The Subantarctic Rayadito (Aphrastura subantarctica), a new bird species on the southernmost islands of the Americas.

We describe a new taxon of terrestrial bird of the genus Aphrastura (rayaditos) inhabiting the Diego Ramírez Archipelago, the southernmost point of the American continent. This archipelago is geographically isolated and lacks terrestrial mammalian predators as well as woody plants, providing a contrasted habitat to the forests inhabited by the other two Aphrastura spp. Individuals of Diego Ramírez differ morphologically from Aphrastura spinicauda, the taxonomic group they were originally attributed to, by their larger beaks, longer tarsi, shorter tails, and larger body mass. These birds move at shorter distances from ground level, and instead of nesting in cavities in trees, they breed in cavities in the ground, reflecting different life-histories. Both taxa are genetically differentiated based on mitochondrial and autosomal markers, with no evidence of current gene flow. Although further research is required to define how far divergence has proceeded along the speciation continuum, we propose A. subantarctica as a new taxonomic unit, given its unique morphological, genetic, and behavioral attributes in a non-forested habitat. The discovery of this endemic passerine highlights the need to monitor and conserve this still-pristine archipelago devoid of exotic species, which is now protected by the recently created Diego Ramírez Islands-Drake Passage Marine Park.

Opportunities for the conservation of migratory birds to benefit threatened resident vertebrates in the Neotropics.

Neotropical countries receive financing and effort from temperate nations to aid the conservation of migratory species that move between temperate and tropical regions. If allocated strategically, these resources could simultaneously contribute to other conservation initiatives. In this study, we use novel distribution maps to show how those resources could aid planning for the recovery of threatened resident vertebrates.Using eBird-based relative abundance estimates, we first identified areas with high richness of Neotropical migrant landbirds of conservation concern (23 species) during the stationary non-breeding period. Within these areas, we then identified threatened species richness, projected forest loss and conducted a prioritization for 1,261 red-listed vertebrates using Terrestrial Area-of-Habitat maps.Richness for migrants was greatest along a corridor from the Yucatan peninsula south to the northern Andes but also included south-west Mexico and Hispaniola. Protected areas account for 22% of this region while 21% is at risk of forest loss. Within this focal region for migrants, all four vertebrate groups showed hotspots of threatened species richness along the west and east Andean slopes. Taxa-specific hotspots included montane areas of southern Mexico and central Guatemala (amphibians/reptiles) and the entire east slope of the Colombian East Andes (mammals).Our prioritization highlighted several areas of importance for conservation due to high threatened species richness and projected forest loss including (a) the Pacific dry forests of south-west Mexico, (b) montane regions of northern Central America and (c) the west Andean slope of Colombia and Ecuador. At a landscape scale in southern Colombia, we show how conservation efforts for six Neotropical migrants could benefit 56 threatened residents that share a similar elevational range. Synthesis and applications. Funding and effort for migratory bird conservation also has potential to benefit threatened resident vertebrates in the Neotropics. Our study highlights how novel, high-resolution information on species distributions and risk of forest loss can be integrated to identify priority areas for the two groups at regional and landscape scales. The approach and data can be further modified for more specific goals, such as within-country initiatives.

