Adaptive divergence in Darwin's small ground finch (Geospiza fuliginosa): divergent selection along a cline

We examine here, in a single year (2005), phenotypic divergence along a 560‐m elevation gradient in Darwin's small ground finch (Geospiza fuliginosa) in the Galápagos Islands. In this sample, four composite measures of phenotypic traits showed significant differences along the 18‐km geographical cline extending from lowlands to highlands. Compared with lowland birds, highland birds had larger and more pointed beaks, and thicker tarsi, but smaller feet and claws. Finches in an intervening agricultural zone had predominantly intermediate trait values. In a second, mark–recapture study we analyse selection on morphological traits among birds recaptured across years (2000–2005) in lowland and highland habitats. Birds were more likely to survive in the highlands and during the wet season, as well as if they had large beaks and bodies. In addition, highland birds exhibited higher survival rates if they had small feet and pointed beaks – attributes common to highland birds as a whole. Lowland birds were more likely to survive if they possessed the opposite traits. Selection therefore reinforced existing morphological divergence, which appears to reflect local adaptation to differing resources during the predominantly drought‐ridden conditions that characterized the 5‐year study. Alternative explanations – including genetic drift, matching habitat choice, deformation by parasites, and the effects of wear – received little or no support. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 45–59.     

Impacts of trichomonosis epidemics on Greenfinch Chloris chloris and Chaffinch Fringilla coelebs populations in Finland

Population consequences of parasites in wild birds are rarely documented. One exception is the decline of British finch populations due to an epidemic caused by the protozoan parasite Trichomonas gallinae. Finch trichomonosis has recently spread from the UK to northern Europe, but its consequences for local finch populations have not been studied. We assessed the extent to which the trichomonosis epidemic affected the Finnish population sizes of European Greenfinch Chloris chloris, Common Chaffinch Fringilla coelebs and a control species, Great Tit Parus major, and the body condition of Greenfinches. The disease was first documented in Finland in 2008 and epidemics were observed mainly in southwestern Finland. Greenfinches showed a significant decline of 47% in breeding numbers and 65% in wintering numbers in southern Finland during 2006–2010. Breeding Chaffinch numbers showed a slight decline (4%) during the same study period that was significant only in central Finland. Great Tit did not show a significant change in breeding numbers. During the initial disease epidemic the body condition of all demographic groups of Greenfinches decreased equally, which suggests that the disease was not selective in respect of age or sex. There were no encounters of Finnish‐ringed Greenfinches or Chaffinches in the UK, which could indicate that the parasite has not necessarily been transferred directly from the UK, but perhaps by migrants from Sweden and Germany.     

The classical case of character release: Darwin's finches (Geospiza) on Isla Daphne Major, Galápagos

The history of how Darwin's medium ground finch (Geospiza fortis) on Isla Daphne Major, Galápagos became a textbook example of character release is reviewed. Four hypotheses for the evolution of the intermediate-sized Daphne fortis are examined, including genetic drift/founder effect, hybridization with the small ground finch (G'. fuliginosa) , food supply (or local adaptation hypothesis) and character release in allopatry.
Modern data suggest that genetic drift is unlikely to have been important, due to inadequate isolation and over-riding selection and introgression on Daphne. All three remaining hypotheses have probably played a role. Hybridization with G. fuliginosa occurs, although it cannot counteract the selection pressures seen during our study. Local adaptation has also occurred, with natural selection changing the relative frequencies of fortis phenotypes in response to changes in Daphne food supplies. The selection resulted from correlations between the size of seeds available, feeding behaviour and morphology. However, recent phenotypic tracking has resulted in larger, not smaller phenotypes. There is also evidence for character release in the form of diet expansion by G. fortis during periods of food shortage, and indirect evidence for interspecific competition between fortis and the cactus ground finch (G. scandens). The Daphne fortis phenotype probably represents a balance between introgression with fuliginosa, selection for larger body size in dry years and selection for smaller body size in wet years. The simple textbook account of a character shift caused by the accidental absence of competitors should be qualified to reflect the ecological complexity of the situation.     

Interspecific competition between introduced house finch populations and two associated passerine species

Summary House Finches (Carpodacus mexicanus), natives of western North America, have expanded their range in the eastern United States since their 1940 release in New York City. Range and the relation of House Finch population growth to the population dynamics of House Sparrows (Passer domesticus) and Purple Finches (Carpodacus purpureus) were examined, using data from the Breeding Bird Survey and the Christmas Bird Count. The House Finch population grew exponentially throughout its eastern range. Significant negative relationships in population density, relative to spatial and temporal control populations, were found between House Finches and House Sparrows in summer and winter, and between House Finches and Purple Finches in summer. Purple Finch and House Sparrow populations outside of the House Finch range appeared to have no effect on each other throughout the study. Neither changes in 74 weather variables, nor changes in forest, field or developed habitat explained the observed trends in population density. The results indicate that House Finches compete with these two species, but winter migration complicates the picture.     

