Cetaceans have historically been object of heavy exploitation, and are still currently subject to different threats. For conservation and management purposes, most of the large cetacean populations have been categorized in stocks, considered as isolated and demographically independent management units. Many research techniques may contribute in the definition of these stocks, such as morphometric studies, mark recapture studies, genetics, satellite tracking. This thesis focuses on the development and application to cetacean populations of chemical markers of geographical origin and spatial movements. The main objective of the thesis is to evaluate the potentialities of chemical markers, such as stable isotopes and element concentrations, to investigate the patterns of distribution, the level of structuring and, when possible, the migration routes, of cetaceans populations. All these variables are essential to ensure the implementation of proper management measures and to guarantee the conservation of these animals. Three case studies were considered: the Eastern North Atlantic sperm whale (Physeter macrocephalus); the South Western Atlantic right whale (Eubalaena australis), and the North Atlantic fin whale (Balaenoptera physalus). δ15N, δ13C and δ18O values were analyzed in the dentinal growth layers of sperm whales from Denmark and NW Spain, in bone of right whales from Southern Brazil and Northern Argentina, and in bone of fin whales from W Iceland and NW Spain; concentrations of fluoride and selected metals (Zn, Pb, Ti, Sr, Cu) were analyzed in bone of fin whales from W Iceland and NW Spain. To establish the validity of each marker, the degree of compliance of each of the following four properties has been investigated: i) the basal levels of the chemical marker in the environment present significant gradients; ii) the chemical marker can be detected and measured in the tissues of whales; iii) the variability of the chemical marker within each population is moderate or, alternatively, it is predictable through measurable parameters, so that its effect can be taken into account; and iv) the values of the chemical marker in the body tissues reflect basal levels in a predictable and consistent manner. Each marker provided, at different levels, valuable information regarding the studied populations. Some difficulties emerged, connected with the complexity of the migratory cycles of whales, the possible intervention of homeostatic regulation mechanisms, and the characteristics of the tissues investigated. Nevertheless, results highlighted the occurrence of structuring within the North Atlantic sperm whale population, as well as within the South Western Atlantic right whale population, which are both currently considered as a single management unit. Moreover, results regarding the North Atlantic fin whale populations contributed to highlight the complexity of migration patterns and of the level of connectivity among the currently accepted stocks, challenging the proper definition of their limits and their degree of isolation within the basin. Some further research is recommended to deepen the knowledge about the North Atlantic fin whale population structure, as well as to investigate the levels of chemical markers in other tissues with different turnover characteristics, such as skin or baleen plates. Overall, the results obtained from the analyses performed in this thesis underline the power of chemical markers as a complementary tool to the traditional techniques used for investigating the structure and demography of cetaceans’ populations. / Los cetáceos han estado y están actualmente sujetos a diferentes tipos de amenazas. Para su correcta gestión y conservación, la mayoría de las poblaciones de grandes cetáceos han sido categorizadas en “stocks”, considerados como unidades de gestión aisladas y demográficamente independientes. Muchas técnicas de investigación pueden contribuir en la definición de estas unidades, como los estudios morfométricos, los estudios de marcaje y recaptura, la genética y el seguimiento por satélite. El objetivo principal de esta tesis es evaluar las potencialidades de los marcadores químicos, tales como los isótopos estables y las concentraciones de algunos elementos, como herramientas para la investigación de los patrones de distribución y del nivel de estructuración de las poblaciones de cetáceos. Para ello, se han considerado tres casos modelo: el cachalote del Atlántico Nororiental, la ballena franca del Atlántico Suroccidental, y el rorcual común del Atlántico Norte. Se analizaron las proporciones isotópicas de δ15N, δ13C y δ18O en muestras de dientes de cachalote y de hueso de ballena franca y rorcual común; y las concentraciones de flúor y de metales pesados (Zn, Pb, Ti, Sr, Cu) en muestras de hueso de rorcual común. Cada marcador proporcionó, a diferentes niveles, información valiosa sobre las poblaciones estudiadas. Al interpretar los resultados, surgieron algunas dificultades relacionadas con la complejidad de los ciclos migratorios de los cetáceos, la posible intervención de mecanismos de regulación homeostática, y las características de los tejidos investigados. Sin embargo, los resultados destacaron cierto nivel de estructuración dentro de las poblaciones de cachalotes del Atlántico Norte y de ballenas francas del Atlántico Suroccidental, que actualmente están ambas consideradas como unidades de gestión únicas. Por otra parte, los resultados obtenidos en las poblaciones de rorcual común del Atlántico Norte contribuyeron a evidenciar la complejidad de los patrones migratorios y del nivel de conectividad entre sus poblaciones, desafiando la adecuada definición de sus límites y de su grado de aislamiento. En general, los resultados de la tesis destacan las potencialidades de los marcadores químicos como herramienta complementaria a las técnicas utilizadas tradicionalmente para la investigación de la estructura y demografía de las poblaciones de cetáceos.

