Evolutionary History of the Fucus Spiralis / Fucus Guiryi Complex Large-Scale Analysis of the Distribution Range and Genetic Structure Across the Northern Hemisphere

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Evolutionary History of the Fucus Spiralis / Fucus Guiryi Complex Large-Scale Analysis of the Distribution Range and Genetic Structure Across the Northern Hemisphere Evolutionary History of the Fucus spiralis / Fucus guiryi Complex Large-scale Analysis of the Distribution Range and Genetic Structure across the Northern Hemisphere Master Thesis by Amelie Prinz; [email protected] Academic Year: 2019/2020 Faculdade de Ciências e Tecnologia Evolutionary History of the Fucus spiralis / Fucus guiryi Complex Large-scale Analysis of the Distribution Range and Genetic Structure across the Northern Hemisphere Master Thesis by Amelie Prinz; [email protected] Master’s in Marine Biology (MSc) Mestrado em Biologia Marinha (MSc) Academic Year: 2019/2020 Supervisor: Ester Serrão, [email protected], (CCMAR) Co-Supervisor: João Neiva, [email protected] (CCMAR) Faculdade de Ciências e Tecnologia II Declaração de autoria de trabalho Statement of authorship and copyright Evolutionary History of the Fucus spiralis / Fucus guiryi Complex Large-scale Analysis of the Distribution Range and Genetic Structure across the Northern Hemisphere Declaro ser o(a) autor(a) deste trabalho, que é original e inédito. Autores e trabalhos consultados estão devidamente citados no texto e constam da listagem de referências incluída I declare to be the author of this work, which is original and unpublished. Authors and works consulted are duly cited in the text and are included in the list of references. _______________________________________ Amelie Prinz, Date: III Copyright A Universidade do Algarve reserva para si o direito, em conformidade com o disposto no Código do Direito de Autor e dos Direitos Conexos, de arquivar, reproduzir e publicar a obra, independentemente do meio utilizado, bem como de a divulgar através de repositórios científicos e de admitir a sua cópia e distribuição para fins meramente educacionais ou de investigação e não comerciais, conquanto seja dado o devido crédito ao autor e editor respetivos. The University of Algarve reserves the right, in accordance with the provisions of the Code of the Copyright Law and related rights, to file, reproduce and publish the work, regardless of the used mean, as well as to disseminate it through scientific repositories and to allow its copy and distribution for purely educational or research purposes and non-commercial purposes, although be given due credit to the respective author and publisher. IV Acknowledgements Firstly, I would like to acknowledge the people that contributed in gathering the samples that were analysed within this study. I was very lucky to have access to samples that were collected over years enabling the exploration of the biogeography of Fucus species on such a large geographic scale. Secondly, I want to thank all the associates of CCMAR and the BEE research group and all the colleagues in the laboratory that had helping advice during the practical working process, especially Cristina Paulino, who introduced me to the lab work, as well as Marta Sofia Mendes Valente, who processed and sequenced my samples. Thanks to Jorge Assis, who gave me an insight into the world of species distribution modelling. Additionally, I want to mention that this thesis would not have been possible without the support of my family and my friends, who always had an open ear for me during very intense times. Thank you to my supervisor, Ester Serrão, who guided me with her sharp mind and great counselling through the complexity of this study topic. Last but not least, I want to acknowledge the support of my co-supervisor, João Neiva, who was an outstanding mentor throughout this whole process. V Abstract This master thesis addresses a large-scale biogeographical question, the distribution and evolutionary history of a marine species complex with a vast distribution in the northern hemisphere. This complex includes two intertidal brown macroalgae, Fucus spiralis and Fucus guiryi, whose boundaries, degree of isolation and ranges are imperfectly known. The species Fucus guiryi has been recently separated from Fucus spiralis, and several studies have been undertaken to investigate its geographical distribution and identifying the factors determining it, especially in the face of global warming and rising sea surface temperature. The essential ecosystem services that are provided by these canopy-forming macroalgae together with factors that threaten this species, highlight the need for enhanced research. In this study a genetic diversity and differentiation analysis were applied, using microsatellites and mitochondrial DNA for assessing the influence of historical and ongoing processes in explaining species distributions, range limits and gene-pools, which are driven by great climate events such as the