Life History of Daphnia Galeata in a Hypertrophic Reservoir and Consequences of Non-Consumptive Mortality for the Initiation of a Midsummer Decline
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Freshwater Biology (2002) 47, 2313–2324 Life history of Daphnia galeata in a hypertrophic reservoir and consequences of non-consumptive mortality for the initiation of a midsummer decline S. HU¨ LSMANN and H. VOIGT Institute of Hydrobiology, Dresden Technical University, Dresden, Germany SUMMARY 1. Field and laboratory investigations were combined in a 2-year study on the initiation of a midsummer decline of Daphnia galeata Sars in a hypertrophic reservoir. Quantitative field samples were taken twice a week, and, adult and juvenile mortality rates were calculated. Patterns of reproduction and survival of daphnids born during spring and early summer under fluctuating food conditions were determined in life-table experi- ments. 2. The abundance of Daphnia increased strongly in early May and declined in June 1998 (midsummer decline). In 1999, Daphnia density increased only slowly in spring and remained constantly high throughout the summer. 3. Food conditions (concentrations of POC<30 lm) for daphnids deteriorated in both years in response to increasing Daphnia densities, resulting in a clear-water phase of about 4 weeks. When Daphnia abundance declined in 1998, POC<30 lm concentrations increased greatly, whereas in 1999 food conditions improved only slightly and Secchi depth remained high. 4. Survival of daphnids in life-table experiments decreased greatly after food became rare and was strongly reduced in those animals born during the clear-water phase compared with those born later. In addition, age at first reproduction was retarded during the clear- water phase, resulting in very low population growth rates. Survivorship patterns in life- table experiments suggest a strong impact of non-consumptive mortality on Daphnia population dynamics. 5. Field data of mortality point to differences in mortality patterns between years, probably resulting from different predation impacts of juvenile fish. In both years, however, adult mortality contributed substantially to overall mortality at the end of the clear-water phase. As bottom-up effects on D. galeata were very similar in both years, the significance of non-consumptive mortality on the initiation of midsummer declines appears to depend largely on recruitment patterns before the clear-water phase. A high impact can be expected when Daphnia populations are dominated by a peak cohort of nearly identical age during the clear-water phase. Keywords: clear-water phase, Daphnia galeata, life history, midsummer decline, mortality Introduction The relative importance of bottom-up and top-down Correspondence: Stephan Huulsmann,€ Netherlands Institute of influences on Daphnia population dynamics and the Ecology, Centre for Limnology, PO Box 1299, 3600 BG Maarssen, induction of midsummer declines of Daphnia abun- the Netherlands. E-mail: [email protected] dance has been the focus of numerous studies (e.g. Ó 2002 Blackwell Science Ltd 2313 2314 S. Huulsmann€ and H. Voigt Luecke et al., 1990; De Stasio et al., 1995; Mehner ment patterns of D. galeata Sars observed in a year et al., 1998a). Predation has been suggested to be the with a midsummer decline (Huulsmann€ & Weiler, main driving force by some investigators (Mills & 2000). Forney, 1983; Cryer, Peirson & Townsend, 1986); Meaningful extrapolation of age-specific mortality others have stressed on the importance of bottom-up patterns from laboratory populations to field situa- effects (Lampert et al., 1986; Boersma, van Tongeren tions is impracticable. Conversely, death rates calcu- & Mooij, 1996). However, only in a few cases could lated from field data encompass all sources of the predominant role of predation be demonstrated mortality and are confounded by large uncertainties by detailed comparisons of consumption and mor- (George & Edwards, 1974; Taylor, 1988). An approach tality patterns (Mills & Forney, 1983; Vijverberg simulating in situ conditions while precluding preda- et al., 1990). Consequently, many investigators con- tion hence would be preferable for estimating the cluded that the combined effects of food limitation importance of non-consumptive mortality in field and predation drive Daphnia populations to decline populations. Most studies that have analysed Daphnia in summer (Luecke et al., 1990; Wu & Culver, 1994; life-history patterns under in situ conditions focused Mehner et al., 1998a). It remains largely speculative, on growth or reproduction (Weglenska, 1971; however, as to how exactly these interactions work, Threlkeld, 1979, 1980, 1985; Langeland, Koksvik & although effects of size-selection and timing of top- Olsen, 1985; Larsson et al., 1985; Muuller-Navarra€ & down and bottom-up factors might be especially Lampert, 1996) and neglected mortality. In situ experi- important (Post et al., 1997; Mehner et al., 1998b; ments related to midsummer declines (Threlkeld, Benndorf et al., 2001). 