The results presented in this thesis have shown the strong dependence of Audouin’s gull from the Ebro Delta on fisheries. Foraging movements of breeding birds showed a great association with fishing activities (specially trawling), determining the most of its at-sea distribution on work days. An increase in the use of rice field habitats was also detected in absence of fishing activities (during weekends) due to the opportunistic feeding on the American crayfish. However, a key aspect presented in this thesis was the significant sex-differences in their foraging behaviour in relation to fisheries. Female Audouin’s gull tended to perform longer at-sea foraging trips on weekends compared to males, probably due to the need of feeding on high quality resources after laying instead of the easily available but lower quality American crayfish. This great dependence on fishing activities was also reflected in the resource allocation into the clutch. An isotopic δ15N enrichment in the albumen of eggs synthesized during the weekends was observed, which is attributed to an increase in the consumption of rice field resources as well as to the mobilization of female reserves. However, rice field diets were associated to lower antioxidant capacity in the eggs. Also, rice field diets were related to smaller egg size. Finally, the limiting nature of calcium (in constraining egg synthesis) and antioxidant molecules (decreasing their concentration especially in the latter eggs of the clutch) were also revealed for females. / Los resultados presentados en esta tesis han mostrado la gran dependencia de las gaviotas de Audouin del Delta del Ebro sobre las pesquerías. Los movimientos de búsqueda de alimento de las aves durante la reproducción mostraron una gran asociación con las actividades pesqueras (especialmente el arrastre durante el día, aunque también es notable la influencia de la pesca de cerco durante noche), determinando la mayor parte de su distribución en el mar los días laborables. Un incremento en el uso de arrozales fue detectado en ausencia de actividades pesqueras (durante los fines de semana). Sin embargo, un aspecto clave presentado en esta tesis fue las diferencias significativas entre sexos para el comportamiento de alimentación en relación a las actividades pesqueras. Las hembras de gaviota de Audouin tendieron a realizar viajes de alimentación más largos que los machos durante los fines de semana. Esta gran dependencia sobre las actividades pesqueras fue también reflejada en la asignación de recursos sobre la puesta. Un enriquecimiento del δ15N fue detectado en el albumen de los huevos sintetizados durante los fines de semana, lo cual puede ser atribuido a un incremento en el uso de recursos de arrozal así como a la movilización de reservas de la hembra. Sin embargo, un mayor uso de dietas de arrozal se asoció con una menor capacidad antioxidante de los huevos. También, dietas de arrozal estaban relacionadas con tamaños del huevo más pequeños. Finalmente, se puso de manifiesto la naturaleza limitante del calcio (restringiendo la síntesis del huevo) y de las moléculas antioxidantes del huevo (decreciendo su concentración especialmente en los últimos huevos de la puesta).

S'avalua la problemàtica de la gestió de la guatlla comuna (Coturnix coturnix) a Europa i Àfrica. S'han recopilat i estudiat les dades del anillamient de 43.020 exemplars i recuperació de 2.631 guatlles a Espanya, definint la tendència i distribució de l'espècie. Es descriu la història de la vida i la seva relació amb la dinàmica de la població de guatlla. Es determinen les rutes de migració de la guatlla que travessen Espanya. S'analitza la taxa de capturabilitat, l'estructura per edats i sexes, i les característiques fenotípiques i fenológiques dels exemplars capturats: pes, longitud de l'ala, longitud de la vuitena rémige primària. Banda de greix intercostal, patró de coloració del plomatge en el pit i la gola. Reproducció, claveguera i ectoparásitos macroscòpics. S'analitza la dimensió socioeconòmica i la gestió de l'espècie a nivell internacional. / Se evalúa la problemática de la gestión de la codorniz común (Coturnix coturnix) en Europa y África. Se han recopilado y estudiado los datos del anillamiento de 43.020 ejemplares y recuperación de 2.631 codornices en España, definiendo la tendencia y distribución de la especie. Se describe la historia de la vida y su relación con la dinámica de la población de codorniz. Se determinan las rutas de migración de la codorniz que atraviesan España. Se analiza la tasa de capturabilidad, la estructura por edades y sexos, y las características fenotípicas y fenológicas de los ejemplares capturados: peso, longitud del ala, longitud de la octava rémige primaria. Banda de grasa intercostal, patrón de coloración del plumaje en el pecho y la garganta. Reproducción, cloaca y ectoparásitos macroscópicos. Se analiza la dimensión socioeconómica y la gestión de la especie a nivel internacional. / The problem of the management of the common quail (Coturnix coturnix) in Europe and Africa is evaluated. We have been studied data from 43,020 banding and of 2,631recovery quails in Spain, defining the trend and distribution of the species. The quail history of life and its relation to population dynamics are described. Migration routes that cross Spain quail are determined. We used weight, wing length, length of the eighth primary rémige, also catchability rate, age and sex structure, phenotypic and phenological characteristics of the captured quails are analyzed. We studied band intercostal fat, and pattern plumage coloration in the chest and throat. Also reproduction, sewer and macroscopic ectoparasites. The socio-economic dimension and the management of the species is discussed internationally.

