Sexual Reproduction in Chydorid Cladocerans (Anomopoda, Chydoridae) in Southern Finland – Implications for Paleolimnology Liisa Nevalainen

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Sexual Reproduction in Chydorid Cladocerans (Anomopoda, Chydoridae) in Southern Finland – Implications for Paleolimnology Liisa Nevalainen Sexual reproduction in chydorid cladocerans (Anomopoda, Chydoridae) in southern Finland – implications for paleolimnology Liisa Nevalainen Academic dissertation To be presented, with the permission of the Faculty of Science of the University of Helsinki, for public criticism in lecture room E204 of Physicum, Kumpula, on November 14th, 2008 at 12 o’clock Publications of the Department of Geology D16 Helsinki 2008 Ph.D. thesis No. 203 of the Department of Geology, University of Helsinki Supervised by: Dr. Kaarina Sarmaja-Korjonen Department of Geology University of Helsinki Finland Reviewed by: Prof. Krystyna Szeroczyńska Institute of Geological Sciences Polish Academy of Sciences Poland Dr. Milla Rautio Department of Environmental Science University of Jyväskylä Finland Discussed with: Dr. Marina Manca The National Research Council (CNR) Institute of Ecosystem Study Italy Cover: morning mist on Lake Hampträsk ISSN 1795-3499 ISBN 978-952-10-4271-3 (paperback) ISBN 978-952-10-4272-0 (PDF) http://ethesis.helsinki.fi/ Helsinki 2008 Yliopistopaino Liisa Nevalainen: Sexual reproduction in chydorid cladocerans (Anomopoda, Chydoridae) in southern Finland - implication for paleolimnology, University of Helsinki, 2008, 54 pp. + appendix, University of Helsinki, Publications of the Department of Geology D16, ISSN 1795-3499, ISBN 978-952-10-4271-3 (paperback), ISBN 978-952-10-4272-0 (PDF). Abstract The relationship between sexual reproduction of littoral chydorid cladocerans (Anomopoda, Chydoridae) and environmental factors in aquatic ecosystems has been rarely studied, although the sexual behavior of some planktonic cladocerans is well documented. Ecological monitoring was used to study the relationship between climate-related and non-climatic environmental factors and chydorid sexual reproduction patterns in nine environmentally different lakes that were closely situated to each other in southern Finland. Furthermore, paleolimnological ephippium analysis was used to clarify how current sexual reproduction is reflected in surface sediments of the same nine lakes. Additionally, short sediment cores from two of the lakes were studied with ephippium analysis to examine how recent climate-related and non-climatic environmental changes were reflected in chydorid sexual reproduction. Ephippium analysis uses the subfossil shells of asexual individuals to represent asexual reproduction and the shells of sexual females, i.e. ephippia, to represent sexual reproduction. The relative proportion of ephippia of all chydorid species, i.e. total chydorid ephippia (TCE) indicates the relative proportion of sexual reproduction during the open-water season. This thesis is part of the EPHIPPIUM-project which aims to develop ephippium analysis towards a quantitative climate reconstruction tool. To be able to develop a valid climate model, the influence of the environmental stressors other than climate on contemporary sexual reproduction and its reflection in sediment assemblages must be clarified so they can be eliminated from the model. During contemporary monitoring a few sexual individuals were observed during summer, apparently forced to sexual reproduction by non-climatic local environmental factors, such as crowding or invertebrate predation. Monitoring also revealed that the autumnal chydorid sexual reproduction period was consistent between the different lakes and climate-related factors appeared to act as the main inducers and regulators of autumnal sexual reproduction. However, during autumn, chydorid species and populations among the lakes exhibited a wide variation in the intensity, induction time, and length of autumnal sexual reproduction. These variations apparently act as mechanisms for local adaptations due to the genetic variability provided by sexual reproduction that enhance the ecological flexibility of chydorid species, allowing them to inhabit a wide range of environments. A large variation was also detected in the abundance of parthenogenetic and gamogenetic individuals during the open-water season among the lakes. On the basis of surface sediment samples, the general level of the TCE is ca. 3-4% in southern Finland, reflecting an average proportion of sexual reproduction in this specific climate. The variation in the TCE was much lower than could be expected on the basis of the monitoring results. This suggests that some of the variation detected by monitoring may derive from differences between sampling sites and years smoothed out in the sediment samples, providing an average of the entire lake area and several years. The TCE is always connected to various ecological interactions