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Zooplankton Adaptation Strategies Against Fish Predation in Turkish Shallow Lakes

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Zooplankton Adaptation Strategies Against Fish Predation in Turkish Shallow Lakes ZOOPLANKTON ADAPTATION STRATEGIES AGAINST FISH PREDATION IN TURKISH SHALLOW LAKES A THESIS SUBMITTED TO THE GRADUATE SCHOOL OF NATURAL AND APPLIED SCIENCES OF MIDDLE EAST TECHNICAL UNIVERSITY BY ÜLKÜ NİHAN YAZGAN TAVŞANOĞLU IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN BIOLOGY DECEMBER 2012 Approval of the thesis: ZOOPLANKTON ADAPTATION STRATEGIES AGAINST FISH PREDATION IN TURKISH SHALLOW LAKES submitted by ÜLKÜ NİHAN YAZGAN TAVŞANOĞLU in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biology Department, Middle East Technical University by, Prof. Dr. Canan Özgen Dean, Graduate School of Natural and Applied Sciences Prof. Dr. Gülay Özcengiz Head of Department, Biology Prof. Dr. Meryem Beklioğlu Yerli Supervisor, Biology Department, METU Prof. Dr. Erik Jeppesen Co-supervisor, Biolociences-Freshwater Biology, University of Aarhus, Denmark Examining Committee Members: Assoc. Prof. Dr. Can Bilgin Biology Department, METU Prof. Dr. Meryem Beklioğlu Yerli Biology Department, METU Prof. Dr. Nuray Emir Akbulut Biology Department, Hacettepe University Prof. Dr. Ahmet Altındağ Biology Department, Ankara University Assoc. Prof. Ayşegül Gözen Biology Department, METU Date: I hereby declare that all information in this document has been obtained and presented in accordance with academic rules and ethical conduct. I also declare that, as required by these rules and conduct, I have fully cited and referenced all material and results that are not original to this work. Name, Last Name: Ülkü Nihan Yazgan Tavşanoğlu Signature: iii ABSTRACT ZOOPLANKTON ADAPTATION STRATEGIES AGAINST FISH PREDATION IN TURKISH SHALLOW LAKES Yazgan Tavşanoğlu, Ülkü Nihan PhD., Department of Biology Supervisor: Prof. Dr. Meryem Beklioğlu Yerli Co-Supervisor: Prof. Dr. Erik Jeppesen December 2012, 181 pages In this study, the factors influencing zooplankton community structure in Turkish shallow lakes were elucidated with four main approaches: (i) space-for-time substitution for shallow lakes using snap-shot sampling in 31 lakes along a latitudinal gradient; (ii) in-situ mesocosm experiments in eleven lakes along a latitudinal gradient using three sets of artificial plants systems; (iii)‘Habitat Choice’ laboratory experiments mimicking a ‘shallow littoral’ zone with plants and a ‘deeper pelagic’ zone with sediments testing the response of Daphnia magna to predation cues; and (iv) long-term monitoring data (1997-2011) from two interconnected lakes. Snap-shot and long-term monitoring showed that eutrophication has a strong influence on the zooplankton community via increased fish predation, nutrient loading and salinization. Here too the zooplankton community shifted towards a smaller sized profile, especially in lakes located at lower latitudes. Moreover, The laboratory and in-situ mesocosm experiments revealed that under predation risk Daphnia preferred to hide near sediment instead of using submerged plants as a refuge. Accordingly, in-situ mesocosm experiments revealed a predation pressure induced size structure shift towards small-medium sized zooplankton and calanoid copepods. iv The long-term monitored lakes experienced (i) drought-induced water level drop, leading to increased salinity and eutrophication, and consequent anoxic conditions and fish kill; as well as (ii) biomanipulation in the downstream. Both conditions resulted in major reduction in the top-down control of fish and ultimate predomination by large sized Daphnia spp. Nevertheless, the excessive exploitation of lakes and ongoing warming entail Turkish shallow lakes to become more eutrophic, making this study indicative for the Mediterranean region. Keywords: diel vertical/horizontal migration, size structure, water level changes, salinity, biomanipulation v ÖZ TÜRKİYE SIĞ GÖLLERİNDE BALIK AVLANMA BASKISI KARŞISINDA ZOOPLANKTONUN UYUM STRATEJİLERİ Yazgan Tavşanoğlu, Ülkü Nihan Doktora, Biyoloji Bölümü Tez Yöneticisi: Prof. Dr. Meryem Beklioğlu Yerli Ortak Tez Yöneticisi: Prof. Dr. Erik Jeppesen Aralık 2012, 181 sayfa Bu tezde, Türkiye’de bulunan sığ göllerdeki zooplankton komünite yapısını etkileyen etmenler dört ana yaklaşım kullanılarak açıklanmıştır: (i) enlemsel hat botunca 31 sığ gölün zaman yerine mekan yaklaşımı kullanılarak anlık fotoğraf çekme yöntemi ile örneklenmesi;(ii) enlemsel hat boyunca onbir gölde üçer set yapay bitki düzeneği kullanılarak gerçekleştirilen yerinde (in-situ) mezokozm deneyleri; (iii) ‘Habitat tercihi’ laboratuvar deneyleri bitki kullanılarak ‘sığ littoral’ bölgenin ve çökel kullanılarak ‘derin pelajik’ bölgenin taklit edilmesi ile Daphnia magna’ nın avcı sinyali karşısında vereceği tepki test edilmiştir; ve (iv) birbirine bağlı iki gölde yapılan uzun dönemli (1997-2011) izleme çalışması. Anlık fotoğraf çekme örneklemesi ve uzun dönemli izleme çalışmasına göre ötrofikasyonun balık avlanma baskısı, besin tuzu yüklemesi ve tuzlanmaya bağlı olarak zooplankton toplulukları üzerinde etkili olduğunu göstermiştir. Aynı şekilde düşük enlemlerdeki göllerde küçük vücutlu zooplankton türlerinin baskın olmasına neden olduğu tespit edilmiştir. Laboratuvar ve yerinde mezokozm deneyleri avlanma riski karşısında Daphnia’nın suiçi bitkiler yerine çökeli barınak olarak tercih ettiğini göstermiştir. Ayrıca yerinde mezokozm deneyleri avlanma baskısı karşısında vi zooplanktonun küçük-orta büyüklükteki türler ile calanoid copepod’ a doğru değiştiğini göstermiştir. İzleme çalışmasına göre göller (i) kuraklığa bağlı olarak meydana gelen su seviyesi düşmesi ile tuzluluğun artmasını ve ötrofikasyonu tetiklemiştir ve bunların sonucunda anoksik koşullar ve balık ölümleri ile; (ii) althavzada biyomanipulasyon etkisi altındadır. Bu koşullar balığın yukardan-aşağı kontrolünü azaltarak büyük vücutlu türlerin özellikle Daphnia spp. ’ın baskın duruma geçtiğini göstermiştir. Ancak göllerin aşırı kullanımı ve süregelen ısınma Türkiye’de bulunan sığ göllerin giderek daha ötrofik olmasına neden olması bu çalışmanın Akdeniz ikliminin hakim olduğu bölgelerdeki göller için belirleyici olabilir. Anahtar Kelimeler: yatay/dikey göç, boy dağılım yapısı, su seviyesi değişimleri, tuzlanma, biyomanipulasyon vii To scientist mothers and their children who are always waiting viii ACKNOWLEDMENTS I believe that “only science can lead us in our life”. However, this adventure of science would have been impossible without the people in my life. I didn’t know how my life was going to change with one knock on a door 6 years ago. Even though words are not enough to express my feelings, at first I want to express my gratitude to my supervisor Prof. Dr. Meryem Beklioğlu Yerli for her guidance, valuable suggestions and encouragement through this long way. I would also like to thank Prof. Dr. Erik Jeppesen, who is the “saint of the lakes”, for his great support, supervision and understanding during this study despite his extremely tight schedule. I want to thank TÜBİTAK (Project No: 105Y332, 110Y125), METU-BAP (BAP-07- 02-2010-00-01) and EU-FP-7 REFRESH (No: 244121) for their financial support. I want to thank Dr. Marianna Meerhoff and Dr. Sandra Brucett for their guidance and help in improving the chapters. I also want to express my gratitude to Prof. Dr. Ahmet Altındağ and Mrs. Liselotte Sander Johansson for helping me during the identification of zooplankton, which was the most painful part of this thesis. I want to express my gratitude to Arda Özen, Şeyda Erdoğan, Gizem Bezirci, Nur Filiz, Jan Coppens, Korhan Özkan, Jennifer Kalvenas for their helps and their friendship. Special thanks go to Eti Ester Levi (angel of the lab) for her endless patience, Tuba Bucak for answering my neverending questions and Sara Banu Akkaş for her precious friendship as well as being my “talking” English dictionary on gtalk. The movie “Hokkabaz” always reminds me of friendship, trust and hope; in other words Ayşe İdil Çakıroğlu. Thank you for being there every single part of this thesis. My deepest graditude goes to my fabulous big family (Yazgan’s, Tavşanoğlu’s, Cengiz’s, Gülan’s, Alnıaçık’s, Erdoğan’s, Turan’s, Ünlü’s, Us’s and Özat’s) for their neverending belief and love… Çağatay and Uygar Tavşanoğlu, the meaning of “Love” and “Devotion”. ix TABLE OF CONTENTS ABSTRACT…………………………………………………………………………iv ÖZ……………………………………………………………………………………vi ACKNOWLEDGEMENTS………………………………………………………….ix TABLE OF CONTENTS……………………………………………………………..x LIST OF TABLES…………………………………………………………………xiii LIST OF FIGURES………………………………………………………………...xiv CHAPTER 1 INTRODUCTION ............................................................................................... 1 1.1. Zooplankton in Shallow Lakes ...................................................................... 1 1.2. Influencing Factors on Zooplankton Community .......................................... 3 1.2.1. Hydrology ............................................................................................... 3 1.2.2. Salinity ................................................................................................... 4 1.2.3. Predation ................................................................................................. 5 1.3. Avoidane Strategies of Zooplankton.............................................................. 6 1.4. Possible Refuges in Shallow Lakes ............................................................. 10 1.4.1. Submerged Macrophyte ....................................................................... 10 1.4.2. Turbidity ..............................................................................................
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