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(Cnidaria, Hydrozoa) in Antarctic Waters

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(Cnidaria, Hydrozoa) in Antarctic Waters FACULTAT DE CIÈNCIES BIOLÒGIQUES PROGRAMA DE DOCTORAT EN BIODIVERSITAT (3001) Diversity, ecology and biogeography of Antarctic benthic hydrozoans: the Weddell Sea and the Scotia Arc as study cases. MEMÒRIA PRESENTADA PER JOAN JOSEP SOTO ÀNGEL, CANDIDAT AL GRAU DE DOCTOR PER LA UNIVERSITAT DE VALÈNCIA DIRECTOR: DR. ÁLVARO LUIS PEÑA CANTERO VALÈNCIA, MAIG DE 2017 iii El Dr. ÁLVARO LUIS PEÑA CANTERO, Professor Titular del Departament de Zoologia de la Facultat de Ciències Biològiques: AUTORITZA la presentació de la memòria titulada "Diversity, ecology and biogeography of Antarctic benthic hydrozoans: the Weddell Sea and the Scotia Arc as study cases" en compliment de la normativa vigent, i CERTIFICA que els resultats inclossos van ser obtinguts sota la seua direcció a l’Institut Cavanilles de Biodiversitat i Biologia Evolutiva per JOAN JOSEP SOTO ÀNGEL, i constitueixen la seua tesi per optar al grau de Doctor per la Universitat de València. I perquè així conste, signa el següent certificat. Dr. Álvaro Luis Peña Cantero València, 29 de maig de 2017 v Si són colonials, acrobàticament, embranquen l’art contemporani dels seus cossos i codifiquen la repartició d’oficis de les seues societats ancestrals. Si són solitaris, dansen sofisticadament a la deriva, vestits amb tela de paraigua, amb dramatisme agestual. Però tots senyalitzen el seu contacte amb jeroglífics urticants. Laia Fontana i Bria. Novembre 2015 Els nostres veïns submarins. Bestiari contemporani de la Mediterrània. The tree of life should perhaps be called the coral of life, base of branches dead; so that passages cannot be seen. Charles Darwin. 1837-1838 Notebook on transmutation of species. On ne voit bien qu'avec le cœur; l'essentiel est invisible pour les yeux. Antoine de Saint-Exupéry. Abril 1943 Le Petit Prince. vii Agraïments Ho reconec: he deixat aquesta secció per al final i molt probablement no manifeste del tot com d’agraït estic a qui, d’una manera o d’altra, ha compartit amb mi aquesta (llarga) temporada de ma vida. Allà vaig: En primer lloc, el meu més profund agraïment a Álvaro Peña: infinites gràcies per oferir-me l’oportunitat de fer aquesta tesi doctoral i per guiar-me durant el procés. Gràcies pel teu consell expert; pels colpets a l’esquena i pels tocs d’atenció (sí, gràcies també per això); per la paciència amb les deadlines, els sprints i amb totes-les-coses paral·leles. I sobretot gràcies per deixar-me recórrer camins desconeguts, i per confiar que aquest dia, finalment, arribaria. A la Conselleria d'Educació, Investigació, Cultura i Esport de la Generalitat Valenciana per recolzar el desenvolupament del projecte mitjançant la concessió d’un contracte pre-doctoral. Als companys que han compartit dependències a l’ICBiBE a l’inici, i al laboratori de Biologia Marina del Departament de Zoologia durant la darrera i més llarga etapa. A Cristina per ser un recolzament enorme, especialment aquestes últimes setmanes. A Mustapha, Elena, Hèctor, Alba, Marina, Júlia, Rebeca G, Núria, Rebeca L, Carlos, Iñaki, Andrea, Sara, Mar, Tamara i Joan entre altres, per les vivències del dia a dia. Un enorme gràcies a Rebeca Genís, per la magnífica il·lustració de la portada de la tesi. Als enamorats de la mar: les xabienques i els xabiencs de l’IROX (Xavi, Encarna, Amadeu, Javier...) i als companys fotosub (Vicente, Sergio, Alfredo, Julio, Juan...), per les vívides hores de blau compartides. Molt especialment a Pablo, Roberto i Alicia, per totes les experiències viscudes tant dins com fora de l’aigua. Per creure en un projecte tan bonic, de nom Bioblau, i donar el millor de vosaltres per dur-ho endavant. Gràcies al meu company d’hidrozous, de mostreigs, de patiments i èxits, Borja, per ser un sí constant, per les discussions científiques, pel seu gran suport informàtic, i per ajudar-me a relativitzar. Menció a banda mereixen Ferran i Romana. La meua admiració cap a vosaltres va molt més enllà del terreny acadèmic. Sou grans referents a molts nivells, però vos esmente ací per la vostra qualitat com a persones. Dir gràcies es queda molt curt! Hem compartit la il·lusió de fer créixer la ciència des de baix. M’haveu fet sentir des de l’inici com vosaltres: professional. Sempre, en qualsevol moment, haveu estat disponibles per a mi i per a molta altra gent. Gràcies per dignificar la docència (la de veritat), i gràcies per fer-me partícip. Espere de tot cor que sempre tinguem projectes comuns. Un càlid gràcies a la gent que està lluny però alhora està molt prop, especialment María i Melissa. Apareguéreu en ma vida molt abans que açò començara i co-definireu qui sóc i com sóc. S’acaba una fase, i nosaltres seguim junts, sempre endavant, sempre aprenent, per un món millor i més sa. Gràcies a la