Estudio genético de la estructura poblacional y conectividad de dos corales endémicos del Mediterráneo"Astroides calycularis" (Pallas, 1766) y "Clodocora caespitosa" (Linnaeus, 1767)

Abstract

The main objective of this thesis is to provide knowledge about the genetic structure and extent of population connectivity in two endemic Mediterranean scleractinian corals, Astroides calycularis (Pallas, 1766) and Cladocora caespitosa (Linnaeus, 1767). These processes are generally linked to different biological characteristics, such as density and population distribution, fertility, reproductive success, the type of larval development, dispersal ability at various life stages, demographic changes and its interactions with other biotic and abiotic factors. These studies are useful for predicting the response of populations that face external disturbances or global changes. Previously, these assessments were mainly restricted to the field of fisheries biology, but recently the number of studies focusing on the genetics of keystone invertebrate species in certain marine ecosystems has grown. The two corals studied here are considered modular organisms and bioconstructor species. They provide a habitat for a diversity of associated micro- and macrofauna, not only in the intricacies of their skeletons (e.g. small polychaetes, crustaceans, molluscs, algae, dwelling organisms, sponges, etc.), but also in their tissues (e.g. zooxantellae, archeas, bacteria, etc.), thereby forming small ecosystems themselves (thus, corals are also known as “holobionts”). To achieve the main objective, different areas of the western Mediterranean Sea were sampled. Sampling for A. calycularis was primarily performed in the Alboran Sea, where the concentration of species is higher, with comparisons of samples from the Algerian Basin and Tyrrhenian Sea; C. caespitosa genetic studies focused on populations in the Balearic Sea and Algerian Basin. For genetic analyses, universal molecular markers were first used, but given the low variability obtained with these markers, microsatellite markers were designed as a basic tool for this study. The findings presented in this thesis also expand the knowledge of the sexual reproductive cycle of A. calycularis, which is important for the interpretation of the genetic analyses. The data based on the analysis of microsatellite markers attempts to clarify the degree of variability and gene flow of populations of these species and the potential impact of oceanographic barriers (i.e. currents, eddy fronts). The results suggest that the degree of genetic differentiation and connectivity among populations of both corals depend mainly on the potential dispersal of their planulae, which is primarily determined by the species sexual reproductive pattern (internal fertilization/incubation vs. external fertilization). A. calycularis and C. caespitosa are representative species of these two different reproductive patterns. In three geographically distant populations of A. calycularis, the planulae were observed to be “crawling larvae” and as and acquiring negative buoyancy at the time of release, showing low dispersal abilities. This description is consistent with the genetic results obtained from the coral populations, which suggest a “stepping-stone” pattern of connectivity and gene flow, with sporadic dispersal events at greater distances by currents and “rafting” processes. In contrast, the fertilization of C. caespitosa occurs in the water column (i.e. a broadcast spawner). Despite the lack of knowledge of the biology of C. caespitosa planulae, its reproductive mode and the high degree of connectivity found among populations suggest that its planulae are able to disperse over long distances, probably via the prevailing surface currents. In terms of the interaction between the planulae of these corals and oceanographic barriers that influence gene flow, it was found that the Almería-Orán front appears to act as a barrier to dispersal in the case of A. calycularis, in synergy with its limited dispersal abilities. In contrast, the Ibiza Channel and Balearic Front do not act as strong impediments to gene flow; although in general, a slightly higher exchange of individuals from north to south and from east to west was observed. The spatial scale of dispersal and connectivity of a keystone species, such as the corals studied in this thesis, should be taken into consideration for the design of marine protected areas (MPAs) and in the current debate of few large or many small (SLOSS). In the case of A. calycularis, the creation of micro-reserves is recommended, thus promoting self-recruitment and allowing the spread into non-protected areas. In the case of C. caespitosa, for their conservation, the scarcity of monospecific reefs should be considered; a suitable MPA would be one that is of sufficient size to protect the wealthiest localities. These two corals have suffered significant regression processes during the Pleistocene and Holocene, possibly due to climatic causes, further aggravated by anthropogenic impacts. A. calycularis is currently catalogued under different national and international legislations, which allows managers and stakeholders interested in conservation to better understand this coral. Thus, the number of studies about its biology and auto-ecology and responses of the coral to changing environments have increased, which is useful for the active management and conservation of the species. Although included in the IUCN red list, C. caespitosa currently has no legal category of protection. However, there are sufficient reasons to consider it a threatened species. Thus, the development of appropriate proposals for the categorization of this species as threatened is recommended. Studies such as those carried out in this thesis are paramount for establishing management and conservation plans, not only for the species studied, but also for other species with similar biological characteristics and life cycles. The results from this thesis provide a model approach for studying anthozoans species with different potential dispersals.