On the Origin of Cryptic Species: Insights from the Stygocapitella Species Complex
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On the Origin of Cryptic Species: Insights from the Stygocapitella species complex José Cerca Thesis submitted for the degree of Philosophiae Doctor Natural History Museum Faculty of Mathematics and Natural Sciences University of Oslo 2019 "The beauty and brilliancy of this insect are indescribable, and none but a naturalist can understand the intense excitement I experienced when I at length captured it. On taking it out of my net and opening the glorious wings, my heart began to beat violently, the blood rushed to my head, and I felt much more like fainting than I have done when in apprehension of immediate death. I had a headache the rest of the day, so great was the excitement produced by what will appear to most people a very inadequate cause." Alfred Russel Wallace “Em cada esquina um amigo, em cada rosto igualdade.” (In each corner a friend, in each face equality) José Afonso – Zeca “Não sou nada. Nunca serei nada. Não posso querer ser nada. À parte disso, tenho em mim todos os sonhos do mundo.” (I am nobody. I will never be anything. I cannot desire to be anything. Other than this, I hold every dream in the world.) Fernando Pessoa Contents Acknowledgements Page 1 List of manuscripts and appendices included Page 5 Summary Page 7 Introduction Page 9 Methods and materials Page 13 Main findings and Discussion Page 21 References Page 46 Manuscripts and appendices Page 57 José Cerca – On the Origin of Cryptic Species: Insights from the Stygocapitella species complex Acknowledgments I belong to a lineage from Portugal’s rural interior. Growing up in an isolated city in the interior of the country, I never had many goals or ambitions. As a 17-year-old I took a ‘career and intelligence test’ which pointed out that I had an IQ far lower than the world average. Going to University was not a tough choice though – everyone else did it, and it was only natural that I followed the herd. My first choice was Sports’ science, but by the time I had made my choice I had already missed the mandatory physical exams. Back then, I had a supportive biology teacher who helped me understand that I was passionate about biology. I went to University to study biology with no concrete goal in mind, but knew I would be happy becoming a biology high-school teacher. During my period at the University of Coimbra I met incredible people who helped me, stimulated me, and trained me – I developed a liking for plant taxonomy and eventually did a MSc in pollinator ecology. During those years I took a liking to student politics and debating which helped me mature. It was not until I read for my MSc thesis that I understood that I felt like a fish out of water doing ecology. During that period, I heard a talk of Rosemary Gillespie (ESEB 2013) on adaptive radiation of spiders which helped me understand I was truly passionate about evolutionary biology. Following my MSc, I was unemployed for a year. I felt undone and spent my time trying to read up on evolutionary biology. I read Nosil’s Ecological Speciation and Schulter’s The Ecology of Adaptive Radiation, which made me understand – that’s where I want to go. I want to work with evolution and its interface with ecology and morphology and other domains. Having a background in taxonomy and in ecology, in a time where evolutionary biology was being transformed by the ‘genomics- revolution’, I could not find a position. I had no network, no support or advice, and did not even understand what I had to do (or the skills to learn) to secure a position. My year of unemployment (note, I took some not-so-relevant jobs) started weighting and defeating me. Nobody was willing to take someone without a skillset in bioinformatics, let alone without much genetics-lab experience. I applied to nearly 40 positions before I took my current position in Oslo. This background fuels my ambition to work as hard as I can to prove my abilities to myself and to the world. But it also highlights the role of those who believed in me and helped me become who I am. To you, I could not be more grateful. There is not a single day that goes by that I do not acknowledge how lucky I am in having found something that I am so passionate about, and for having had the support to mature and to navigate my way to evolutionary biology. I was lucky I had food, a roof, a bed and a peaceful environment to grow. That being said, I do not think acknowledgements should be laid in paper. Acknowledgements should be done on a daily basis, embodied through caring actions and words, smiles and celebrating each other’s success. I thus write these words for personal memories, but also to make sure that in the