last glacial maximum as well as, potentially, historical and ongoing hybridization. In addition to the genetic methods, species distribution models (SDM) were developed to investigate niche differences and assess potential near-future range shifts under two contrasting climate change scenarios. Genetic analyses revealed that the investigated species complex does not consist of the aforementioned species, instead we recognize a third genetic entity currently under F. guiryi: Fucus macroguiryi. Besides the distinct lower vertical position along the shore in comparison to Fucus spiralis and Fucus guiryi, when in sympatry, Fucus macroguiryi shows a clear genetic and morphologically separation to its relatives. Most interesting results regarding species boundaries, ranges, structure and hybridization SDMs showed that the fate of the three described entities are highly different. Fucus spiralis will experience a great increase in suitable habitat for the future within the “worst case” climate scenario, contrastingly to Fucus guiryi, which will potentially have to face extreme reduction in its population sizes in the southern margins and might even face local extinctions. Fucus macroguiryi showed a poleward shift and relict populations in Morocco could as well vanish in the near future. Key words: Fucus spp., biogeography, intertidal, hybridization, range shifts, species distribution model, species complex VI Resumo Algardo Esta dissertação de mestrado aborda uma questão biogeográfica de larga-escala, a distribuição e a história evolutiva de um complexo espécies marinhas com uma distribuição muito vasta no hemisfério norte. Os modelos usados são duas macroalgas castanhas da zona entremarés, Fucus spiralis e Fucus guiryi. A espécie Fucus guiryi foi recentemente descrita, e vários estudos têm sido realizados para investigar sua distribuição geográfica e identificar os fatores que a influenciam, especialmente face ao aquecimento global e ao aumento da temperatura da superfície do mar. De acordo com o site AlgaeBase, Fucus spiralis foi registrada na costa africana , europa, e, de forma mais interessante, nas costas americanas (Atlântico e Pacífico). Por outro lado, a distribuição de Fucus guiryi está aparentemente restrita entre a Irlanda e a Grã-Bretanha e o seu limite sul na costa do Marrocos. Observações empíricas sugerem que a distribuição de Fucus guiryi pode ser maior do presentemente se conhece. Outro aspecto investigado neste estudo foi impulsionado por descobertas anteriores, que indicam que Fucus spiralis é composta por dois morfótipos distintos. Estes morfótipos ocorrem em níveis de maré distinctos chamados de Fucus spiralis Baixo e Fucus spiralis Alto. Existe também alguma incerteza à escala geográfica, como a diferenciação entre "Fucus guiryi alopátrico" (anteriormente "Fucus spiralis Sul") e "Fucus guiryi simpátrico”. “Fucus guiryi alopátrico” ocorre atualmente nas margens meridionais do complexo de espécies, Marrocos, Sahara Ocidental, Ilhas Canárias e no Sudoeste de Portugal, enquanto “Fucus guiryi simpátrico” é mais comum no Noroeste da Península Ibérica, França e ilhas britânicas. Neste estudo, o uso de microssatélites e DNA mitocondrial permitiu avaliar a influência de diversos processos históricos responsáveis pela diversificação e distribuição das espécies (limites de distribuição e “pools” de genes), e identificar uma terceira entidade dentro deste complexo de espécies: Fucus macroguiryi. Esta espécie apresenta tipicamente uma posição vertical mais baixa em comparação com Fucus spiralis e Fucus guiryi (quando em simpatria), e também mostra uma clara separação genética e morfológica. Apesar de ser possível a hibridização entre estas espécies, mecanismos pré-zigóticos e fatores ambientais adversos aparentemente garantem que as espécies mantenham sua integridade genética mesmo quando em simpatria. Além do mais, a hipótese de uma recente colonização de habitats marginais e de áreas antes não conhecidas, como o litoral do Noroeste da América, por Fucus guiryi após o último máximo glacial, se mostrou equivocada. Baixos níveis de diversidade sugerem que a América do Norte foi recentemente (re)colonizada por Fucus spiralis. Além disso, identificamos o Nordeste Atlântico como a principal área de persistência a longo prazo, caracterizada pela presença das VII três entidades e geralmente elevada diversidade. Neste estudo destacamos o Noroeste da Península Ibérica como uma região de persistência a longo prazo e o Sudoeste de Portugal como uma zona de contato entre Fucus spiralis e Fucus guiryi. Ademais, foi confirmada a hipótese de que as populações localizadas nos Açores são geneticamente separadas das populações do continente, sendo provavelmente o resultado de um isolamento crónico devido à sua distância. Este estudo também analisou potenciais padrões de conectividade de Fucus guiryi ao tentar obter uma abordagem holística através da compreensão de sua dinâmica de reprodução,
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