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Life Cycles and Secondary Production of Ephemeroptera, Plecoptera, and Trichoptera (Insecta) Under an Extreme Continental Climate (River Kharaa, Mongolia)

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Life Cycles and Secondary Production of Ephemeroptera, Plecoptera, and Trichoptera (Insecta) Under an Extreme Continental Climate (River Kharaa, Mongolia) Faculty of Environmental Sciences Life cycles and secondary production of Ephemeroptera, Plecoptera, and Trichoptera (Insecta) under an extreme continental climate (River Kharaa, Mongolia) Dissertation for awarding the academic degree Doctor rerum naturalium (Dr. rer. nat.) Submitted by MSc. Saulyegul Avlyush born on 10 December, 1982 in Darkhan-Uul province, Mongolia Reviewer: Prof. Dr. Dietrich Borchardt Helmholtz Centre for Environmental Research (UFZ) Department of Aquatic Ecosystem Analysis and Management Prof. Dr. Markus Weitere Helmholtz Centre for Environmental Research (UFZ) Department of River Ecology Prof. Dr. Rüdiger Wagner University of Kassel Institute for Biology Dresden, 2013 Defense date: 12 December 2013 Explanation of the doctoral candidate This is to certify that this copy is fully congruent with the original copy of the dissertation with the topic: „Life cycles and secondary production of Ephemeroptera, Plecoptera, and Trichoptera (Insecta) under an extreme continental climate (River Kharaa, Mongolia)“ Magdeburg, 24.05.2013 …………………………………………............ Place, Date Avlyush, Saulyegul ………………………………………….............. Signature (surname, first name) Acknowledgements The thesis described here was performed within the research and development project ‘Integrated Water Resources Management in Central Asia: Model Region Mongolia’ (IWRM- MoMo) funded by the German Ministry of Education and Research (BMBF Grant 0033L003A) under the “Research for Sustainability” Programme and supported by the German Academic Exchange Program-DAAD (A/07/99089). Foremost, I owe my deepest gratitude to my supervisor Prof. Dr. Dietrich Borchardt for giving me the chance to accomplish my Ph.D research under his supervision. He opened the door for me to find the diverse cultural environment of researchers and students and enjoyable scientific atmosphere at the Center for Environmental Systems Research (CESR), University Kassel and at the Department of Aquatic Ecosystem Analysis and Management (ASAM) at the Helmholtz Centre for Environmental Research (UFZ) in Magdeburg. I thank him especially for introducing me to the scientific branch known as aquatic ecology. Also for his assistance in my understanding of Mongolia’s unique ecological conditions in running water ecosystems and how different they are in Mongolia compared to European conditions. Further, special thanks for his time and countless hours of scientific discussions and proofreading of my research article and thesis. I also extend my sincere thanks to Dr. Ch. Javzan from the Institute of Geoecology, Mongolian Academy of Science and Prof. Dr. R. Samiya, and Prof. Dr. N. Soninkhishig from the National University of Mongolia for giving me the opportunities to work in this research field, encouraging my ideas and accepting my requests. I am indebted to all my current and former colleagues at CESR and UFZ including Daniel, Madlen, Philipp, Ralf, Rehan and other colleagues in the ASAM group for their friendship and encouragement. I am grateful to Melanie Hartwig and Michael Schäffer for the translation of my theses and summary into German and Andrew Kaus and Philipp Theuring for helping me correct my writing in English. I am very thankful to Michael Schäffer and Mario Brauns for the review of my thesis and giving valuable comments. I would like to express my special thanks to my handling supervisor, colleague and friend Michael Schäffer. I have got uncountable and invaluable knowledge from him. He accompanied me working in the field, analyzing the samples in the laboratory, analyzing the data and writing the research article. Beside the research work, he has shared in my happiness, troubles and sorrows. Further, I would like to express my appreciation to Ts. Batchuluun and N. Natsagnyam, A. Jargal, Emee, Zorigoo as well as several Mongolian students for their help during the expeditions and laboratory analysis. I sincerely thank B. Byambajav and Ts. Otgonkhuu and I call them as sisters. Thank you very much for your moral support and patience when I have expressed different personal characteristics of mine. I also wish to thank all of my friends for being there for me when I needed them; they all know who they are. i Finally, and most importantly, my deepest appreciation and love to my parents, my sister and brother. I would never reach this far in my academic career without the support of my parents. I dedicate this dissertation to my sister Erkyegul, you are in my heart every day since you left us. I am never disillusioned from my deepest hope to meet you again one day. ii Theses 1. Since the 