Los países neotropicales reciben financiamiento y recursos esfuerzos de países templados para ayudar a la conservación de las aves migratorias que se mueven entre estas dos regiones. Si se asignan estratégicamente, estos recursos podrían beneficiar simultáneamente a otras iniciativas de conservación. En esta investigación, usamos novedosos mapas de distribución para mostrar cómo esos recursos podrían ayudar a planificar la conservación de especies residentes amenazadas.Utilizando estimaciones de abundancia de eBird, delineamos la Región Focal con la mayor riqueza de aves migratorias neotropicales de interés para la conservación durante el periodo invernal (23 especies). Dentro de esta región, determinamos la riqueza de especies residentes amenazadas, la proyección de pérdida de bosque, y realizamos una priorización para 1261 especies de vertebrados incluidos en la lista roja de la IUCN utilizando mapas del área de hábitat terrestre.En la Región Focal, la riqueza de aves migratorias fue mayor a lo largo de un corredor desde el sur de la península de Yucatán hasta el norte de los Andes, y en el suroeste de México y la isla La Española. Las áreas protegidas representan 22% de esta región mientras que el 21% está en riesgo de pérdida de bosque. Dentro de la Región Focal para las aves migratorias, todos los vertebrados residentes mostraron puntos de mayor riqueza de especies amenazadas en los Andes occidentales y orientales. Las regiones con mayor riqueza para taxones específicos incluyeron las montañas del sur de México y del centro de Guatemala (anfibios y reptiles) y la vertiente oriental de la cordillera oriental de Colombia (mamíferos).La priorización destacó varias áreas de importancia para la conservación debido a la alta riqueza de especies amenazadas y a la pérdida de bosque proyectada incluyendo 1) los bosques secos del Pacífico del suroeste de México, 2) las regiones montañosas del norte de Centroamérica y 3) la vertiente occidental de los Andes Occidental de Colombia y Ecuador. A escala del paisaje en el sur de Colombia, mostramos cómo los esfuerzos para la conservación de 6 aves migratorias podrían beneficiar a 56 especies residentes amenazadas que comparten un rango altitudinal similar. Síntesis y aplicaciones. La financiación y esfuerzos para la conservación de las aves migratorias tienen el potencial a beneficiar a los vertebrados residentes amenazados en el Neotrópico. Nuestro estudio resalta como se puede integrar información novedosa y de alta resolución acerca de la distribución de especies y el riesgo de pérdida de bosques para identificar áreas prioritarias para los dos grupos a escala regional y paisajística. El enfoque y los datos se pueden modificar para objetivos más específicos, como por ejemplo iniciativas dentro de cada país.

Evidence of low within-pair genetic relatedness in a relict population of Thorn-tailed Rayadito despite long-term isolation.

Investigating whether mating patterns are biased in relation to kinship in isolated populations can provide a better understanding of the occurrence of inbreeding avoidance mechanisms in wild populations. Here, we report on the genetic relatedness (r) among breeding pairs in a relict population of Thorn-tailed Rayadito (Aphrastura spinicauda) in north-central Chile that has experienced a long-term history of isolation. We used simulations based on 8 years of data to assess whether mating is random with respect to relatedness. We found that mean and median population values of pair relatedness tended to be lower than randomly generated values, suggesting that mating is not random with respect to kinship. We hypothesize that female-biased dispersal is the main mechanism reducing the likelihood of mating among kin, and that the proportion of related pairs (i.e., r > 0.125) in the study population (25%) would presumably be higher in the absence of sex-biased dispersal. The occurrence of other mechanisms such as extra-pair copulations, delayed breeding, and active inbreeding avoidance through kin discrimination cannot be dismissed and require further study.

Local selection signals in the genome of blue tits emphasize regulatory and neuronal evolution.

Understanding the genomic landscape of adaptation is central to understanding microevolution in wild populations. Genomic targets of selection and the underlying genomic mechanisms of adaptation can be elucidated by genome-wide scans for past selective sweeps or by scans for direct fitness associations. We sequenced and assembled 150 haplotypes of 75 blue tits (Cyanistes caeruleus) of a single Central European population by a linked-read technology. We used these genome data in combination with coalescent simulations (i) to estimate an historical effective population size of ~250,000, which recently declined to ~10,000, and (ii) to identify genome-wide distributed selective sweeps of beneficial variants probably originating from standing genetic variation (soft sweeps). The genes linked to these soft sweeps, but also those linked to hard sweeps based on new beneficial mutants, showed a significant enrichment for functions associated with gene expression and transcription regulation. This emphasizes the importance of regulatory evolution in the population's adaptive history. Soft sweeps were further enriched for genes related to axon and synapse development, indicating the significance of neuronal connectivity changes in the brain potentially linked to behavioural adaptations. A previous scan of heterozygosity-fitness correlations revealed a consistent negative effect on arrival date at the breeding site for a single microsatellite in the MDGA2 gene. Here, we used the haplotype structure around this microsatellite to explain the effect as a local and direct outbreeding effect of a gene involved in synapse development.

