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Biogeochemistry of Marine Phanerogams Soils

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Biogeochemistry of Marine Phanerogams Soils TESE DE DOUTORAMENTO BIOGEOCHEMISTRY OF MARINE PHANEROGAMS SOILS Nerea Piñeiro Juncal ESCOLA DE DOUTORAMENTO INTERNACIONAL PROGRAMA DE DOUTORAMENTO EN MEDIO AMBIENTE E RECURSOS NATURAIS SANTIAGO DE COMPOSTELA 2020 DECLARACIÓN DO AUTOR/A DA TESE Biogeochemistry of marine phanerogams soils D./Dna. Nerea Piñeiro Juncal Presento a miña tese, seguindo o procedemento axeitado ao Regulamento, e declaro que: 1) A tese abarca os resultados da elaboración do meu traballo. 2) De selo caso, na tese faise referencia ás colaboracións que tivo este traballo. 3) A tese é a versión definitiva presentada para a súa defensa e coincide coa versión enviada en formato electrónico. 4) Confirmo que a tese non incorre en ningún tipo de plaxio doutros autores nin de traballos presentados por min para a obtención doutros títulos. En Santiago de Compostela, 19 de Novembro de 2020 Asdo Nerea Piñeiro Juncal AUTORIZACIÓN DO DIRECTOR / TITOR DA TESE [Biogeochemistry of marine phanerogams soils] D. Antonio Martinez Cortizas e D. Miguel Ángel Mateo Mínguez INFORMA/N: Que a presente tese, correspóndese co traballo realizado por Dna. Nerea Piñeiro Juncal, baixo a miña dirección, e autorizo a súa presentación, considerando que reúne os requisitos esixidos no Regulamento de Estudos de Doutoramento da USC, e que como director desta non incorre nas causas de abstención establecidas na Lei 40/2015. De acordo co artigo 37 do Regulamento de Estudos de Doutoramento, declara tamén que a presente tese de doutoramento é idónea para ser defendida en base á modalidade de COMPENDIO DE PUBLICACIÓNS, nos que a participación do/a doutorando/a foi decisiva para a súa elaboración e as publicacións se axustan ao Plan de Investigación. En Santiago de Compostela, 20 de Novembro de 2020 Antonio Martinez Cortizas Miguel Ángel Mateo Mínguez This thesis is bases in the following manuscripts: Manuscript 1: Piñeiro-Juncal, N., Leiva-Dueñas, C., Serrano, O., Mateo, M.A., Martinez Cortizas, A., 2020. Pedogenic processes in a Posidonia oceanica mat. Soil Systems 4,18, 1-15. Contribution of the PhD student: Conceptualization, laboratory analysis, statistical analysis, data interpretation and manuscript writing. Manuscript 2: Piñeiro-Juncal, N., Díaz-Almela, E., Leiva-Dueñas, Deulofeu, O., Frigola, J., Soler, M., Martinez Cortizas, A., Giralt, S., García-Orellana, J., Mateo, M.A., Processes driving seagrass soils composition along the western Mediterranean: the case of the southeast Iberian Peninsula. Under review at Science of the Total Environment. Contribution of the PhD student: Conceptualization, statistical analysis, data interpretation and manuscript writing. Manuscript 3: Piñeiro-Juncal, N., Mateo, M.A., Holmer, M., Martinez Cortizas, A., 2018. Potential microbial functional activity along a Posidonia oceanica soil profile. Aquatic Microbial Ecology, 81, 189-200. JCR IF (2018): 2.788, Q1 in Marine and fresh water biology Contribution of the PhD student: Laboratory analysis, statistical analysis, data interpretation and manuscript writing. Manuscript 4: Piñeiro-Juncal, N., Leiva-Dueñas, C., Serrano, O., Mateo, M.A., Martinez Cortizas, A., 2020. Cover loss in a seagrass Posidonia oceanica meadow accelerates soil organic matter turnover and alters soil prokaryotic communities. Organic Geochemistry. JCR IF (2019): 3.231, Q2 in Geochemistry and Geophysics Contribution of the PhD student: Part of the laboratory and statistical analysis, data interpretation and manuscript writing. Manuscript 5: Piñeiro-Juncal, N., Martinez Cortizas, A., Mateo, M.A. Reviewing soil seagrass data for its preliminary description and classification. In preparation to be submitted. Contribution of the PhD student: Conceptualization, review and manuscript writing. NEREA PIÑEIRO JUNCAL TABLE OF CONTENTS RESUMO ...................................................................................................... 5 SUMMARY ................................................................................................ 17 GENERAL INTRODUCTION ................................................................. 25 1. Marine phanerogams: the seagrasses ............................................... 25 2. Seagrass meadows and soil diagenesis ............................................. 27 3. Seagrass substrates as soils .............................................................. 29 4. Implications of soil recognition ........................................................ 34 GENERAL OBJECTIVES ........................................................................ 37 METHODS ................................................................................................. 39 GENERAL DISCUSION ........................................................................... 