Isolation, characterization and multiplex polymerase chain reaction of novel microsatellite loci for the avian parasite Philornis downsi (Diptera: Muscidae)

An enrichment technique was used to isolate 11 di-, tri-, and tetra microsatellites for the parasitic fly Philornis downsi (Diptera: Muscidae). These loci were polymerase chain reaction amplified in singleplexes or two-plexes for P. downsi. The loci showed low to moderate polymorphism, exhibited between three and four alleles, and observed heterozygosity ranged from 0.05 to 0.86. These new markers will be useful for population-level and paternity analyses and will provide valuable information about the ecology of this high-impact parasite of vulnerable bird species.     

Sex differences in survival selection in the serin,Serinus serinus

Natural selection is demonstrated in most natural populations which suggests that populations are dispatched from their adaptive peaks as a result of selection on correlated characters, or conflicting selection between the sexes. We analysed patterns of survival selection in a population of serins (Serinus serinus) outside Barcelona over a period of 13 years. There was directional selection for increased wing length in males and females accompanied by strong disruptive selection on both tail and wing length in males and a selection against a positive correlation between the two characters in males. In females there was directional selection for increased bill width but decreased bill depth, which should be contrasted to the stabilizing selection acting on bill depth in males. There were conflicting selection on the characters within a sex and conflicting selection of the same characters between sexes, which constrain the rate of access to the nearest adaptive peak.     

Host‐specific associations affect the microbiome of Philornis downsi,an introduced parasite to the Galápagos Islands

The composition and diversity of bacteria forming the microbiome of parasitic organisms have implications for differential host pathogenicity and host–parasite co‐evolutionary interactions. The microbiome of pathogens can therefore have consequences that are relevant for managing disease prevalence and impact on affected hosts. Here, we investigate the microbiome of an invasive parasitic fly Philornis downsi, recently introduced to the Galápagos Islands, where it poses extinction threat to Darwin's finches and other land birds. Larvae infest nests of Darwin's finches and consume blood and tissue of developing nestlings, and have severe mortality impacts. Using 16s rRNA sequencing data, we characterize the bacterial microbiota associated with P. downsi adults and larvae sourced from four finch host species, inhabiting two islands and representing two ecologically distinct groups. We show that larval and adult microbiomes are dominated by the phyla Proteobacteria and Firmicutes, which significantly differ between life stages in their distributions. Additionally, bacterial community structure significantly differed between larvae retrieved from strictly insectivorous warbler finches (Certhidea olivacea) and those parasitizing hosts with broader dietary preferences (ground and tree finches, Geospiza and Camarhynchus spp., respectively). Finally, we found no spatial effects on the larval microbiome, as larvae feeding on the same host (ground finches) harboured similar microbiomes across islands. Our results suggest that the microbiome of P. downsi changes during its development, according to dietary composition or nutritional needs, and is significantly affected by host‐related factors during the larval stage. Unravelling the ecological significance of bacteria for this parasite will contribute to the development of novel, effective control strategies.     

Evolution of bite force in Darwin's finches: a key role for head width

Studies of Darwin's finches of the Galapagos Islands have provided pivotal insights into the interplay of ecological variation, natural selection, and morphological evolution. Here we document, across nine Darwin's finch species, correlations between morphological variation and bite force capacity. We find that bite force correlates strongly with beak depth and width but only weakly or not at all with beak length, a result that is consistent with prior demonstrations of natural selection on finch beak morphology. We also find that bite force is predicted even more strongly by head width, which exceeds all beak dimensions in predictive strength. To explain this result we suggest that head width determines the maximum size, and thus maximum force generation capacity of finch jaw adductor muscles. We suggest that head width is functionally relevant and may be a previously unrecognized locus of natural selection in these birds, because of its close relationship to bite force capacity.     

Reconstructing ancestral ecologies: challenges and possible solutions

There are several ways to extract information about the evolutionary ecology of clades from their phylogenies. Of these, character state optimization and 'ancestor reconstruction' are perhaps the most widely used despite their being fraught with assumptions and potential pitfalls. Requirements for robust inferences of ancestral traits in general (i.e. those applicable to all types of characters) include accurate and robust phylogenetic hypotheses, complete species-level sampling and the appropriate choice of optimality criterion. Ecological characters, however, also require careful consideration of methods for accounting for intraspecific variability. Such methods include 'Presence Coding' and 'Polymorphism Coding' for discrete ecological characters, and 'Range Coding' and 'MaxMin Coding' for continuously variable characters. Ultimately, however, historical inferences such as these are, as with phylogenetic inference itself, associated with a degree of uncertainty. Statistically based uncertainty estimates are available within the context of model-based inference (e.g. maximum likelihood and Bayesian); however, these measures are only as reliable as the chosen model is appropriate. Although generally thought to preclude the possibility of measuring relative uncertainty or support for alternative possible reconstructions, certain useful non-statistical support measures (i.e. 'Sharkey support' and 'Parsimony support') are applicable to parsimony reconstructions.