FROM GENES TO GENOMES: LOCAL ADAPTATION AND ADAPTIVE POTENTIAL IN TWO ARCTIC SEABIRDS

By investigating the mechanisms underlying the evolution and the maintenance of local adaptations we can help predict how species will adapt to future environmental change. In this thesis I investigate local adaptation and adaptive potential in thick-billed and common murres (Uria lomvia and U. aalge), two arctic seabirds of international conservation concern. Thanks to the recent development of new genomic methods, I address three major themes that are relevant for both the development of evolutionary theory and conservation: 1) the role of gene flow in the origin and maintenance of adaptation; 2) levels and distribution of standing genetic variation, and their contribution to adaptive potential; and 3) the genomic mechanisms maintaining an adaptive dimorphism within a single interbreeding population. First, I review the literature on genomics of local adaptation with gene flow and find that adaptation can be maintained despite gene flow, that gene flow itself can promote adaptation, and that genetic architecture is important in the origin and maintenance of local adaptations. Second, I genotype genome-wide markers and toll-like receptor genes (TLRs) to investigate local adaptation and adaptive potential in thick-billed murres. Thick-billed murres do not show signatures of local adaptation to their breeding grounds, but outlier loci group birds according to their non-breeding distributions, suggesting that selection and/or demographic connectivity in the winter may explain patterns of differentiation in this species. Genetic variation at TLRs does not decrease with increasing latitude as predicted, but tests of selection and measures of genetic diversity suggest differences in local selective regimes at most genes. Thick-billed murres show high levels of standing genetic variation and their adaptive potential will mostly depend on rate and magnitude of environmental change. Finally, I improve and annotate the assembly of the highly heterozygous genome of the thick-billed murre. Using this assembly as a reference, I perform whole genome analyses to investigate the genomic basis of an adaptive dimorphism in Atlantic common murres. I show for the first time that a 60 kb complex copy number variant in a non-coding region maintains differences in plumage and cold adaptation despite high gene flow.

Pronounced inter-colony variation in the foraging ecology of Australasian gannets : influence of habitat differences

The predictability of prey due to oceanographic features can result in large aggregations of apex predators. Central place foragers, such as seabirds, are limited in their foraging duration and range during the breeding period, which can restrict their ability to reach such locations. Segregation by colony and sex can further restrict foraging range and may have direct implications for the foraging ecology of a species. In parts of its range, the Australasian gannet Morus serrator breeds in colonies of relatively close proximity and has recently been found to display sexual dimorphism. Two neighbouring breeding colonies in Bass Strait that experience divergent environmental conditions were investigated to determine whether these conditions or the sex of the individual are important variables influencing foraging behaviour in this species. GPS tracking and accelerometry were paired with stable isotope analysis to compare differences in foraging effort, habitat use and diet. Birds from Point Danger, a large colony located near a seasonally strong upwelling, travelled considerably further (77%) than birds from Pope's Eye, a small colony in a nutrient-poor embayment. However, within colonies no sexual differences in foraging effort were found. While the colonies did not overlap in foraging areas, a degree of sexual segregation was apparent within both colonies (Point Danger 46.3% overlap and Pope's Eye 73.7% overlap in home range). Furthermore, stable isotope analysis indicated birds from each colony fed at different trophic niches. This study reveals differences in habitat use and, consequently, dietary niches between and within neighbouring colonies.

Sexual segregation in habitat use is smaller than expected in a highly dimorphic marine predator, the southern sea lion

Sexual segregation in habitat use is widely reported in many taxa and can profoundly influence the distribution and behaviour of animals. However, our knowledge of the mechanisms driving sexual segregation is still in its infancy (particularly in marine taxa) and the influence of extrinsic factors in mediating the expression of sex differences in foraging behaviour is underdeveloped. Here, we combine data from biologging tags, with stable isotope analysis of vibrissae, to assess sexual segregation in southern sea lions (SSL) (Otaria flavescens) breeding at the Falkland Islands in the South Atlantic. We found evidence to support segregation, most notably in δ13C and δ15N values. However, in spite of extreme sexual size dimorphism and differing constraints related to female-only parental care, adult male and adult female SSL overlapped considerably in isotopic niches and foraging area, and shared similar foraging trip characteristics (such as distance and duration). This is in contrast to SSL breeding in Argentina, where prior studies report sexual differences in foraging locations and foraging trip characteristics. We posit that sexual segregation in SSL is influenced by habitat availability (defined here as the width of the Patagonian Shelf) and individual foraging preferences, rather than commonly invoked individual-based limiting factors per se.

Sexual selection predicts brain structure in dragon lizards

Phenotypic traits such as ornaments and armaments are generally shaped by sexual selection, which often favours larger and more elaborate males compared to females. But can sexual selection also influence the brain? Previous studies in vertebrates report contradictory results with no consistent pattern between variation in brain structure and the strength of sexual selection. We hypothesize that sexual selection will act in a consistent way on two vertebrate brain regions that directly regulate sexual behaviour: the medial preoptic nucleus (MPON) and the ventromedial hypothalamic nucleus (VMN). The MPON regulates male reproductive behaviour whereas the VMN regulates female reproductive behaviour and is also involved in male aggression. To test our hypothesis, we used high-resolution magnetic resonance imaging combined with traditional histology of brains in 14 dragon lizard species of the genus Ctenophorus that vary in the strength of precopulatory sexual selection. Males belonging to species that experience greater sexual selection had a larger MPON and a smaller VMN. Conversely, females did not show any patterns of variation in these brain regions. As the volumes of both these regions also correlated with brain volume (BV) in our models, we tested whether they show the same pattern of evolution in response to changes in BV and found that the do. Therefore, we show that the primary brain nuclei underlying reproductive behaviour in vertebrates can evolve in a mosaic fashion, differently between males and females, likely in response to sexual selection, and that these same regions are simultaneously evolving in concert in relation to overall brain size.

