Pronounced Fixation, Strong Population Differentiation and Complex Population History in the Canary Islands Blue Tit Subspecies Complex

Evolutionary molecular studies of island radiations may lead to insights in the role of vicariance, founder events, population size and drift in the processes of population differentiation. We evaluate the degree of population genetic differentiation and fixation of the Canary Islands blue tit subspecies complex using microsatellite markers and aim to get insights in the population history using coalescence based methods. The Canary Island populations were strongly genetically differentiated and had reduced diversity with pronounced fixation including many private alleles. In population structure models, the relationship between the central island populations (La Gomera, Tenerife and Gran Canaria) and El Hierro was difficult to disentangle whereas the two European populations showed consistent clustering, the two eastern islands (Fuerteventura and Lanzarote) and Morocco weak clustering, and La Palma a consistent unique lineage. Coalescence based models suggested that the European mainland forms an outgroup to the Afrocanarian population, a split between the western island group (La Palma and El Hierro) and the central island group, and recent splits between the three central islands, and between the two eastern islands and Morocco, respectively. It is clear that strong genetic drift and low level of concurrent gene flow among populations have shaped complex allelic patterns of fixation and skewed frequencies over the archipelago. However, understanding the population history remains challenging; in particular, the pattern of extreme divergence with low genetic diversity and yet unique genetic material in the Canary Island system requires an explanation. A potential scenario is population contractions of a historically large and genetically variable Afrocanarian population, with vicariance and drift following in the wake. The suggestion from sequence-based analyses of a Pleistocene extinction of a substantial part of North Africa and a Pleistocene/Holocene eastward re-colonisation of western North Africa from the Canaries remains possible

Characterization of microsatellite markers for the endangered Daphne rodriguezii (Thymelaeaceae) and related species

[Premise] The endangered shrub Daphne rodriguezii (Thymelaeaceae) is endemic to the Balearic island of Menorca, where fragmentation and severe population decline are ongoing threats to this taxon. We developed a set of microsatellite markers to analyze the fine-scale genetics of its few extant populations.[Methods and Results] Fifteen microsatellite markers were obtained through Illumina high-throughput sequencing and tested in two populations. Twelve of these loci showed no evidence of null alleles and were highly polymorphic, with a mean number of 8.3 alleles per locus. Levels of observed and expected heterozygosity ranged from 0.100 to 0.952 and from 0.095 to 0.854, respectively. Seven to nine of these loci were successfully amplified in five other Daphne species.[Conclusions] This set of markers provides a useful tool for investigating the factors driving fine-scale population structure in this threatened species, and it represents a novel genetic resource for other European Daphne species.J.C.I. was funded by a GRUPIN research grant from the Regional Government of Asturias (Ref.: IDI/2018/000151), and C.G.‐V. was supported by a Vicenç Mut postdoctoral fellowship awarded by the Govern de les Illes Balears–Conselleria d'Innovació, Recerca i Turisme. The study was funded by the Ministerio de Economía, Industria y Competitividad (project Islet‐Foodwebs CGL2017‐88122‐P)

Selection for functional performance in the evolution of cuticle hardening mechanisms in insects

Calcified tissues have repeatedly evolved in many animal lineages and show a tremendous diversity of forms and functions. The cuticle of many insects is enriched with elements other than Calcium, a strategy of hardening that is taxonomically widespread but apparently poorly variable among clades. Here, we investigate the evolutionary potential of the enrichment with metals in insect cuticle at different biological levels. We combined experimental evidence of Zinc content variation in the mandibles of a target species (Chorthippus cazurroi [Bolívar]) with phylogenetic comparative analyses among grasshopper species. We found that mandibular Zinc content was repeatable among related individuals and was associated with an indicator of fitness, so there was potential for adaptive variation. Among species, Zinc enrichment evolved as a consequence of environmental and dietary influences on the physical function of the jaw (cutting and chewing), suggesting a role of natural selection in environmental fit. However, there were also important within and transgenerational environmental sources of similarity among individuals. These environmental influences, along with the tight relationship with biomechanics, may limit the potential for diversification of this hardening mechanism. This work provides novel insights into the diversification of biological structures and the link between evolutionary capacity and intra- and interspecific variation.We were supported by grants 4278 of the British Ecological Society, CGL2017-85191-P/AEI/FEDER.UE from the Spanish Ministry of Science and Innovation, LIQUENES from FICYT and EDP-HC Energía, and IDI/2018/000151 from Principado de Asturias.Peer reviewe