in lake ecosystems and therefore is always lake-specific. Hypothetically, deterioration of climate conditions can be detected in the TCE as an increase in ephippia of all chydorid species, since a shortening open-water season is reflected in the relative proportions of the two reproduction modes. Such an increase was clearly detected for the time period of the Little Ice Age in a sediment core. The paleolimnological results also indicated that TCE can suddenly increase due to ephippia of one or two species, which suggests that at least some chydorids can somehow increase the production of resting eggs under local environmental stress. Thus, some environmental factors may act as species-specific environmental stressors. The actual mechanism of the “increased” sexual reproduction seen in sediments has been unknown but the present study suggests that the mechanism is probably the increased intensity of gamogenesis, i.e. that a larger proportion of individuals in autumnal populations reproduce sexually, which results in a larger proportion of ephippia in sediments and a higher TCE. The results of this thesis demonstrate the utility of ephippium analysis as a paleoclimatological method which may also detect paleolimnological changes by identifying species-specific environmental stressors. For a quantitative TCE-based climate reconstruction model, the natural variation in the TCE of surface sediments in different climates must be clarified with more extensive studies. In addition, it is important to recognize the lakes where the TCE is not only a reflection of the length of the open-water season, but is also non-climatically forced. The results of ephippium analysis should always be interpreted in a lake-specific manner and in the context of other paleoecological proxies. Contents List of publications Author’s contribution to the publications 1. Introduction 8 1.1. Biology of cladocerans 8 1.2. Cladoceran reproduction 12 1.3. Cladocerans in paleolimnology 15 1.4. Ephippium analysis 17 1.5. Objectives of this thesis 19 2. Material and Methods 20 2.1. Sites 20 2.2. Sampling and sample analyses 23 2.2.1. Contemporary samples 23 2.2.2. Sediment samples 24 2.3. Data analysis 25 3. Results and Discussion 25 3.1. Chydorid distribution 25 3.1.1. Species occurrence in sweep net and surface sediment samples 25 3.1.2. First findings of intact Unapertura latens sp. n. 28 3.2. Monitoring of modern sexual reproduction 29 3.2.1. Timing of gamogenesis (I) 29 3.2.2. Intensity of autumnal gamogenesis (II) 31 3.2.3. Population sizes and abundance of gamogenesis (III) 35 3.3. Sexual reproduction reflected in sediments 37 3.3.1. Chydorid ephippia in the Hampträsk core (IV) 37 3.3.2. Chydorid ephippia in surface sediments (V) 38 3.3.3. Chydorid ephippia in the Pieni Majaslampi core (V) 40 4. Conclusions 41 Acknowledgements References Appendix Publications I-V List of publications This thesis is based on the following papers, which in the text are referred to by Roman numerals: I Nevalainen, L. and Sarmaja-Korjonen, K. 2008. Timing of sexual reproduction in chydorid cladocerans (Anomopoda, Chydoridae) from nine lakes in southern Finland. Estonian Journal of Ecology 57, 21-36. II Nevalainen, L. and Sarmaja-Korjonen, K. 2008. Intensity of autumnal gamogenesis in chydorid (Cladocera, Chydoridae) communities in southern Finland, with a focus on Alonella nana (Baird). Aquatic Ecology 42, 151-163. III Nevalainen, L. 2008. Parthenogenesis and gamogenesis in seasonal succession of chydorids (Crustacea, Chydoridae) in three low-productive lakes as observed with activity traps. Polish Journal of Ecology 56, 85-97. IV Luoto, T.P., Nevalainen, L. and Sarmaja-Korjonen, K. 2008. Multiproxy evidence for the ‘Little Ice Age’ from Lake Hampträsk, Southern Finland. Journal of Paleolimnology 40, 1097-1113. V Sarmaja-Korjonen, K., Nevalainen L. and Gąsiorowski, M. Chydorid ephippia in surface sediments and a short core from southern Finland – evidence of increased sexual reproduction caused by environmental forcing. Submitted to Journal of Paleolimnology. In addition to the original papers, this thesis includes previously unpublished material analyzed by the author. Author’s contribution to the publications I K. Sarmaja-Korjonen planned the study, L. Nevalainen performed the sampling and the sample and data analyses, L. Nevalainen and K. Sarmaja-Korjonen interpreted the results and prepared the manuscript II L. Nevalainen and K. Sarmaja-Korjonen planned the study, L. Nevalainen performed the sampling and the sample and data analyses, L. Nevalainen and K. Sarmaja-Korjonen interpreted the results and prepared the manuscript III L. Nevalainen planned the study, performed the sampling, the sample and data analyses, interpreted the results and prepared the manuscript IV K. Sarmaja-Korjonen, T.P. Luoto and L. Nevalainen planned the study, L. Nevalainen and T.P. Luoto performed the sampling, T.P. Luoto
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