midst of all our cultural differences, you understand how deeply grateful I am to all of you. First, and foremost, I am thankful to my main supervisor Torsten H. Struck. I cannot thank you enough for believing in my potential from the beginning. For accepting that you had to train me and that you had to be patient. You invested an incredible amount of time in me – helping me in the lab, discussing 1 José Cerca – On the Origin of Cryptic Species: Insights from the Stygocapitella species complex PCR-gel results as the photography came out, letters of recommendation, grant proposals, in the field, in the terminal, in manuscripts, research ideas, talks and posters. There are many lessons I have learnt from you, but most importantly, you have taught me to always have a high ethical standard – that piece of rigor that separates excellent science from shenanigans. I hope one day I become a scientist like you and make you proud that I came from your laboratory. To Mark Ravinet, for all your advice and support. Hearing how you faced the adversities of your career, and your advice on navigating through graduate school made me spend 4 years admiring you. To Lutz Bachmann, for hours and hours of advice – which made me think twice, and perhaps even become less reckless. To Hugo de Boer for all your time, advice and encouragement. Your ambition and strategy-minded thinking will always be a reference (and a lesson) to me. To Dimitar Dimitrov for your friendship, advice, collaboration and mentorship. It was truly great to have regular lunches with you, as well as to learn from you. To Mike Nowak, for always having an open door. To Günter Purschke for all your support as my co-advisor and expertise in invertebrate morphology. To Mark Blaxter and Julian Catchen for receiving me and treating me as one of your own. I hope to carry a bit of Torsten, MarkR, Lutz, Hugo, Mike, Dimitar, Günter, MarkB, Julian with me. To everyone who has helped me building up this thesis. This includes the three lab warlocks Lisbeth, Jarl Andreas and Audun for your kind support in laboratory work. To Christian Meyer for helping in sampling and photographing worms. To those who provided me with laboratory space, funding or help in obtaining specimens including Andrew Mackie, Claudia Miller, Christer Erséus, Gustav Paulay, Henning Reiss, Inês Modesto, Lis Lindal Jørgensen, Natalyia Budaeva, Nicola Pennisi, Tim Worsfold, Vasily Radashevsky, and relevant funding bodies. To my kind and supportive girlfriend Tora Kjærnes Knutsen, who I want to spend my life next to. For all your love and comprehension (i.e. wasted summer holidays, lonely weeks and weekends). À minha mãe e ao meu pai, porque apesar de pensarmos de forma diferente, eu sempre vos amei. Aos meus avós, que estariam em lágrimas por terem criado ‘um doutor’. Ao meu irmão Luis Nuno por tanto ser besta como bestial, mas por estar sempre presente quando preciso. To my close friends who kept me going and made me constantly feel saudade (Wikipedia translation: ‘deep emotional state of nostalgic or profound melancholic longing for an absent something or someone that one cares for and/or loves (…). One English translation of the word is missingness, although it might not convey the feeling of deep emotion attached to the word "saudade"’). It is not easy to fight alone and I felt the weight of distance was counteracted by your messages and caring support. Thank you urso-Manso (por estares sempre presente quando preciso, e por juntos sermos as duas maiores, mas mais felizes, bestas do mundo), Ruizinho (possas – porque não conseguiria ser feliz sem mandar postas contigo!), Eli (por seres um amor e um grande apoio; já sabes, se o Rui não der..), Ivo (o iron-palhaço que me deu sempre casa e apoio sempre que precisei – mesmo quando não tinha tempo), Rocha (pelo constante apoio, desde que nascemos e até aos nossos últimos dias), Chichorro (pelo apoio e entendimento nesta jornada de doutoramento), Pardal (por estares constantemente preocupada comigo), Maria Palma (por teres sempre uma palavra carinhosa), 2 José Cerca – On the Origin of Cryptic Species: Insights from the Stygocapitella species complex MafaldaJ (por me teres acompanhado desde sempre e me entenderes como ninguém), Paula e Pedro (olhem que vou precisar de espaço no vosso casamento – obrigado por estarem sempre presentes), MafaldaM (por me diagnosticares que sou obcessivo-compulsivo, hiperactivo e ter um défice de atenção – e por aturares toda uma pancada nestes anos), Lucas (por partilharmos ambição e ‘background’ e crescermos lado-a-lado), David (pela nossa amizade de infância e por ter orgulho das tuas causas), Tiago da Rapoula (possas, tu sabes – sem ti eu não seria eu), Tiago orelhas (por estares sempre presente), restantes ursos do C.d.B.