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Ecosystem Modelling in the Eastern Mediterranean Sea: the Cumulative Impact of Alien Species, Fishing and Climate Change on the Israeli Marine Ecosystem

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Ecosystem Modelling in the Eastern Mediterranean Sea: the Cumulative Impact of Alien Species, Fishing and Climate Change on the Israeli Marine Ecosystem Ecosystem modelling in the Eastern Mediterranean Sea: the cumulative impact of alien species, fishing and climate change on the Israeli marine ecosystem PhD Thesis 2019 Xavier Corrales Ribas Ecosystem modelling in the Eastern Mediterranean Sea: the cumulative impact of alien species, fishing and climate change on the Israeli marine ecosystem Modelización ecológica en el Mediterráneo oriental: el impacto acumulado de las especies invasoras, la pesca y el cambio climáti co en el ecosistema marino de Israel Memoria presentada por Xavier Corrales Ribas para optar al título de Doctor por la Universidad Politécnica de Cataluña (UPC) dentro del Programa de Doctorado de Ciencias del Mar Supervisores de tesis: Dr. Marta Coll Montón. Instituto de Ciencias del Mar (ICM-CSIC), Barcelona, España Dr. Gideon Gal. Centro de Investigación Oceanográfica y Limnológica de Israel (IOLR), Migdal, Israel Tutor: Dr. Manuel Espino Infantes. Universidad Politécnica de Cataluña (UPC), Barcelona, España Enero 2019 This PhD thesis has been framed within the project DESSIM ( A Decision Support system for the management of Israel’s Mediterranean Exclusive Economic Zone ) through a grant from the Israel Oceanographic and Limnological Research Institute (IORL). The PhD has been carried out at the Kinneret Limnological Laboratory (IOLR) (Migdal, Israel) and the Institute of Marine Science (ICM-CSIC) (Barcelona, Spain). The project team included Gideon Gal (Kinneret Limnological Laboratory, IORL, Israel) who was the project coordinator and co-director of this thesis, Marta Coll (ICM- CSIC, Spain), who was co-director of this thesis, Sheila Heymans (Scottish Association for Marine Science, UK), Jeroen Steenbeek (Ecopath International Initiative research association, Spain), Eyal Ofir (Kinneret Limnological Laboratory, IORL, Israel) and Menachem Goren and Daphna DiSegni (Tel Aviv University, Israel). “In nature there are no rewards or punishments, there are consequences” “En la naturaleza no hay recompensas ni castigos, hay consecuencias” Bob Ingersoll ACKNOWLEDGEMENTS I would like to think about this thesis as a race, with accelerated and decelerated moments. It has been a quite long trip (5 years). During this time I lived in 4 different places, I worked in 4 research institutes, I met a lot of people and I made good friends. During all this time I have felt accompanied and I would like to thank to all the people that during one part of the trip or even all of it have contributed to the fact that today I am able to write this part of the thesis. I would like to mention all the people, so I apologize if I forget someone. Here it is my short tribute to all of them. First and foremost, this thesis would has not been possible without the opportunity that my supervisors, Dr. Marta Coll and Dr. Gideon Gal, gave me quite a long time ago. Thank you for all your support, guidance, encouragement and patience, allowing me to take a valuable experience in terms of research and personal enrichment. Thanks also to the members of the project team as well as co-authors on the different chapters of my thesis. Thank you for your time and advice on my work. Many thanks to Dr. Eyal Ofir for all the help during my stay in Israel, for all the discussions, for all the conversations, for all the trips, etc. Definitively, for being a friend. Many thanks extended also to Jeroen Steenbeek, who helped a lot during the thesis, especially for all the discussions and fixing Ecopath/Ecosim bugs. A special thanks to Dr. Sheila Heymans, who helped me on model development and kindly received in Oban. I am grateful to Dr. Stelios Katsanevakis, for the opportunity to participate in a European project, for my stay in Lesbos and for all your help and support in one of the chapters of the thesis. Many thanks to Dr. Dor Edelist and Dr. Menachem Goren for all the data on the study area and all the discussions, contributions, insightful comments and confidence they had in me. I want to warmly thank Dr. Chiara Piroddi, for all your lessons, valuable comments and discussions about Ecopath and ecological aspects. At the same time, I would like to thanks all the people who during my scientific training have brought some contribution, especially Dr. Josep Lloret and Dr. Isabel Palomera. Thanks for your advice, support, encouragement and your friendship. During the PhD, I had the fortune to meet many people. Thanks very much to my colleagues from the Kinneret Limnological Laboratory, especially Noam, Ruthy, Peleg, Yaron and Yohi. Many thanks to the colleagues and friends from Renewable Marine I Resources Department from the Institute of Marine Science (ICM-CSIC). Firstly, a special thanks to my office mates: Marc F., Morane and John. Thanks for your patience with your chatterbox office mate and all your help. A special thanks to Daniel