The aim of the present thesis is directed towards the comprehension of the mechanisms involved in the regulation of lipid and carbohydrate metabolism in relevant species in European aquaculture. Special attention has been placed on adipose tissue due to its key role in the control of the overall energy homeostasis and its endocrine functions. Six diets were formulated to determine the feasibility of replacing dietary lipids (from 23 to 17%) by starch (from 12 to 28%) and subsequent replacements in starch by fibre up to levels of 18% in gilthead sea bream. The dietary treatments did not affect fish growth nor feed utilization efficiency and triggered only moderate changes in lipid and carbohydrate metabolism. Gilthead sea bream adipose tissue exhibited an activation of the lipogenic pathway and a transcriptional regulation of relevant transcription factors directed to modulate the preferential use of lipids or carbohydrates according to availability. Despite of a lack of hepatic regulation of gluconeogenesis, an efficient metabolic hepatic adaptation to dietary starch was found through the modulation of enzymes directed to increase glucose metabolization and balance fatty acid uptake and lipolysis. These metabolic adaptations encourage the formulation of diets with a relatively high content of starch or fibre. In addition, the capacity of Atlantic salmon adipocytes to synthesise fatty acids from glucose has been demonstrated for the first time. However, the capacity of the pathway is relatively low, which might be contributing to the previously reported glucose intolerance in salmonids. Atlantic salmon adipocytes mainly transformed glucose to glycerol and Krebs-cycle intermediates. The effect of glucose on the transcriptional regulation of enzymes involved in glycolysis, the pentose phosphate pathway and glyceroneogenesis, highlights the relevance of carbohydrate metabolism in fish adipocytes as well as the metabolic adaptation of these cells to ensure the availability of necessary factors and other substrates for lipid synthesis and storage. The implication of adiponectin and TNFα in the regulation of glucose uptake in rainbow trout adipocytes has been explored, showing that both adipokines stimulated glucose uptake without modifying AKT or TOR phosphorylation. However, adiponectin failed to stimulate glucose uptake in insulin-treated adipocytes. Insulin regulated the transcription of the adiponectin system enhancing the mRNA levels of the peptide and reducing those of adiponectin receptors, following the mammalian model. However, neither adiponectin nor TNFα mRNA levels were affected by each other, concluding that the reciprocal suppressive effects reported in mammals are not conserved in fish. TNFα supressed PPARγ at a protein level, corroborating its anti-adipogenic role. However, TNFα enhanced glucose uptake in insulin-stimulated adipocytes, suggesting that the interactions between this cytokine and insulin are not so well defined in fish. Finally, the transcriptomic profile along primary culture of rainbow trout preadipocytes has been characterized revealing that adipogenesis is a complex process involving the coordinated activation or repression of genes associated to the different stages of cell development. It presented high similarities to the mammalian model, being characterized by two main phases: proliferation and differentiation. Proliferation phase was enriched with genes involved in cellular remodelling, autophagy and angiogenesis. In addition, the implication of the eicosanoid signalling pathway was highlighted during this phase. Differentiation phase was enriched with genes involved in energy production, lipid and carbohydrate metabolism. During this phase, the activation of the TR/RXR and the PPAR signalling pathways were revealed. The whole process was driven by a coordinated activation of transcription factors and epigenetic modulators. Overall, these studies have contributed to better understand the mechanisms regulating lipid and carbohydrate metabolisms in teleost fish, as well as adipose tissue growth and development. All this information might be applied to modulate and optimize farming practices.

The winter period is typically associated with harsh weather conditions and reduced food availability, which may lead to increased mortality. In birds, a common strategy to avoid such risks is to migrate to more favourable grounds, but its demographic costs are largely unknown. The wintering season remains a black box in our knowledge of the life cycle of birds. From the perspective of a wintering area, I study the population ecology of a migratory species, based on the hypothesis that winter aggregations of birds constitute true populations, instead of random aggregations of individuals. I look at the population composition, structure and dynamics across several spatial scales, and the link between the wintering and breeding areas through connecting migratory routes. My study model is a population of wintering Mediterranean Gulls Larus melanocephalus made of birds from multiple origins, a significant proportion of which have been individually marked with individually-coded colour-rings in their natal or breeding colonies. The study system constitutes the species’ main winter quarters globally and is situated on the Mediterranean coast of E-NE Spain. Capture-recapture analyses reveal high estimates of apparent local interannual survival after the first year (0.81 on average), indicating a high individual winter site-fidelity that does not vary with age. This attachment to the wintering area models the distribution of individuals, shaping them into a spatially structured population with metapopulation dynamics, as each subpopulation has independent demography, determined by area-specific survival and low levels of dispersal. Using a probability model, I infer the migratory route followed by first-winter birds; the most likely path is a combination of fluvial and coastal routes, seeking minimal distance along favourable terrain. In the focus species, wintering populations are made of individuals from different sex and age classes, in one of the few documented cases of nondifferential migration. Exceptionally, the species also exhibits synheimy (the population in each distinct wintering area is made of birds from all the breeding populations), possibly as a result of its recent history. Individual choice of wintering area is made in the first migration according to a probability rule determined by distance, and