Data from: Selection for functional performance in the evolution of cuticle hardening mechanisms in insects

[Methods] We measured Zinc content and strengths of grasshopper mandibles, by means of scanning electron microscope and calibrated multifocus pictures. We used biological and ecological information obtained from field and experimental data on the grasshopper species inhabiting the Cantabrian Mountains (NW Spain). [Usage Notes] The different datasets of the study can be found in separate sheets. An informative title on sheet content has been given to each sheet. In tables, column names refer to the variable measured (and the unit of measurement).Calcified tissues have repeatedly evolved in many animal lineages and show a tremendous diversity of forms and functions. The cuticle of many insects is enriched with elements other than Calcium, a strategy of hardening that is taxonomically widespread but apparently poorly variable among clades. Here, we investigate the evolutionary potential of the enrichment with metals in insect cuticle at different biological levels. We combined experimental evidence of Zinc content variation in the mandibles of a target species (Chorthippus cazurroi) with phylogenetic comparative analyses among grasshopper species. We found that mandibular Zinc content was repeatable among related individuals and was associated with an indicator of fitness, so there was potential for adaptive variation. Among species, Zinc enrichment evolved as a consequence of environmental and dietary influences on the physical function of the jaw (cutting and chewing), suggesting a role of natural selection in environmental fit. However, there were also important within and transgenerational environmental sources of similarity among individuals. These environmental influences, along with the tight relationship with biomechanics, may limit the potential for diversification of this hardening mechanism. This work provides novel insights into the diversification of biological structures and the link between evolutionary capacity and intra and interspecific variation.Ministerio de Ciencia e Innovación, Award: CGL2017-85191-P/AEI/FEDER.UE. Gobierno del Principado de Asturias, Award: IDI/2018/000151. British Ecological Society, Award: 4278. FYCIT Asturias.Peer reviewe

The roles of geography, climate and sexual selection in driving divergence among insect populations on mountaintops

Aim Analysing the drivers of intraspecific variation and how reproductive barriers arise is an essential step to infer the mechanisms of biogeographic differentiation. In populations of a specialized alpine species, we explore the role of geography and climate in the divergence of genetic, morphological and acoustic characters, and analyse the functional consequences of variation on mate choice. Taxon Chorthippus cazurroi (Orthoptera: Caelifera, Acrididae, Gomphocerinae). Location The entire distribution of the species (23 populations from six massifs of the Cantabrian Mountains, NW Spain). Methods First, we analysed the extent of intraspecific spatial divergence and the covariation among climatic niche, genetic (mtDNA), acoustic (song structure) and morphological (body size) traits. Then, we analysed the consequences of phenotypic variation by means of a crossing experiment among populations from different elevations. This served to test for differences in sexual selection among body size‐divergent populations and for the relationship between male traits, female preference and reproduction. Results Genetic, morphologic and acoustic divergence increased with geographic distance. Female morphology was also affected by climate variation, while male one tightly covaried with the song differentiation. Females more closely approached males investing more time in song activities, but weakly responded to the rest of acoustic features and morphological variation. They also distanced themselves slightly more from males from different populations, although this behaviour did not lead to clear differences in reproductive parameters. Main conclusions The process of colonization of mountain massifs has led to significant genetic and phenotypic changes in C. cazurroi. Phenotypic divergence does not constitute a strong intrinsic barrier to reproduction and is largely unpaired from female preference, overall suggesting that sexual selection is a minor actor in the process of differentiation as compared, for instance, to drift. This does not exclude that traits associated with individual condition are under strong selection and, therefore, do not vary so extensively. This study dismisses the idea that alpine specialists with narrow distributions lack genetic and phenotypic variability, and highlights the importance of synthesizing biogeographic and experimental approaches to obtain stronger and deeper inferences about the dynamics and mechanisms of biological differentiation.Funding for this study was provided by grant 4278 of the British Ecological Society, grants CGL2011‐28177, CGL2014‐53899‐P and CGL2017‐85191‐P from the Spanish Ministry of Economy, Industry and Competitiveness, and grant LIQUENES 2014 from FICYT and edp‐HC Energía.Peer reviewe