meua família (biològica) per fer l’esforç d’entendre què és el que faig, especialment a ma mare. No ho saps (o sí?) però t’admire: molt poca gent ha patit i alhora fet tant pels demés com tu. Gràcies per ensenyar-me a lluitar amb el cor i a no rendir-me mai. A Laia, per totes i cadascuna de les coses que enumere abans, i per moltíssimes altres coses que saps ben bé. Has sigut la meua companya de beca, de Departament, de docència, i de tesi, però també de cultura, sapiència, política i amistat. Aquesta tesi és també mèrit teu, perquè sense cap no hi ha tesi, i tu m’has ajudat a no perdre’l (Stop Moixon inclós). Gràcies i enhorabona a tu també. I per últim a tu, Luis. M’has fet retrobar el nord, fins acabar convertint-te en ell. Aquesta tesi va dedicada a tu. Gràcies per fer-me coneixedor del vertader significat de felicitat. ix Contents Abstract ............................................................................................................................. 1 Introduction ..................................................................................................................... 3 1. The Antarctic benthos: a song of ice and forest ............................................................... 3 2. Introducing the hydrozoans ............................................................................................... 5 3. The study of Antarctic benthic hydroids: past and present ........................................... 6 Objectives ...................................................................................................................... 21 Structure of the thesis .................................................................................................. 22 Chapter I: Benthic hydroids from the Scotia Arc...................................................... 23 Chapter II: Faunistic affinities of the benthic hydroids from Scotia Arc .............. 59 Chapter III: Aglaophenia in Antarctic waters ............................................................. 77 Chapter IV: Benthic hydroids from the Weddell Sea .............................................. 93 General discussion ..................................................................................................... 155 Conclusions ................................................................................................................. 161 Resum en valencià ...................................................................................................... 163 Appendices .................................................................................................................. 167 Appendix A: Original publication reprint ........................................................................ 167 Appendix B: Supplementary tables and figures .............................................................. 171 1 Abstract Benthic hydrozoans are one of the most speciose and characteristic taxa living in Antarctic waters, with highly diverse genera, such as Antarctoscyphus, Halecium, Oswaldella, Schizotricha, Staurotheca, and Symplectoscyphus, being common in the area. Our knowledge of these and other Antarctic hydrozoans has recently increased thanks to several studies, but much remains to be done in some areas of the Southern Ocean and with some particular taxa. In the present dissertation, traditional morphology, cnidome determination, multivariate analyses, and comprehensive literature searches were employed to improve our current understanding of the group, with a strong focus in the relevant regions of the Scotia Arc and Weddell Sea, providing novel data on the reproductive phenology, distribution, use of substrate, and epibionts of the studied species. A complete inventory of benthic hydrozoans from the Patagonian and Antarctic regions was produced, allowing the analysis of faunal affinities, endemism, and trends in the relevant scientific literature in the area. In the Scotia Arc and surrounding waters, 45 species belonging to 9 families and 20 genera were found, including the new species Halecium stoloniferum and Schizotricha discovery. Leptothecata was the dominant taxon. Multivariate analysis based on two similarity indexes showed that the Scotia Arc has greater affinity with Antarctica than Patagonia, also highlighting the effectiveness of the Polar Front as a major oceanographic barrier, and strongly supporting the placement of the Scotia Arc within the Antarctic region. In the Weddell Sea, 77 species belonging to 21 families and 28 genera were found in what constitutes the most prolific collection of Antarctic benthic hydroids hitherto analyzed. Leptothecata was again the dominant taxon, but there was a higher percentage of Anthoathecata species than in other collections from Antarctic waters. A new species, Aglaophenia baggins, was collected in the Weddell
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