is maintained through life. On a local scale, I find a very high turnover of individuals and a frequent movement, in no particular direction, among sites. The low monthly probability of local residency (0.56) contrasts with the species’ strong site-fidelity to the general wintering area. Individual turnover shows that a single site may be visited by a large proportion of the regional (43%) and global (18%) populations every season. Thus, the population home range extends over an area much larger than a single locality or a protected area, showing evidence of a mismatch between the distribution of protected areas and the spatial needs of the population. Compared with data from 60 years ago, Mediterranean Gulls have undergone a huge expansion of their breeding distribution range (750% area increase) while the wintering distribution has increased only moderately (25%). The global population size has remained stable, indicating that its growth may be limited by events taking place in the wintering season. As the breeding distribution continues to expand, where breeding and wintering populations come in contact, they interact little and follow independent dynamics. This work contributes to a better understanding of bird population dynamics in the wintering season and shows how the detailed study of a species’ ecology during one part of the annual cycle can help comprehend what happens during the rest of the year. The ‘view from the winter quarters’ can bring important and complementary information to the study of birds during the breeding period. / Des de la perspectiva d’una àrea d’hivernada, el meu estudi es centra en l’ecologia de poblacions d’una espècie migratòria, a partir de la hipòtesi que els agrupaments hivernals d’ocells constitueixen veritables poblacions. Investigo la composició, estructura i dinàmiques de la població a diverses escales espacials, i el lligam entre les àrees de reproducció i hivernada connectades mitjançant les rutes migratòries. El meu model d’estudi és una població hivernant de Gavines Capnegres Larus melanocephalus formada per individus provinents de diversos orígens, una fracció dels quals han estat marcats individualment amb anelles de colors. El sistema d’estudi a la costa catalana constitueix la principal àrea d’hivernada de l’espècie a nivell mundial. Mitjançant anàlisis de captura-recaptura, estimo alts nivells de supervivència interanual local (0,81), indicatius d’una alta fidelitat a l’àrea d’hivernada que no canvia amb l’edat individual. La població s’estructura espacialment com a metapoblació, amb subpoblacions independents demogràficament. Un model probabilístic permet inferir la ruta migratòria dels ocells joves en llur primera migració; aquesta és una combinació de rutes fluvials i costaneres, cercant la mínima distància sobre terreny favorable. A escala local, l’espècie mostra alts nivells de rotació d’individus i freqüents moviments entre localitats, sense destí concret, i una baixa probabilitat de residència local (0,56). Les necessitats espacials de la població, superiors a una sola localitat, demostren la falta d’adequació amb la distribució de les àrees protegides per a espècies amb alta mobilitat. Comparant els darrers 60 anys, la distribució reproductora ha experimentat una espectacular expansió (increment areal, 750%) mentre que la distribució hivernal ha augmentat de manera moderada (25%). La població mundial ha romàs estable, indicant que el seu creixement pot estar limitat per esdeveniments de l’època hivernal. Allà on entren en contacte les poblacions reproductora en expansió i la hivernant, interactuen poc i mantenen dinàmiques independents. Aquest treball contribueix a millorar la comprensió de les dinàmiques poblacionals durant l’hivern i demostra com l’estudi de l’ecologia durant una part del cicle anual pot ajudar a comprendre el que succeeix la resta de l’any. La ‘visió des dels quarters d’hivernada’ pot aportar una informació important i complementària als estudis durant l’època reproductora.

This work is the result of extensive research on sexual and ontogenetic variation of parameters with mechanical significance in a radius bone in two species of odontocetes: Stenella coeruleoalba (Sc) and Pontoporia Blainvillei (Pb) (Cetacea, Mammalia). Our assumption was that the proportions of the bone structures were not only limited by the constraints of mechanical nature, but also by factors of adaptive type, including those arising from the selective pressure of possible modes of transport. In Cetacean, it has been reported that the size and shape of the pectoral limbs are related to the ecology of each species. Allometric scaling relationship of the bones of the fins in response to the ecological requirements has also been reported. Our results show negative allometric patterns in the radius of Sc, as in other oceanic dolphins and conversely, positive allometric patterns in Pb. These biomechanical differences in radius in both species might be due to an adaptation to the respective means (oceanic vs. estuarine). Gender differences in Sc are associated to the corresponding effort made males and females according to their complex social structure, in which males have greater physical activity than females. In Sc, the mechanical parameters grow more and faster in females than in males, reaching peak faster. It may be due to a compensation of loss of bone mass relative to males, caused by hormonal changes and lactation. Pb growth is slower and no such compensation is observed. Pb has a hypermineralized bone compared to what normal mineral content is considered, among which is Sc. As a result, their bones offer less resistance against transverse forces. The Ca / P ratio found for both species is greater in sexually mature animals than in other vertebrates and is similar to those cases where the bone has undergone a metabolic diseases This result suggests a non-pathological secondary osteoporosis in these dolphins, which agrees with the already describe Delphinus delphis. Bone microarchitecture is a spatial and temporal function. From the spatial point of view, according to the results of microhardness and nanohardness, bone is harder from the center outward. From the point of view of time, the average hardness of the bone (microhardness) changes with age, increasing in the whole bone in Sc and in the cortical bone in Pb whereas it decreases in the trabecular Pb bone. This observation, along with the fact that the bone is denser, would lead to a bone with more strength.

