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Phylogenetic Analysis of the Family Rissoidae (Mollusca: Gastropoda) in the Azores Archipelago (NE Atlantic)

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Phylogenetic Analysis of the Family Rissoidae (Mollusca: Gastropoda) in the Azores Archipelago (NE Atlantic) MSc 2.º CICLO FCUP CIBIO InBio 2017 (NE (NE ( Phylogenetic Mollusca Atlantic : Gastropoda ) analysis Phylogenetic of ) in ) analysis of the the the familyy Azores family Rissoidae Rissoidae Archipelago (Mollusca: Gastropoda) in the Azores Archipelago Lara Valéria Couto Baptista Valéria Lara (NE Atlantic) Lara Valéria Couto Baptista Dissertação de Mestrado apresentada à Faculdade de Ciências da Universidade do Porto em Biodiversidade, Genética e Evolução 2017 Phylogenetic analysis of the family Rissoidae (Mollusca: Gastropoda) in the Azores Archipelago (NE Atlantic) Lara Valéria Couto Baptista MSc in Biodiversity, Genetics and Evolution Department of Biology 2017 Supervisor Sérgio P. Ávila (Ph.D), Researcher, University of Azores, CIBIO-Azores Co-supervisors António Múrias dos Santos (Ph.D), Assistant Professor, Faculty of Sciences of the University of Porto, CIBIO-InBIO M. Pilar Cabezas (Ph.D), PostDoc Researcher, CIBIO-InBIO Todas as correções determinadas pelo júri, e só essas, foram efetuadas. O Presidente do Júri, Porto, ______/______/_________ “Science never solves a problem without creating ten more.” George Bernard Shaw FCUP i Phylogenetic analysis of the family Rissoidae (Mollusca: Gastropoda) in the Azores Archipelago (NE Atlantic) Acknowledgments Before ending this chapter of my life and presenting the work developed for this dissertation, I would like to acknowledge all the people that, somehow, contributed for my personal and professional enrichment: To my supervisor, Sérgio Ávila, firstly, for giving me the opportunity to see what “real- life” scientists do in fieldwork back in 2009, definitely influencing my choice on following Genetics and Evolution as a study area in university; secondly, for accepting my request to supervise this master dissertation, accepting me in the Marine PaleoBiogeography (MPB) research group and handing me such an interesting and challenging theme about marine life in “my” archipelago; finally, for all the support and knowledge shared throughout these past years. To my co-supervisor, António Múrias dos Santos, for sharing his experience, teaching me how to deal with the bioinformatics component of this work and helping me to overcome the difficulties that appeared throughout the analyses with insightful comments and explanations. I also thank him the funding for all laboratorial work. To my co-supervisor, Pilar Cabezas Rodriguez, for the incredible support since day one, all the patience and tips about laboratorial procedures and analyses performed posteriorly. I would also like to express my gratitude for all the productive discussions to clear my doubts and for always been available to help me when I needed. It would have been very hard to finish this work without you. To the members of the MPB research group for welcoming me so well in São Miguel Island and helping me with everything in your reach. To Ricardo Cordeiro, for collecting samples used in this work and for the precious tips about laboratorial procedures and molecular analyses in family Rissoidae, after warning me about the “headaches” I would probably get with it. To Carlos Melo, for being the best “coleguinha” ever, all the jokes, the helpful discussions and, also, for his patience in creating some of the figures presented in this dissertation. To my master course colleagues, for these amazing two years, for the friendship, funny dinners, knowledge and experience shared. A special acknowledgment to Filipa and Sophia, which became close friends, for all the memorable sunsets, incredible nights out and all the laughter that helped us relax during the past year. To Beatriz and Bruna, for always be by my side even in different cities, for the craziest daily talks about everything, for making me see the results of this work from different FCUP ii Phylogenetic analysis of the family Rissoidae (Mollusca: Gastropoda) in the Azores Archipelago (NE Atlantic) perspectives, for the enthusiasm with breakthroughs and, of course, for helping me maintaining my mental health, especially in the past year. Thank you for being such amazing friends! To Sara, for supporting me in every situation ever since we were little girls. No more words are needed to express my gratitude towards you. Finally, to my family, for being the most important pillar in my life. Thanks for always encouraging me to follow my dreams and making me see “beyond today”. To my grandparents, Humberto and Gilda, for believing that the almost-microscopic organisms studied in this work are real and for supporting me unconditionally. To my sister, Joana, for all her affection and craziness, for making my life funnier and for