Vilas, for our friendship, support, help, valuable comments… I also would like to thanks Marc B., Mariona and Cristina, for our smoking breaks, your friendship and great (and bad) moments in front of the ICM. I would like to express many thanks to Alejandro, Marta Albo, Ivan, Marta Carretón, Carol, Claudio, Núria, Dafni, Susana, Anabel, Vanesa, Amalia, Federico, Elena Lloret, Elena Fernández, Fernando, Joan Navarro, Lucia, David, Joan Mir, Alizée, Javi, and Diego. I am also deeply thankful to Natalia Serpetti for your help while I was in Oban and to Athanasios Evagelopoulus, Maria Sini and Vasilis Trygonis for your warm reception during my stay in Mytilene. A really warm thanks to all my friends. Per la nostre amistat, per tots els bons moments i sobretot pel vostre suport en els moments dolents. En primer lloc a la colla d’amics de tota la vida de Portbou. Sempre aprop (tot i la distancia), actuant com a vàlvules d’escapament i desconexió en aquest petit racó de la Costa Brava. Moltes gràcies Pedro, Carlos (i els teus fills Leo i Enzo), Natalia, Eric, Paula, Anna, Didi, Davinia, Antonio, Laura, Joel, Carrillo, Edgar, Borja, Saida, Abraham, Manel, ... També m’agradaria donar les gràcies als companys de pis que he tingut a Barcelona. Moltes gràcies Andreu, Pablo, Marta, Janire i Elena. A totes les amistats barcelonines que hem continuat mantenint la amistat: Jordi, Virginia, Jenifer, Miquel, Uriol, David, Nuria, Gemma, Iris, Salva, Anna, Nuria ... I com no, un agraïment especial també al grup “Bailones” . Per tots el moments compartits, riures, balls, sortides, activitats, ... Heu sigut una via d’escapament molt important durant la última part de la tesis. Moltes gràcies Roberto, Eli, Alba, Eva, Mel, Mónica, David, Oscar, Laia, Elena, Laura, Paula, Joan, Robert, Toni, … I would like to conclude thanking my family. Especialment als meus pares, al meu germà i a la meva àvia. Moltes gràcies per tot: per fer-me la persona que soc ara, per tots els valors que m’heu inculcat, pel vostre amor, confiança i recolzament incondicional. Per aprendre a lluitar, a créixer. Gràcies per estar sempre al meu costat. A vosaltres us ho dec tot! Mil gràcies! II Abstract ABSTRACT The ecosystems of the Eastern Mediterranean Sea have undergone significant ecological changes caused by multiple human stressors, including the impact of invasive alien species (IAS), overfishing and sea warming. Firstly, I reviewed modelling approaches that have been used to assess the impact of IAS in aquatic ecosystems. According to the review, multispecies/ecosystem mechanistic models dominated the applications, with dynamic and non-spatial models being the most prevalent. Most of the models included an additional human stressor, mainly fisheries, climate change and/or nutrient loading. In the review, I summarised the main features of these applications and analysed their capabilities and limitations. Based on my conclusions, I reflected on future directions of development and applications of suitable modelling tools. In the review, I found that the Ecopath with Ecosim (EwE) approach had been frequently used to assess the impacts of already established IAS. In addition, the review highlighted EwE capabilities to forecast existing, emerging and potential new IAS. Other available literature reviews had shown EwE capabilities in assessing impacts of fishing and climate change, the other two stressors included in this thesis. Therefore, as a second step in the thesis, I first developed two static Ecopath ecosystem models from EwE approach representing the food web of the Israeli Mediterranean coast in 1990s and 2010s to characterize the structure and functioning of the ecosystem and assess past and current impacts of IAS and fishing. I then used the time-dynamic Ecosim module of EwE to fit the 1990s model to available time series of observations between both periods and to explore the historical dynamics of the ecosystem considering the effects of IAS, fishing dynamics and sea warming. Finally, the time- dynamic ecosystem model was used to perform future simulations of ecosystem change. Particularly, and after interacting with key stakeholders, I assessed the effects of a new set of fishing regulations currently being implemented in Israel, future changes in sea temperature following IPCC (Intergovernmental Panel on Climate Change) scenario projections and the potential increases in IAS biomass in the future. Firstly, I investigated the impacts of the stressors separately, and then I combined them to evaluate their cumulative effects. V Abstract Results from the static Ecopath models highlighted that the Israeli marine ecosystem, despite productivity differences, shared some structural and functional traits with other Mediterranean ecosystems such as the dominance of the pelagic domain in term of flows, the important role of detritus through low trophic levels and the importance of the benthic-pelagic coupling. In both time periods investigated, the same keystone groups were identified with the exception of hake in 2010s, which showed a decline in its keystone role, and may indicate that hake had lost its ecological role due to population declines.
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