Identification of deep water lantern sharks (Chondrichthyes : Etmopteridae) using morphometric data and multivariate analysis

Taxonomic distinction to species level of deep water sharks is complex and often impossible to achieve during fisheries-related studies. The species of the genus Etmopterus are particularly difficult to identify, so they often appear without species assignation as Etmopetrus sp. or spp. in studies, even those focusing on elasmobranchs. During this work, the morphometric traits of two species of Etmopterus, E. spinax and E. pusillus were studied using 27 different morphological measurements, relatively easy to obtain even in the field. These measurements were processed with multivariate analysis in order to find out the most important ones likely to separate the two species. Sexual dimorphism was also assessed using the same techniques, and it was found that it does not occur in these species. The two Etmopterus species presented in this study share the same habitats in the overlapping ranges of distribution and are caught together on the outer shelves and slopes of the north-eastern Atlantic.

LMS tables for waist circumference and waist-to-height ratio among Colombian children and adolescents: The FUPRECOL study

Background: Indices predictive of central obesity include waist circumference (WC) and waist-to-height ratio (WHtR). The aims of this study were 1) to establish a Colombian youth smoothed centile charts and LMS tables for WC and WHtR and 2) to evaluate the utility of these parameters as predictors of overweight and obesity. Method: A cross-sectional study whose sample population comprised 7954 healthy Colombian schoolchildren [boys n=3460 and girls n=4494, mean (standard deviation) age 12.8 (2.3) years old]. Weight, height, body mass index (BMI), WC and WHtR and its percentiles were calculated. Appropriate cut-offs point of WC and WHtR for overweight and obesity, as defined by the International Obesity Task Force (IOTF) definitions, were selected using receiver operating characteristic (ROC) analysis. The discriminating power of WC and WHtR was expressed as area under the curve (AUC). Results: Reference values for WC and WHtR are presented. Mean WC increased and WHtR decreased with age for both genders. We found a moderate positive correlation between WC and BMI (r= 0.756, P < 0.01) and WHtR and BMI (r= 0.604, P < 0.01). The ROC analysis showed a high discrimination power in the identification of overweight and obesity for both measures in our sample population. Overall, WHtR was slightly a better predictor for overweight/obesity (AUC 95% CI 0.868-0.916) than the WC (AUC 95% CI 0.862-0.904). Conclusion: This paper presents the first sex- and age-specific WC and WHtR percentiles for both measures among Colombian children and adolescents aged 9–17.9 years. By providing LMS tables for Latin-American people based on Colombian reference data, we hope to provide quantitative tools for the study of obesity and its comorbidities.

Triceps and subscapular skinfold thicknesses percentiles and cut-offs for overweight and obesity in a population-based sample of schoolchildren in Bogota, Colombia

OBJECTIVES: The aims of this study were to establish a Colombian smoothed centile charts and LMS tables for tríceps, subscapular and sum tríceps+subscapular skinfolds; appropriate cut-offs were selected using receiver operating characteristic analysis based in a populationbased sample of schoolchildren in Bogota, Colombia and to compare them with international studies. METHODS: A total of 9 618 children and adolescents attending public schools in Bogota, Colombia (55.7% girls; age range of 9–17.9 years). Height, weight, body mass index (BMI), waist circumference, triceps and subscapular skinfold measurements were obtained using standardized methods. We have calculated tríceps+subscapular skinfold (T+SS) sum. Smoothed percentile curves for triceps and subscapular skinfold thickness were derived by the LMS method. Receiver operating characteristics curve (ROC) analyses were used to evaluate the optimal cut-off point of tríceps, subscapular and sum tríceps+subscapular skinfolds for overweight and obesity based on the International Obesity Task Force (IOTF) definitions. Data were compared with international studies. RESULTS: Subscapular, triceps skinfolds and T+SS were significantly higher in girls than in boys (P <0.001). The median values for triceps, subscapular as well as T+SS skinfold thickness increased in a sex-specific pattern with age. The ROC analysis showed that subscapular, triceps skinfolds and T+SS have a high discrimination power in the identification of overweight and obesity in the sample population in this study. Based on the raw non-adjusted data, we found that Colombian boys and girls had high triceps and subscapular skinfolds values than their counterparts from Spain, UK, German and US. CONCLUSIONS: Our results provide sex- and age-specific normative reference standards for the triceps and subscapular skinfold thickness values in a large, population-based sample of 3 schoolchildren and adolescents from an Latin-American population. By providing LMS tables for Latin-American people based on Colombian reference data, we hope to provide quantitative tools for the study of obesity and its complications.

Hematological parameters of three freshwater stingray species (Chondrichthyes: Potamotrygonidae) in the middle Rio Negro, Amazonas state.

This paper aimed to study and compare the hematology of newborns, young, subadults, adult males, adult females and pregnant females of Potamotrygon wallacei (cururu stingray), Potamotrygon motoro and Paratrygon aiereba. Newborn cururu stingrays had lower red blood parameters than those of other development stages. Thrombograms and leukograms showed a conservative pattern between development stage, sexual dimorphism and pregnancy. In P. motoro and P. aiereba, variables relating to red blood parameters, biochemistry and leukograms showed little variation between the species' biological characteristics, thus showing that these variables are not good criteria for differentiating them within the same species. In conclusion, the development stage is an important factor for differentiating hematological properties in the cururu stingray, while this has not been observed in P. motoro and P. aiereba stingrays.