making an effort to understand the life I wish to live. I would also like to acknowledge her skills in finishing the edition of shell’s images in 15 minutes, whilst I would take 2 hours. To my parents, Paulo and Ivone, for their unconditional love, for trusting me, for the incredible opportunities provided throughout my life and for raising me not to fear the future, but also for staying by my side in case I tremble, holding reservations about the “unknown”. To all those who directly or indirectly contributed for the realization of this work… Obrigada! FCUP iii Phylogenetic analysis of the family Rissoidae (Mollusca: Gastropoda) in the Azores Archipelago (NE Atlantic) Abstract The mollusc caenogastropod family Rissoidae Gray, 1847 comprises the largest number of microgastropod species, which can be found occupying distinct marine environments worldwide. The systematic classification of this family has been considered a difficult task due to the minute size of individuals, their diverse morphology and the convergence of shell characters, encouraging several attempts to revise classification schemes throughout the years. The most recent revision was based in molecular data and resulted on the reorganization of the Rissoidae into 20 presently recognized genera. A total of 12 of these genera are presently found in the Azores Archipelago and no phylogenetic studies about their taxonomic status are available. This study aims to analyse molecular phylogenies of the family Rissoidae and to establish the systematic position of the Azorean rissoids, which is lacking in the current scientific knowledge. Furthermore, the reconstruction of a Rissoidae species tree, calibrated with palaeontological observations in Santa Maria Island, was performed, allowing to infer the evolutionary history of the family in the Azores Archipelago and to test a theoretical biogeographical hypothesis. The results from phylogenetic analyses of nDNA 28S gene and mtDNA Cytochrome C Oxidase Subunit I (COI) and 16S genes, both by Bayesian and Maximum Likelihood methods performed using sequences of 11 rissoid species from Azorean waters and others retrieved from GenBank database, emphasize the need of a detailed revision of the genera assigned to Rissoidae. Moreover, Botryphallus ovummuscae, Setia alexandrae and S. ermelindoi displayed a considerable degree of genetic differentiation regarding the remaining rissoids, suggesting that the relationships established with other rissoids are not in accordance with the current classification of the family Rissoidae. Therefore, a systematic reclassification of the former species is proposed. Molecular similarities of Rissoa guernei and some haplotypes of Setia subvaricosa, despite some minor morphological differences, were depicted in the analyses and interpreted as a possible occurrence of sexual dimorphism in R. guernei. A theoretical biogeographical hypothesis, already sustained by ecological and palaeontological observations, was tested in this study with two non-planktotrophic species, currently reported for the Azores and Madeira and differing in their bathymetrical range, included in the species tree reconstruction. According to this hypothesis proposed by Ávila (2013, 2006), the simultaneous existence of two congeneric non-planktotrophic species, only differing in their bathymetrical range, in two nearby archipelagos can be explained if the deeper, subtidal/circalittoral species is longer-lived, in a geological FCUP iv Phylogenetic analysis of the family Rissoidae (Mollusca: Gastropoda) in the Azores Archipelago (NE Atlantic) sense, than the shallower, intertidal species. The estimation of an older split event between the subtidal and intertidal lineages provides, for the first time based on genetic data, support to this theoretical hypothesis as the subtidal species Alvania sleursi is indeed longer-lived than the intertidal Alvania mediolittoralis. The integration of palaeoceanographical data and life-traits of Azorean rissoids in the interpretation of the results obtained throughout this study led to hypothesise the potential role of Southern Azores Seamounts Chain as stepping-stones in the colonisation process of the Azores Archipelago by rissoids and littoral benthic fauna in general. The privileged setting of the Azores Archipelago in the middle of the Atlantic Ocean and the presence of fossil rissoids specimens in Santa Maria Island, make this archipelago an ideal location to study the evolution of the family Rissoidae, to hypothesise plausible processes of colonisation of these islands by marine littoral benthic gastropods, as well as to test biogeographical hypotheses. Keywords Phylogenetics, Mollusca, Gastropoda, Rissoidae, Azores Archipelago, COI, 16S rRNA, 28S rRNA, Theoretical biogeographical hypothesis, Colonization FCUP v Phylogenetic analysis of the family Rissoidae (Mollusca: Gastropoda) in the Azores Archipelago (NE
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