1979, 1985; Larsson et al., 1985) focused only on the Bottom-up effects influence Daphnia dynamics in actual decline phase. If, however, non-consumptive different ways. Starvation-induced mortality may be mortality of adults and preadults is important for the expected especially in juvenile stages (Threlkeld, 1976; decline, the life history of the daphnids born during Tessier et al., 1983) and has been suggested as the the build-up of the population or during the period of main cause of midsummer declines in several lakes high Daphnia abundance would be most critical. (e.g. Boersma et al., 1996), although this mechanism Gries & Guude€ (1999) recently gave evidence of the might not hold for every species and situation significance of non-consumptive mortality for the (Matveev & Gabriel, 1994). Reduced fecundity owing dynamics of D. galeata in Lake Constance. High to food limitation during the spring clear-water phase population losses resulting from sedimentation were is commonly regarded as key mechanism (Lampert attributed to an unidentified infection. However, size– et al., 1986; Sommer et al., 1986). This mechanism is frequency distributions of D. galeata in that study difficult to demonstrate, however, because birth and point to ageing effects as a partially alternative death rates must be considered simultaneously and explanation. That is, it is possible that the infection both parameters are related in a complex way invol- ‘only’ influenced reproduction, whereas mortality ving time lags (e.g. George & Edwards, 1974). The resulted simply from senescence. question remains therefore: Which part of the popula- The exact mechanism leading to midsummer tion dies during a decline and why? Although declines of Daphnia remains speculative as long as mortality of older and larger specimens of Daphnia is the life history under natural conditions is not commonly attributed to selective predation (Gliwicz known. Therefore, the main goal of this study was & Pijanowska, 1989), the synchronous die-off of a to elucidate the life-history pattern of D. galeata peak-cohort might also be an important source of during spring and early summer in a reservoir. Data adult mortality and contribute to the induction of a from life-table experiments approximating field con- midsummer decline (Huulsmann€ & Weiler, 2000; ditions were related to population dynamics and Benndorf et al., 2001). The proposed mechanism demography of D. galeata between May and July in involves a reduced mean life span of a peak cohort two successive years, one with a midsummer decline caused by interactions between starvation-induced and one without. The mortality of juveniles and and age-specific mortality (Threlkeld, 1976). Evidence adults was calculated from both field data and for this idea came from the analysis of long-term data estimates of juvenile growth, and compared with (Benndorf et al., 2001), and the mortality and recruit- Daphnia survival in life-table experiments to get Ó 2002 Blackwell Science Ltd, Freshwater Biology, 47, 2313–2324 Life history of Daphnia galeata Sars 2315 insight into the relative importance of non-con- where Nti+1,obs., Nti,obs. is observed abundance at ti+1 sumptive and consumptive mortality for the initi- and ti, respectively and Nti+1,calc., the calculated )1 ation of a midsummer decline. abundance at ti+1. Mortality rates (m) (day )of juvenile and adult daphnids were calculated by subtracting r from r¢, which results in: Methods À 1 m ¼ðln Nti þ 1;calc: À ln Ntiþ1;obs:ÞDt ðday Þð3Þ Field sampling and determination of population parameters Life-table experiments Daphnia galeata and particulate organic carbon A total of nine life-table experiments were per- (POC l ) were sampled twice a week from May <30 m formed in 1998 and 1999. Large, fecund individuals to July 1998 and 1999 in the pelagic zone of Bautzen of D. galeata from Bautzen Reservoir were obtained Reservoir, Germany. Samples were taken with a tube by net tows (780 lm) and kept in filtered reservoir sampler (diameter 95 mm, 2 L; Limnos, Finland) at water for 12 h. Six to ten neonates were then placed 1-m intervals over the whole water column. Samples individually in glass vessels (15 mL) filled with from three stations with a similar water depth reservoir water. Water was changed daily (1998) or (10–12 m) were pooled. Secchi depth was recorded every other day (1999), but gently stirred everyday. and water temperature was measured in depth At least three additional neonates were taken to intervals of 1 m. Daphnids from zooplankton samples estimate the initial length of the experimental were enumerated in at least three subsamples. About animals. All experiments were run at 18 ± 0.5 °C 100 individuals were measured and the number of and a 16 : 8 h light–dark cycle. eggs in the brood pouch, as well as their develop- Depth-integrated water samples from the reservoir mental stage (Threlkeld, 1979) was recorded. The were collected twice a week and stored in the dark proportion of adult daphnids was calculated after at 4 °C. In 1998, the water was passed over a 30-lm determination of the size at maturity (SAM) according screen to remove non-edible size fractions of the to Stibor & Lampert (1993). seston. In 1999, a 250-lm screen was used to mimic Juvenile growth was estimated once per week in more closely the food situation experienced by flow-through chambers at 18 °C.