Female extra-pair behavior is not associated with reduced paternal care in Thorn-tailed Rayadito.

Extra-pair behavior is present in 76% of socially monogamous bird species with biparental care. This behavior may produce costs to females related to a reduction in paternal care. We estimated the percentage of extra-pair offspring and quantified paternal care in 44 nests of Thorn-tailed Rayadito (Aphrastura spinicauda) to assess whether males reduce their parental care when females obtain extra-pair fertilizations. We used data from a sub-Antarctic population of Rayadito located on Navarino Island (55°4'S, 67°40'W), southern Chile. We found no statistical support for a relationship between variation in paternal care and the percentage of extra-pair offspring. We discuss how the inability of breeding males to assess their genetic paternity and potential restrictions on behavioral flexibility may explain this result. Additionally, if paternal care is subjected to sexual selection, this could limit a facultative response to female extra-pair behavior by males. Finally, it is possible that a reduction in paternal care might not have evolved in this particular locality given the low frequency of extra-pair paternity in our study population.

On the relationship between baseline corticosterone levels and annual survival of the thorn-tailed rayadito.

Baseline concentrations of glucocorticoids (i.e., cortisol and/or corticosterone) can moderately increase with the degree of energy demands that an individual faces. This could be a mechanism based on which glucocorticods (GCs) can mediate life history trade-offs, and therefore fitness. The 'cort-fitness hypothesis' predicts a negative relationship between GCs and fitness, meanwhile the 'cort-adaptation hypothesis' predicts the opposite pattern. Field studies on the relation between baseline GCs and survival rate have shown mixed results, supporting both positive and negative effect. These ambiguous results could be partially consequence of the short time frame in that most of the studies are carried on. In this study, we tested the predictions of the 'cort-fitness hypothesis' and 'cort-adaptation hypothesis' by using long-term data (eight-year of capture-mark-recapture) of Thorn-tailed Rayadito (Aphrastura spinicauda) in two populations at different latitudes. We assessed whether survival varied as a function of Cort levels and whether it varied in a linear (positive: 'cort-adaptation hypothesis' or negative: 'cort-fitness hypothesis') or curvilinear way. The two populations in our study had different baseline Cort levels, then we evaluated whether the association between baseline Cort and survival probability varied between them. In the high latitude population (i.e., lower baseline Cort levels), we observed a marginally quadratic relationship that is consistent with the cort-fitness hypothesis. In contrast, in the low altitude population we did not find this relation. Our findings suggests that the association between baseline Cort and survival probability is context-dependent, and highlights the importance of comparing different populations and the use of long-term data.

Heterozygosity-Fitness Correlations in a Continental Island Population of Thorn-Tailed Rayadito.

Heterozygosity-fitness correlations (HFCs) have been used to monitor the effects of inbreeding in threatened populations. HFCs can also be useful to investigate the potential effects of inbreeding in isolated relict populations of long-term persistence and to better understand the role of inbreeding and outbreeding as drivers of changes in genetic diversity. We studied a continental island population of thorn-tailed rayadito (Aphrastura spinicauda) inhabiting the relict forest of Fray Jorge National Park, north-central Chile. This population has experienced a long-term, gradual process of isolation since the end of the Tertiary. Using 10 years of field data in combination with molecular techniques, we tested for HFCs to assess the importance of inbreeding depression. If inbreeding depression is important, we predict a positive relationship between individual heterozygosity and fitness-related traits. We genotyped 183 individuals at 12 polymorphic microsatellite loci and used 7 measures of reproductive success and estimates of apparent survival to calculate HFCs. We found weak to moderate statistical support (P-values between 0.05 and 0.01) for a linear effect of female multi-locus heterozygosity (MLH) on clutch size and nonlinear effects on laying date and fledging success. While more heterozygous females laid smaller clutches, nonlinear effects indicated that females with intermediate values of MLH started laying earlier and had higher fledging success. We found no evidence for effects of MLH on annual fecundity or on apparent survival. Our results along with the long-term demographic stability of the study population contradict the hypothesis that inbreeding depression occurs in this population.

