Population Dynamics of Daphnia Galeata in the Biomanipulated Bautzen Reservoir: Life History Strategies Against Food Deficiency and Predation
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Institut für Hydrobiologie, Technische Universität Dresden Population dynamics of Daphnia galeata in the biomanipulated Bautzen Reservoir: life history strategies against food deficiency and predation Dissertation zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) an der Fakultät Forst-, Geo- und Hydrowissenschaften der Technischen Universität Dresden vorgelegt von Stephan Hülsmann aus Melle Dresden 2000 Gutachter: Prof. Dr. Jürgen Benndorf, Institut für Hydrobiologie, Technische Universität Dresden Prof. Dr. Winfried Lampert, Max-Planck-Institut für Limnologie, Plön PD Dr. Thomas Mehner, Institut für Gewässerökologie und Binnenfischerei, Berlin Tag der mündlichen Prüfung: 27. Februar 2001 Contents Table of contents Abstract.......................................................................................................................................1 1 General introduction...........................................................................................................3 2 Introduction to the biomanipulation experiment in Bautzen Reservoir..............................7 3 Overview: Bottom-up and top-down factors influencing population dynamics of Daphnia galeata in Bautzen Reservoir ..............................................................................9 3.1 Introduction.................................................................................................................9 3.2 Methods ....................................................................................................................10 3.3 Results ......................................................................................................................12 3.4 Discussion.................................................................................................................19 4 Food limitation of fecundity and reproduction of Daphnia galeata as determined in field samples..........................................................................................23 4.1 Introduction...............................................................................................................23 4.2 Methods ....................................................................................................................24 4.3 Results ......................................................................................................................27 4.4 Discussion.................................................................................................................36 5 Life history and juvenile growth of Daphnia galeata under natural food conditions ......45 5.1 Introduction...............................................................................................................45 5.2 Methods ....................................................................................................................46 5.3 Results ......................................................................................................................47 5.4 Discussion.................................................................................................................56 6 Demography and life history characteristics of Daphnia galeata as related to food conditions and age-0 fish predation .....................................................................61 6.1 Introduction...............................................................................................................61 6.2 Methods ....................................................................................................................62 6.3 Results ......................................................................................................................62 6.4 Discussion.................................................................................................................70 7 Mortality pattern of Daphnia galeata in Bautzen Reservoir: The importance of non-consumptive mortality ................................................................77 7.1 Introduction...............................................................................................................77 7.2 Methods ....................................................................................................................79 7.3 Results ......................................................................................................................82 7.4 Discussion.................................................................................................................88 8 General discussion and conclusions .................................................................................93 References...............................................................................................................................111 Contents Danksagung ............................................................................................................................127 List of publications .................................................................................................................129 Abstract Abstract The population dynamics and demography of Daphnia galeata was analysed in a five year study in the biomanipulated Bautzen Reservoir. Samples were taken two times a week during the period May-July in the pelagic zone of this highly eutrophic water. Major bottom-up and top-down factors were determined during the study period and analysed with regard to their influence on Daphnia dynamics and life history. Field data on fecundity and population structure of D. galeata were combined with results from life table and growth experiments performed under approximately in situ conditions to gain insight into the mechanisms leading to a midsummer decline of this cladoceran species which dominates the zooplankton community in Bautzen Reservoir. Two main patterns of Daphnia dynamics emerged: In years without a midsummer decline the population increased slowly in spring, starting from low densities. High water transparency was observed already during the build-up of the population of D. galeata. Despite considerable fluctuations, Daphnia abundance remained on a high level throughout summer. In years with a midsummer decline the population started from relatively high densities in early May and more than doubled during one week. Peak densities were reached before the clear-water stage emerged. At the end of this period the population declined to low values which lasted for the rest of the summer. Fecundity of the Population of D. galeata declined, whereas the mean egg volume increased at the beginning of the clear-water stage as a result of declining food levels. The size at maturity (SAM) remained high during this period. Additionally, juvenile growth was reduced and the age at maturity was retarded. Survival probability was low for those daphnids born shortly before or during the clear-water stage compared to those born later. It can be concluded from these results that recruitment to adult stages is strongly reduced during the clear-water stage. The end of this period is marked by an alternation in generations. Only at that time can SAM be reduced because the new generation of adults matures at a smaller size, carrying small eggs. A high impact of non-predatory adult mortality can be expected when the population is dominated by a strong peak-cohort during the clear-water stage according to recruitment patterns during the build-up of the population. The most drastic decline both of Daphnia abundance and SAM was observed in those years when the biomass of juvenile fish exceeded 20 kg ha-1 at the end of the clear-water stage. Due to gape-size limitation juvenile fish mainly feed on juvenile daphnids during this period and thus, they reinforce bottom-up effects on the Daphnia population. When fish change their size selection towards adult daphnids at the time when the new generation takes over, this seems to represent the worst case for the Daphnia population. Consequently, the timing between bottom-up effects and the feeding pressure of juvenile fish determines the extent of the decline. 1 Chapter 1 1 General introduction The genus Daphnia has attracted scientific interest ever since the 17th century (reviewed by Edmondson 1987) and is considered as a model organism in aquatic ecology (Lampert and Sommer 1993). The most obvious reason for this attractiveness is that daphnids dominate the zooplankton community in many temperate meso- to eutrophic water bodies. Apart from that, practical reasons may explain the popularity. Daphnia has an appropriate size for being used in laboratory cultures, it is easy to cultivate and has short generation times. As an (relatively) unspecific filter feeder, able to feed on a broad size range of particles in the water (Burns 1968; Wagner 1998), Daphnia is able to increase water transparency significantly. Uhlmann (1954) was the first to notice that high densities of Daphnia may turn waste water ponds from a turbid into a clear-water state due to their filtering activity. The high importance of daphnids for the development of a clear-water stage has been confirmed in many studies (see review by Lampert 1988 a). This potentially high grazing pressure of Daphnia on phytoplankton and other seston-particles has drawn considerable attraction to this particular group of cladocera in biomanipulation experiments. Biomanipulation is an ecotechnological strategy to improve water quality of eutroficated systems via top-down manipulation of the food web (Shapiro