Tesi realitzada a l'Institut de Biologia Evolutiva (CSIC-UPF) / A major challenge in evolutionary biology is understanding the main drivers that underlie morphological and species diversity. Ecological opportunity—access to new or previously inaccessible niches—has been identified as one of the most important drivers of both phenotypic and species diversification. This is because the exploitation of new ecological niches is often accompanied by phenotypic differentiation among closely related taxa. This can in turn facilitate species diversification if phenotypic differentiation is associated with the appearance of reproductive isolation. The main goal of this thesis is to explore the extent in which a major source of ecological opportunity—the colonization of islands—have driven evolutionary diversification in different taxonomic and geographic contexts, specifically in the geckos of Arabia and the Socotra Archipelago and the Australasian diplodactyloid geckos. Island colonization is thought to provide a context of many available resources with few competitors and predators. This allows to colonizing groups the possibility to experience an “ecological release” and use a wider array of niches compared to their continental close-relatives. In such a situation, we expect an expansion of the morphospace in island groups typically associated with high rates of phenotypic and species diversification. We have found compelling evidence for this in both of the two mainland-island systems studied. The Australasian diplodactyloid geckos clearly expanded the range of phenotypic variation existing in the continent, producing the biggest and the smallest species in the radiation and were associated to accelerated rates of body size diversification compared to their closest relatives in the continent. Likewise, in the Hemidactylus geckos from Arabia-Socotra, island species were also associated to accelerated rates of phenotypic diversification and, as in the Australasian system, produced the most extreme sizes in the radiation. In this last mainland-island system, aside of reaching the maximum and the minimum sizes, the body size disparities in continental species assemblages were always significantly greater than the disparities computed by continental species assemblages. Nonetheless, when we compare Hemidactylus to other groups diversifying in the same islands, it appears that not all groups equally tend to diversify in body size. In fact, the comparison of the stages of diversification between the three gecko genera occurring in the Archipelago of Socotra revealed that not a single path of intra-island diversification was shared by all genera. Hemidactylus and Haemodracon greatly diversified in body size. However, in Pristurus diversification was strongly mediated by climatic shifts with size diversification being displaced to a subtle role. This is an important result as suggests that not all groups respond in the same way to similar amounts of ecological opportunity and that group-dependent (intrinsic) components can potentially play a role at defining the different stages of diversification. Aside of these examples of island diversification, in this thesis I also exposed remarkable examples of “island-like” patterns of diversification occurring in the continent. This was the case for the Australasian Pygopodidae, which attained rates of phenotypic diversification (possibly species diversification too) comparable to those found in island groups and was mostly mediated by the acquisition of a remarkable key innovation: a “snake-like” phenotype. Another remarkable example of continental diversification revealed in this thesis is a complex of highly divergent species existing within the subspecies Pristurus rupestris rupestris. This diversification took place in truly “island-like” setting, the Hajar Mountains in southeastern Arabia, formed by three main isolated blocks or “sky islands”. However, in this case diversification failed to take place within each of the “islands” and mostly was driven through intermittent pulses of dispersal and isolation taking place between two of the two main mountain blocks. Examples like this provide an example of how continental groups in some contexts, as the ones offered by mountain ranges, can fuel substantial amounts of diversification. In this case one of the greatest vertebrate diversification in Arabia. / Uno de los retos más importantes en biología evolutiva es entender las causas principales de la diversidad de fenotipos y de especies. La oportunidad ecológica (o el acceso a recursos nuevos y previamente inaccesibles) se considera como uno de los factores más importantes en la generación de nuevas especies y fenotipos. El objetivo principal de esta tesis es estudiar el papel generador de especies y fenotipos de uno de los grandes tipos de oportunidad ecológica, la colonización de islas. Para ello, se han usado dos sistemas continente-isla como modelo: los gecos de Arabia y el Archipiélago de Socotra y los gecos diplodactiloideos australasiáticos. Dado que las islas proporcionan ambientes con pocos competidores y depredadores, esto permite a los grupos insulares experimentar una gran expansión de nicho que puede estar asociada a una expansión del morfoespacio mediada por tasas elevadas de diversificación fenotípica y de especies. Los resultados de esta tesis respaldan totalmente esta hipótesis. Tanto en los gecos australasiáticos como en los gecos de Arabia y Socotra, la colonización de islas vino acompañada con aumentos sustanciales de disparidad fenotípica asociados con incrementos en las tasas de diversificación de tamaños del cuerpo y de especies. Aunque esto subraya la gran capacidad que tienen los entornos insulares en promover diversificación, diferentes grupos diversificando en las mismas islas no necesariamente siguen caminos evolutivos equivalentes. En el archipiélago de Socotra, la diversificación intra-isla en los géneros Hemidactylus y Haemodracon estuvo acompañada de grandes cambios del tamaño del cuerpo, que sólo posteriormente derivaron en cambios macroecológicos. Por el contrario, en el género Pristurus, la diversificación intra-isla fue principalmente mediada por cambios macroecológicos, siendo la diversificación del tamaño del cuerpo casi inapreciable. Esto demuestra que grupos cercanos diversificando en las mismas islas, pueden presentar patrones de diversificación completamente dispares. Al margen de las diversificaciones intra-isla expuestas, los resultados de esta tesis también han expuesto importantes patrones de diversificación también en el continente. Uno de ellos es el encontrado en los gecos pigopódidos australasiáticos, que después de adquirir una innovación clave (un fenotipo en forma de serpiente), entraron en dinámicas de diversificación comparables a los grupos insulares. Otro ejemplo destacado trabajado en esta tesis la diversidad encontrada dentro de la subespecie de geco Pristurus rupestris rupestris. Esta subespecie, en realidad, está formada por 14 especies altamente divergentes que diversificaron en las montañas del noroeste de Arábico, constituyendo uno de los casos más extremos de diversificación en vertebrados en Arabia.