Extrapair paternity in two populations of the socially monogamous Thorn-tailed Rayadito Aphrastura spinicauda (Passeriformes: Furnariidae).

Studies on extrapair paternity (EPP) are key to understanding the ecological and evolutionary drivers of variation in avian mating strategies, but information is currently lacking for most tropical and subtropical taxa. We describe the occurrence of EPP in two populations of a South American socially monogamous bird, the Thorn-tailed Rayadito, based on data from 266 broods and 895 offspring that were sampled during six breeding seasons in north-central and southern Chile. In the northern population, 21% of the broods contained at least one extrapair young and 14% of all offspring were sired by an extrapair male, while in the southern population, we detected extrapair offspring (EPO) in 14% of the broods, and 6% of all offspring were EPO. Variation in the frequency of EPP could stem from population differences in the duration of the breeding season or the density of breeding individuals. Other factors such as differences in breeding synchrony and variation in food availability need to be evaluated. More reports on EPP rates are necessary to determine the patterns of taxonomic and geographic variation in mating strategies in Neotropical birds, and to better understand the differences in ecological dynamics between northern and southern hemisphere populations.

Negative effects of individual heterozygosity on reproductive success in a wild bird population.

The evolutionary consequences of individual genetic diversity are frequently studied by assessing heterozygosity-fitness correlations (HFCs). The prevalence of positive and negative HFCs and the predominance of general versus local effects in wild populations are far from understood, partly because comprehensive studies testing for both inbreeding and outbreeding depression are lacking. We studied a genetically diverse population of blue tits in southern Germany using a genome-wide set of 87 microsatellites to investigate the relationship between proxies of reproductive success and measures of multilocus and single-locus individual heterozygosity (MLH and SLH). We used complimentary measures of MLH and partitioned markers into functional categories according to their position in the blue tit genome. HFCs based on MLH were consistently negative for functional loci, whereas correlations were rather inconsistent for loci found in nonfunctional areas of the genome. Clutch size was the only reproductive variable showing a general effect. We found evidence for local effects for three measures of reproductive success: arrival date at the breeding site, the probability of breeding at the study site and male reproductive success. For these, we observed consistent, and relatively strong, negative effects at one functional locus. Remarkably, this marker had a similar effect in another blue tit population from Austria (~400 km to the east). We suggest that a genetic local effect on timing of arrival might be responsible for most negative HFCs detected, with carry-over effects on other reproductive traits. This effect could reflect individual differences in the distance between overwintering areas and breeding sites.

Range-wide genetic structure in the thorn-tailed rayadito suggests limited gene flow towards peripheral populations.

Understanding the population genetic consequences of habitat heterogeneity requires assessing whether patterns of gene flow correspond to landscape configuration. Studies of the genetic structure of populations are still scarce for Neotropical forest birds. We assessed range-wide genetic structure and contemporary gene flow in the thorn-tailed rayadito (Aphrastura spinicauda), a passerine bird inhabiting the temperate forests of South America. We used 12 microsatellite loci to genotype 582 individuals from eight localities across a large latitudinal range (30°S-56°S). Using population structure metrics, multivariate analyses, clustering algorithms, and Bayesian methods, we found evidence for moderately low regional genetic structure and reduced gene flow towards the range margins. Genetic differentiation increased with geographic distance, particularly in the southern part of the species' distribution where forests are continuously distributed. Populations in the north seem to experience limited gene flow likely due to forest discontinuity, and may comprise a demographically independent unit. The southernmost population, on the other hand, is genetically depauperate and different from all other populations. Different analytical approaches support the presence of three to five genetic clusters. We hypothesize that the genetic structure of the species follows a hierarchical clustered pattern.