45 GENERAL CONCLUSIONS .................................................................... 59 RECOMMENDATION FOR FUTURE RESEARCH ........................... 61 GENERAL REFERENCES ...................................................................... 63 CHAPTER I ................................................................................................ 91 CHAPTER II .............................................................................................. 95 CHAPTER III ........................................................................................... 153 CHAPTER IV ........................................................................................... 155 CHAPTER V ............................................................................................ 157 APPENDIX. SUPPLEMENTARY MATERIAL .................................. 179 1. Chapter II ..................................................................................... 179 2. Chapter IV .................................................................................... 219 3. Chapter V ...................................................................................... 249 ACKNOWLEDGMENTS ....................................................................... 319 3 RESUMO 4 NEREA PIÑEIRO JUNCAL RESUMO As fanerógamas mariñas, ou herbas mariñas, forman parte dunha segunda colonización das augas costeiras pouco profundas por parte de plantas superiores. A recolonización levouse a cabo por tres camiños evolutivos diferentes, pero todas as herbas mariñas desenvolveron adaptacións semellantes á vida no mar, sinalando a presión ambiental como o principal factor de selección evolutiva. Como os cetáceos, volveron ao mar levando consigo características propias dos seus devanceiros terrestres. Algúns destes trazos morfolóxicos e fisiolóxicos, desenvolvidos para a súa adaptación á vida na terra, poden ter promovido a desenvolvemento de mecanismos diferentes a aqueles empregados polas algas para a adaptación a algúns dos retos da vida mariña. Por exemplo, o reforzo dos seus tecidos con lignina para manter a posición ergueita fóra da auga pode ter dado soporte ao desenvolvemento do sistema lacunar, o cal permite ás herbas mariñas 2+ 2+ bombear O2 á súa rizosfera para reducir fitotoxinas como o Fe , Mn ou os sulfuros. O desenvolvemento de órganos soterrados é un dos principais trazos das plantas terrestres e o trazo máis salientable das herbas mariñas desde o punto de vista da interacción co substrato onde medran. Os prados mariños están distribuídos nas áreas costeiras de todos os continentes agás da Antártida. As seus doseis promoven a deposición de sedimento e, tanto a dosel como os seus órganos subterráneos, reducen a resuspensión deste sedimento, promovendo a formación de solos ben estruturados seguindo una secuencia cronolóxica onde máis profundo é equivalente a máis vello. Os cambios promovidos pola planta no substrato ocorren na rizosfera e poden variar coa madurez da pradaría mariña. Os rizomas das herbas mariñas e as puntas das súas raíces liberan O2 e compostos orgánicos cara ao substrato, promovendo a formación de micronichos bioxeoquímicos ao seu redor. As herbas 5 RESUMO mariñas non son as únicas que promoven estes cambios no substrato baixo o prado; macro e microinvertebrados así como comunidades de microorganismos tamén xogan un papel na diaxénese destes solos. As herbas mariñas do xénero Posidonaceae desenvolven solos cunha alta carga orgánica, coñecidos como matas. O papel destas matas como reservas de carbono a longo prazo ten promovido un grande interese na comunidade científica sobre o destino da materia orgánica soterrada nos solos das herbas mariñas. A acumulación de materia orgánica nestes solos ten sido relacionada con (1) a promoción da retención de partículas debido ao dosel das herbas mariñas, (2) as condicións anóxicas que se acadan dentro do solo e (3) o carácter refractario dos tecidos das herbas mariñas, que poden ser preservados durante milleiros de anos. Tradicionalmente, os substratos baixo os prados de herbas mariñas teñen sido considerados como sedimentos. Con todo, co paso do tempo, a presenzas das herbas mariñas transforma radicalmente estes substratos mediante adicións, subtraccións, transferencias e transformacións (i.e., procesos de formación de solo) dunha mera acumulación de sedimentos a un solo. A definición de solo do Unite States Department of Agriculture Soil Taxonomy recoñece a posibilidade da formación de solos baixo as herbas mariñas dende a súa primeira edición. Non obstante, os solos subacuáticos teñen sido ignorados en gran medida na investigación das ciencias do solo. Por outra banda, os solos subacuáticos son apenas recoñecidos baixo a World Reference Base for Soil Resources até os dous metros de profundidade de auga (medido
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