Les espècies d'aranyes del gènere Loxosceles són originàries de les zones temperades i tropicals d'Amèrica i d'Àfrica. La major part de les espècies tenen àrees de distribució relativament petites, tot i que existeixen algunes excepcions a aquest patró general, i algunes espècies amb hàbits antropòfils han estat transportades de forma passiva i introduïdes en diverses zones del planeta. Aquest és el cas, per exemple, de L. rufescens (Dufour 1820), l'única espècie descrita d'Europa. Actualment, la diversitat del gènere es concentra a Amèrica del Sud i del Nord, d'on s'han descrit la major part de les 107 espècies del gènere. L'objectiu principal d'aquesta tesi ha estat estudiar la diversitat del gènere Loxosceles a la Mediterrània, el Nord d'Àfrica i les Illes Canàries, i situar-la en un context filogenètic. L'intens mostreig de camp portat a terme per tota l'àrea d'estudi a posat al descobert l'existència d'una elevada diversitat. S'han identificat 11 llinatges mitocondrials dins de L. rufescens amb dos patrons filogeogràfics contrastats: (1) uns llinatges amb poblacions ben estructurades al Marroc i la Península Ibèrica, i (2) uns llinatges que manquen d'estructura geogràfica al llarg de la Mediterrània. Les estimes d'edats han situat la major part de diversificacions durant el Pleistocè, i la diferenciació al·lopàtrica dels llinatges és compatible amb els múltiples refugis Pleistocènics identificats amb la modelització de nínxol ecològics (ENM). Sembla que el transport indirecte pels humans ha complicat la biogeografia actual en aquesta espècie. En el cas de les Illes Canàries, les anàlisis filogenètiques han posat al descobert l'existència d'un clade ben recolzat endèmic de les Illes Canàries, format per set llinatges evolutius distribuïts de forma al·lopàtrica. La major part de dispersions entre Illes van succeir durant el Miocè superior. A més, també s'han trobat representants de l'espècie cosmopolita Loxosceles rufescens en l'arxipèlag. Sis dels llinatges pels quals es disposava de representants adults, s'han estudiat morfològicament i s'han descrit com a espècies nominals, quedant la diversitat de Loxosceles en les Illes Canàries amb una espècie endèmica de Fuerteventura i Lanzarote, dues de Gran Canària, dues més de Tenerife i una de La Gomera i El Hierro. Per tal d'aprofundir en la filogeografia de l'espècie endèmica de les illes i illots orientals, s'han desenvolupat set microsatèl·lits polimòrfics utilitzant les noves tecnologies de seqüenciació. Aquests nous marcadors han servit per a ressaltar la importància dels canvis en el nivell del mar i el vulcanisme en l'estructuració genètica d'aquesta espècie. Pel que fa a la diversitat de Loxosceles del Nord d'Àfrica, s'ha descrit una nova espècie de Loxosceles de Tunísia, L. mrazig, representant la segona espècie del gènere en la Conca Mediterrània. Tot i descriure's primerament d'una sola localitat, durant mostrejos posteriors s'han localitzat en diverses zones àrides del Nord d'Àfrica, des del Marroc fins a Israel. A més, l'estudi en profunditat a la regió del Souss-Massa del Marroc ha posat al descobert l'existència de diversos llinatges evolutius molt divergents molecularment i amb notables diferències morfològiques. L'estudi del verí de diferents llinatges de L. rufescens i de dues espècies de les Illes Canàries ha demostrat que existeixen diferències respecte a la composició principalment entre L. rufescens i les dues espècies canàries, que aquestes últimes expressen una diversitat menor de paràlegs de la família gènica SicTox, i que l'activitat esfingomielinasa D és en tots els llinatges i espècies estudiades tan elevada com en espècies americanes, les quals s'ha comprovat que poden provocar casos de loxoscelime. / The spider genus Loxosceles Henieken and Lowe, 1832 is mainly distributed across the Americas and Africa. Most of the species show a narrow, circumscribed, distribution, although a few exceptions exist, for example L. rufescens (Dufour 1820), described from Europe. The aim of this thesis is to study the diversity of the genus Loxosceles in the Mediterranean Basin, North Africa and the Canary Islands, and to situate this diversity within a phylogenetic framework. The extensive sampling across the studied areas revealed the existence of hidden diversity. Using mitochondrial data, 11 different evolutionary lineages were identified within L. rufescens, presenting two contrasting phylogeographic patterns: (1) lineages with well-structured populations in Morocco and Iberia, and (2) lineages lacking geographic structure across the Mediterranean Basin. Dating analyses placed main diversification events in the Pleistocene, and multiple Pleistocene refugia, identified using ecological niche modelling (ENM), are compatible with allopatric differentiation of the lineages. Human-mediated transportation appears to have complicated the current biogeography of this medically important and synanthropic spider. This same species was found in the Canary Islands, where phylogenetic analyses revealed the existence of a well-supported clade formed exclusively by Canarian Loxosceles specimens, comprising seven allopatrically distributed evolutionary lineages. Morphological and molecular data confirmed the distinctiveness of six lineages and were described accordingly. Seven newly developed microsatellite markers were used to study the phylogeographic patterns of the species endemic to Fuerteventura and Lanzarote, and the study revealed the importance of sea-level changes and volcanism in shaping the genetic diversity in this species. Also, a new species of Loxosceles was described from Tunisia based on molecular and morphological data. Although this species was first described from a single locality, further sampling across North Africa expanded its distribution from Morocco to Israel. In the Souss-Massa region of Morocco, several highly divergent lineages were detected using molecular data and were shown to be also different morphologically. Using multiple approaches to study venom variation in selected species and lineages from the Mediterranean Basin and the Canary Islands, we found that SMase D activity, the key bioactive component of Loxosceles venom, is comparable to that of the American species that are confirmed to have medically relevant bites.

