quality, and plays a role in mate assessment. Australian Reed Warblers built two types of nest structures: ‘type I’ nests, which were used for eggs and nestlings, and ‘type II’ nests, which were structurally distinct from type I nests, did not support eggs, nestlings or adults and were not essential for successful breeding. The number of type II nests built in each territory varied. Type II nests were only built before breeding had commenced in a territory and females were not observed participating in their construction, supporting a role in female mate choice. Birds provided with supplementary food built significantly more type II nests than control birds. However, supplementary‐fed birds did not have greater pairing success, and the addition of further type II nests to territories did not increase the pairing rate or type II nest construction in those territories. There was no relationship between the presence of type II nests and either reproductive success or likelihood of nest predation. We discuss the implications of these results in light of previous suggestions regarding the function of multiple nest‐building in birds.