Gradual Distance Dispersal Shapes the Genetic Structure in an Alpine Grasshopper

The location of the high mountains of southern Europe has been crucial in the phylogeography of most European species, but how extrinsic (topography of sky islands) and intrinsic features (dispersal dynamics) have interacted to shape the genetic structure in alpine restricted species is still poorly known. Here we investigated the mechanisms explaining the colonisation of Cantabrian sky islands in an endemic flightless grasshopper. We scrutinised the maternal genetic variability and haplotype structure, and we evaluated the fitting of two migration models to understand the extant genetic structure in these populations: Long-distance dispersal (LDD) and gradual distance dispersal (GDD). We found that GDD fits the real data better than the LDD model, with an onset of the expansion matching postglacial expansions after the retreat of the ice sheets. Our findings suggest a scenario with small carrying capacity, migration rates, and population growth rates, being compatible with a slow dispersal process. The gradual expansion process along the Cantabrian sky islands found here seems to be conditioned by the suitability of habitats and the presence of alpine corridors. Our findings shed light on our understanding about how organisms which have adapted to live in alpine habitats with limited dispersal abilities have faced new and suitable environmental conditions.This study was funded by grant 4278 of the British Ecological Society, grants CGL2011-28177, CGL2014-53899-P, CGL2017-85191-P, and RYC-2015-18241 from the Spanish Ministry of Science, grant LIQUENES 2014 (CN-13-058) from FICYT and edp-HC Energía, and a GRUPIN research grant from the Regional Government of Asturias (Ref.: IDI/2018/000151).Peer reviewe

The evolutionary convergence of avian lifestyles and their constrained coevolution with species’ ecological niche

The fit between life histories and ecological niche is a paradigm of phenotypic evolution, also widely used to explain patterns of species co-occurrence. By analysing the lifestyles of a sympatric avian assemblage, we show that species' solutions to environmental problems are not unbound. We identify a life-history continuum structured on the cost of reproduction along a temperature gradient, as well as habitat-driven parental behaviour. However, environmental fit and trait convergence are limited by niche filling and by within-species variability of niche traits, which is greater than variability of life histories. Phylogeny, allometry and trade-offs are other important constraints: lifetime reproductive investment is tightly bound to body size, and the optimal allocation to reproduction for a given size is not established by niche characteristics but by trade-offs with survival. Life histories thus keep pace with habitat and climate, but under the limitations imposed by metabolism, trade-offs among traits and species' realized niche.This work was supported by the Spanish Ministry of Economy and Competitiveness (CGL2009-11302; CGL2011-28177; CGL2014-53899-P; Ramón y Cajal program) and Fundación Biodiversidad.Peer reviewe

Switching TNF antagonists in patients with chronic arthritis: An observational study of 488 patients over a four-year period

The objective of this work is to analyze the survival of infliximab, etanercept and adalimumab in patients who have switched among tumor necrosis factor (TNF) antagonists for the treatment of chronic arthritis. BIOBADASER is a national registry of patients with different forms of chronic arthritis who are treated with biologics. Using this registry, we have analyzed patient switching of TNF antagonists. The cumulative discontinuation rate was calculated using the actuarial method. The log-rank test was used to compare survival curves, and Cox regression models were used to assess independent factors associated with discontinuing medication. Between February 2000 and September 2004, 4,706 patients were registered in BIOBADASER, of whom 68% had rheumatoid arthritis, 11% ankylosing spondylitis, 10% psoriatic arthritis, and 11% other forms of chronic arthritis. One- and two-year drug survival rates of the TNF antagonist were 0.83 and 0.75, respectively. There were 488 patients treated with more than one TNF antagonist. In this situation, survival of the second TNF antagonist decreased to 0.68 and 0.60 at 1 and 2 years, respectively. Survival was better in patients replacing the first TNF antagonist because of adverse events (hazard ratio (HR) for discontinuation 0.55 (95% confidence interval (CI), 0.34-0.84)), and worse in patients older than 60 years (HR 1.10 (95% CI 0.97-2.49)) or who were treated with infliximab (HR 3.22 (95% CI 2.13-4.87)). In summary, in patients who require continuous therapy and have failed to respond to a TNF antagonist, replacement with a different TNF antagonist may be of use under certain situations. This issue will deserve continuous reassessment with the arrival of new medications. © 2006 Gomez-Reino and Loreto Carmona; licensee BioMed Central Ltd