La coevolució hoste-paràsit descriu el procés de canvi evolutiu recíproc en dos organismes, hoste i paràsit, que interactuen. La coevolució hoste-paràsit s'ha estudiat fonamentalment a nivell interespecífic, i encara que processos similars poden donar-se també a nivell intraspecific, hi han molt pocs estudis en aquesta línia. A nivell microevolutiu, la dinàmica i la coevolució de les interaccions hoste-paràsit pot dependre en última instancia de la variabilitat genètica i de la seva estructuració en ambdós grups. En el present projecte de tesi, hem escollit les baldrigues del gènere Calonectris diomedea i els seus ectoparàsits, tres especies de polls de la ploma (Halipeurus abnormis, Austromenopon echinatum i Saemundssonia peusi) i una puça (Xenopsylla gratiosa); com a model d'estudi. L'objectiu principal d'aquesta tesi és investigar l'estructuració genètica de les baldrigues del gènere Calonectris i els seus ectoparàsits. Com a objectius específics ens proposem: i) Estudiar l'estructura filogeogràfica de les espècies del complex Calonectris combinant dades morfologiques i moleculars, ii) Estudiar l'estructuració genètica de les poblacions de l'espècie de baldriga cendrosa al llarg de tot el seu rang de distribució i identificar-ne els mecanismes de diferenciació implicats, iii) Estudiar l'estructuració ecològica de la comunitat d'ectoparàsits de la baldriga cendrosa, iv) Investigar la congruència en l'estructuració genètica d'hoste i ectoparàsits. Tanmateix, examinar si una especificitat diferencial per l'hoste es reflecteix en diferències en el grau de similitud de les relacions genètiques d'hoste i ectoparàsits, i finalment v) Avaluar la utilitat dels marcadors intrínsecs com a eines per assignar les baldrigues a les seves poblacions d'origen. Els nostres resultats indiquen: i) Les baldrigues del complex Calonectris mostren una estructura filogeogràfica molt marcada. Tant les dades morfològiques com els anàlisi filogenètics agrupen les poblacions en quatre clades principals d'acord amb les seves distribucions espacials segregades i la taxonomia convencional, ii) Les poblacions de baldriga cendrosa s'estructuren genèticament en dos grups corresponents les subespècies de baldriga cendrosa Atlàntica i Mediterrània. El front oceanogràfic Almeria- Orà apareix com la barrera efectiva entre les dues subespècies, iii) Les comunitats d'ectoparàsits de la baldriga cendrosa i la baldriga de Cap Verd s'estructuren a diferents nivells jeràrquics d'organització i diversos factors espacials hi juguen un paper molt important, iv) Tot i l'estructuració filogeogràfica i poblacional de l'hoste, no hem trobat estructuració genètica en cap de les tres espècies de polls de la ploma que apareixen genèticament indiferenciats. Contràriament, la puça mostra una nivells de variabilitat interespecífica alts, v) L'avaluació de diversos marcadors intrínsecs per l'assignació de baldrigues cendroses a les seves colònies de cria, demostra que és possible utilitzar marcadors biogeoquímics en plomes, elements traça i isòtops estables, com una eina efectiva en l'anàlisi de la mortalitat d'ocells marins al mar a una escala local. / In this project we chose Cory's shearwater ("Calonectris diomedea"), which includes the Mediterranean ("C. d. diomedea") and the Atlantic ("C. d. borealis") Cory's shearwater subspecies, and the Cape Verde ("C. edwardsii") shearwater, three closely related seabird taxa that host three lice ("Halipeurus abnormis", "Austromenopon echinatum" and "Saemundssonia peusi") and one flea species ("Xenopsylla gratiosa"); as a study model. The main objective of this thesis is to investigate the genetic structure of "Calonectris" shearwaters and their ectoparasites. More specifically: i) to investigate phylogenetic relationships and the biogeographic history of the "Calonectris" species complex, using both molecular and morphometric data, ii) to evaluate the population genetic structure of Cory's shearwater ("Calonectris diomedea") across its whole distribution and identify causal mechanisms of population differentiation, iii) to investigate ectoparasite community structure in Calonectris shearwaters at three levels of organization of parasite assemblages (infra-, component and regional community levels), iv) to investigate congruence between patterns of genetic structure and levels of genetic differentiation among Calonectris ectoparasites and their host taxa, and to examine whether host specificity is reflected in differences in the degree of similarity between host and ectoparasite population genetic relationships, and v) to assess the existence of spatial patterns of variation in Calonectris shearwaters by using genetic, morphometric and biogeochemical markers and evaluate the utility of all four markers as effective tools assigning Calonectris shearwaters to source populations.