Latitudinal gradients of haemosporidian parasites: Prevalence, diversity and drivers of infection in the Thorn-tailed Rayadito (Aphrastura spinicauda).

Latitudinal gradients are well-suited systems that may be helpful explaining distribution of haemosporidian parasites and host susceptibility. We studied the prevalence, diversity and drivers of haemosporidian parasites (Leucocytozoon, Plasmodium and Haemoproteus) along a latitudinal gradient (30°-56° S), that encompass the total distribution (~3,000 km) of the Thorn-tailed Rayadito (Aphrastura spinicauda) in the South American temperate forests from Chile. We analyzed 516 individuals from 18 localities between 2010 and 2017 and observed an overall prevalence of 28.3% for haemosporidian parasites. Leucocytozoon was the most prevalent parasite (25.8%). We recorded 19 distinct lineages (13 for Leucocytozoon, five for Plasmodium, and one for Haemoproteus). Differences in haemosporidian prevalence and diversity by genus and type of habitat were observed in the latitudinal gradient. Further, we support the existence of a latitudinal associate distribution of Leucocytozoids in South America, where prevalence and diversity increase toward higher latitudes. Distribution of Leucocytozoon was associated with sub-antarctic habitat (higher latitude) and explained by cold temperature and high precipitation. On the other hand, we lacked to find a latitudinal associate pattern for Plasmodium and Haemoproteus, however low prevalence and high diversity were recorded in areas considered as a hotspot of biodiversity in Central Chile. Our findings confirmed the importance of habitat and climatic variables explaining prevalence, diversity and distribution of haemosporidian parasites in a huge latitudinal gradient, belonging the distribution of the Thorn-tailed Rayadito in the world's southernmost forests ecosystems.

Variation in fine-scale genetic structure and local dispersal patterns between peripheral populations of a South American passerine bird.

The distribution of suitable habitat influences natal and breeding dispersal at small spatial scales, resulting in strong microgeographic genetic structure. Although environmental variation can promote interpopulation differences in dispersal behavior and local spatial patterns, the effects of distinct ecological conditions on within-species variation in dispersal strategies and in fine-scale genetic structure remain poorly understood. We studied local dispersal and fine-scale genetic structure in the thorn-tailed rayadito (Aphrastura spinicauda), a South American bird that breeds along a wide latitudinal gradient. We combine capture-mark-recapture data from eight breeding seasons and molecular genetics to compare two peripheral populations with contrasting environments in Chile: Navarino Island, a continuous and low density habitat, and Fray Jorge National Park, a fragmented, densely populated and more stressful environment. Natal dispersal showed no sex bias in Navarino but was female-biased in the more dense population in Fray Jorge. In the latter, male movements were restricted, and some birds seemed to skip breeding in their first year, suggesting habitat saturation. Breeding dispersal was limited in both populations, with males being more philopatric than females. Spatial genetic autocorrelation analyzes using 13 polymorphic microsatellite loci confirmed the observed dispersal patterns: a fine-scale genetic structure was only detectable for males in Fray Jorge for distances up to 450 m. Furthermore, two-dimensional autocorrelation analyzes and estimates of genetic relatedness indicated that related males tended to be spatially clustered in this population. Our study shows evidence for context-dependent variation in natal dispersal and corresponding local genetic structure in peripheral populations of this bird. It seems likely that the costs of dispersal are higher in the fragmented and higher density environment in Fray Jorge, particularly for males. The observed differences in microgeographic genetic structure for rayaditos might reflect the genetic consequences of population-specific responses to contrasting environmental pressures near the range limits of its distribution.