1990s water quality monitoring projects using aquatic insects or macroinvertebrates as bioindication in Mongolia has mostly occurred in rivers drainage to the Arctic Ocean. They have been conducted to identify different anthropogentic stressors and impacts upon these running water ecosystems. However, there are still knowledge gaps and uncertainties concerning the research of these macroinvertebrates, in particular, a life cycle study of representative species are one such section of information missing. Therefore, this study was conducted within the framework of the biomonitoring program in the Kharaa River Basin under the research aims of the project entitled ‘Integrated Water Resources Management (IWRM) in Central Asia: Model Region Mongolia (MoMo)’. 2. The specific aim for the study was to determine their life cycles and secondary production of selected species in the Kharaa River Basin, Mongolia, where these animals are exposed to harsh environment conditions. Mongolia’s climate is characterised by long, dry and cold winters and hot summers with low precipitation and a high number of clear, sunny days per year. An important hydrological characteristic of the Kharaa River is the occurrence of complete ice coverage from November to April. The seasonal variation in the Kharaa River has been documented by the continuous air and water temperature recordings during the sampling years of 2007 - 2009. Another character of the Kharaa River is the substrate composition which changes substantially from upstream to downstream. 3. The main challenges for the research project were selecting the most suitable methods for use in the field sampling campaigns as well as establishing biomonitoring criteria for the target species under the extreme harsh climatic conditions. The research also sorts to address the pre-existing taxonomical identification problems. Consequently, a multi- habitat quantitative sampling method, and emergence traps type ‘Model week’ were selected. Five specific traits were chosen as selection criteria from the literature, where the life cycles of numerous species were investigated under comparable conditions to this study. Based on those five distinct criteria, a total of 18 species from EPT group (Ephemeroptera, Plecoptera, and Trichoptera) were selected for deeper analysis. 4. The main purpose of this study was to provide basic autecological information of 18 EPT species in the Kharaa River Basin, Mongolia, focusing on (i) larval and emergence densities, (ii) life cycle periods, and (iii) secondary production and growth rates. The key hypothesis was to test whether or not EPT species in the Kharaa River have special adaptations to survive the harsh winter conditions in Mongolia and reduced secondary production. Diverse life cycles and estimations of density, secondary production and growth rates from field data showed a wide spectrum of structural responses to a variety of natural and anthropogenic stressors in the aquatic ecosystem. iii 5. The 3-year study (2007-2009) of the life cycles of eighteen selected species according to the larval body size distribution and emergence period showed five different life cycle types including strict univoltinism, a flexible life cycle, a short and a long life cycle. The results of these life cycles for the selected 18 species and their development periods were compared to other studies at different taxonomical levels. However, a comparison at a species level was only possible for one macroinvertebrate (Ephemeroptera: Ephemera orientalis) within the literature available. On the one hand, it indicated a characteristic biogeographic distribution of these species; while on the other hand, it illustrated a knowledge gap within the species description. Considering the above mentioned hypothesis, Ephemera orientalis had a flexible life cycle that displayed an extended larval development period in the Kharaa River. 6. The production parameters were available to estimate the total of 15 species based on data from one year of sampling. The estimated annual productions of larvae at three investigated sites in the upper and middle stretches of the river in the year 2009 was in the range from 0.08 g DW m-2 y-1 to 18.7 g DW m-2 y-1. Biomass was in the range from 10.6 mg DW m-2 to 907.8 mg DW m-2, and the annual production to biomass rate (P/B) was in the range from 0.6 y-1 to 23.5 y-1 respectively. The value from the secondary production estimates from 13 species was lower than pre-determined criteria for low production (5.4 g DW m-2 y-1). The growth rates were calculated only for univoltine summer and winter cycle species. According to the general classification of growth rate for macroinvertebrates, all univoltine winter cycle species were classified as species with a low growth rate (up to 1.4% day-1) and univoltine summer cycle species were slightly higher than the criteria range. 7. This thesis provided the first quantitative results on the life
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