Tesi realitzada a l'Institut de Ciències del Mar (ICM - CSIC) / The Mediterranean Sea can be considered an excellent natural laboratory for benthic ecologists, due to its peculiar environmental conditions, its broad-scale gradients, and its deep-sea fauna. Primary production levels in the Mediterranean follow a longitudinal gradient, decreasing from west to east. The present thesis focused on the fine-mud benthos habitat of the Mediterranean Sea, on its bathyal and abyssal zones, and the compartment considered was demersal megafauna. The main objective of the present thesis was to describe and characterize the deep habitats of the Mediterranean Sea, relating their biodiversity and ecosystem functioning with the varying environmental conditions along the geographic, bathymetric, and temporal axes. The work was divided into four parts. In the first part, the patterns of distribution, biomass and abundance of bathyal and abyssal megafauna were studied, in relation with environmental variables, along longitudinal and bathymetric gradients. This was a large geographic scale study, across the three basins of the Mediterranean at depths of 1200-3000 m. Benthic biomass strongly decreased with depth and longitude (west-to-east). Results showed a significant correlation between the dynamics of surface layers, the amount of food available on the bottom, and the composition of benthos. In the second part, the seasonal fluctuations of benthic communities in the Catalan continental margin were observed. A highly replicated, multi-period study was conducted at depths of 900-1500 m. Total biomass followed an inverted U-shaped pattern, peaking at depths of 1050-1350 m. Range-related ecological forcings between shallower and deeper species may have caused this biomass accumulation at intermediate slope depths. The arrival of a new water mass from the deep basin to the slope in spring may have also driven an accumulation of biomass at 900-1050 m depth over the same period. An adjacent submarine canyon was also studied, comparing communities between the canyon and the open slope area. Analyses revealed higher diversity, but not biomass, inside the canyon than in the adjacent open slope, and a significantly different assemblage between the habitats. These results strengthen the concept of submarine canyons as hotspots of biodiversity and underline the importance of their conservation as diversity repositories. The third part is a study on trophic relations of deep-sea fauna, over the same large spatial transects considered in the first part, by means of natural-abundances stable isotopes analyses and the use of novel statistical methods to analyse heavy datasets all in once. Bulk-tissue d13C and d15N isotope ratios were analysed for benthic megafauna and associated surface and mesopelagic components from the 3 basins of the Mediterranean Sea. The trophic niche width and the amplitude of primary carbon sources were positively correlated with both primary and secondary surface production indicators. Moreover, mesopelagic organic matter utilization processes showed an intermediate position between surface and deep benthic components. Thus, both primary and secondary production processes taking place at surface layers are key drivers of deep-sea food web structuring. Finally, the fourth part integrates the whole collected dataset into the first ecotrophic model of a deep-water ecosystem in the Mediterranean Sea, evaluating the general ecosystem functioning as well as the degree of vulnerability of these ecosystems. The Ecopath with Ecosim approach was used to model the flows and biomasses of the Catalan Sea continental slope ecosystem at depths of 1000-1400 m. Network analysis identified low levels of consumer biomass cycling and low system omnivory index when compared with expected values of marine ecosystems, and higher cycling and omnivory when compared with available models of shallower areas of the Mediterranean. A simulation of the possible expansion of the red-shrimp benthic trawl fishery that currently operates at shallower depths, showed reductions in fish biomass and that the state of the deep continental slope ecosystem seems to be result of a long-term succession process, which has reached ecological stability, and is particularly vulnerable to human impact and, specifically, to fisheries exploitation. / En la presente tesis se han estudiado los patrones de biodiversidad, funcionamiento trófico y ecosistémico de los sistemas bentónicos de gran profundidad en el Mar Mediterráneo. El objetivo principal fue de describir y caracterizar los hábitats profundos, relacionando su biodiversidad y sus procesos de funcionamiento con los gradientes ambientales a lo largo de los ejes longitudinal, batimétrico y estacional. El estudio de la composición de comunidades y la respectiva recogida de muestras para los análisis tróficos se han realizado a través de arrastres profundos. Los resultados de análisis de composición de comunidades hallaron una correlación significativa entre las dinámicas de las capas oceánicas de superficie, la cantidad de alimento disponible en el fondo marino, y la composición de la megafauna estudiada. A escala reducida y considerando las variaciones estacionales, la biomasa bentónica mostró un patrón a U-invertida, con un pico a profundidades de 1000-1400 m. Una posible explicación de este fenómeno podría ser la combinación de forzajes ecológicos ligados a las diferentes distribuciones batimétricas de las especies. Para estudiar las relaciones tróficas en los ejes batimétricos y geográficos de gran escala, se utilizó el análisis de isotopos estables de las componentes bentónicas, de superficie, y de capas pelágicas intermedias. El amplitud de los nichos tróficos de las comunidades bentónicas se encontraron relacionados con indicadores de productividad de superficie, tanto primaria como secundaria. Asimismo, los procesos de utilización mesopelágica del carbono orgánico se hallaron en posición intermedia entre los de superficie y los del bentos profundo. Finalmente, un análisis ecosistémico a través de “Ecopath with Ecosim”, reveló que los ecosistemas de los taludes continentales del Mediterráneo son el resultado de un largo proceso de sucesión ecológica, han alcanzado un estado de estabilidad, y son particularmente vulnerables a factores de impacto humano, entre ellos, la explotación pesquera.