Detailed information about the location and extent of zooplankton patches is fundamental to understand how abiotic and biotic forces interact to structure the spatial distribution of zooplankton. We mapped zooplankton patchiness in a Minnesota lake during spring, summer, and autumn with high-frequency (192-kHz) single-beam sonar. Conventional plankton samples of aggregations detected acoustically revealed that Daphnia pulicaria (mean body length 1.6 mm, mean target strength Ϫ120 dB) scattered most (ϳ63%) of the sound. Other taxa were smaller (Ͻ½ the length of D. pulicaria) and were usually less abundant and therefore scattered much less sound than D. pulicaria. Our acoustic estimates of Daphnia concentrations illustrate extreme patchiness, with concentrations varying by as much as four orders of magnitude over vertical distances of less than 1 m. Seasonal patterns of patchiness were related to predation by rainbow trout and to abiotic factors associated with stratification. Daphnia concentrations were highest from June to October in a deep-water ''refuge zone'' where oxygen concentrations were between 3 and 5 mg L Ϫ1 . These oxygen levels are suitable for Daphnia but are lower than those required by rainbow trout. Heterogeneity in Daphnia concentration along the lake's long axis was highest in May and June, when the population resided primarily in the oxic hypolimnion during the daytime. From July to October, as oxygen concentrations declined in the hypolimnion, the population became more metalimnetic and more uniformly distributed in the horizontal dimension. A diel study of the population in October indicated that the patchiness of population also changed dramatically between day and night. During the day the population aggregated densely in a thin layer (ϳ2 m thick) in the thermocline. After sunset the population dispersed into the epilimnion, where concentrations were ϳ100,000 m Ϫ3 less than they were during the day in the thermocline.Patchy spatial distributions and the low resolution of conventional sampling methods have impeded analyses of zooplankton populations. Zooplankton concentrations have been shown to vary by a factor of 1,000 within distances of meters horizontally or vertically, and the sampling resolution of conventional plankton nets and hoses is usually too coarse to identify the spatial limits of aggregations precisely (Coyle 2000). Variation in population density estimates due to sampling often cannot be distinguished from real changes of population density, and the effects of biological processes are difficult to distinguish from those of advective transport (Megard et al. 1997). Acoustic and optical plankton samplers developed during recent decades are major advances. They have very high sampling rates and spatial resolution, comparable to modern instruments (e.g., CTD profilers) used to measure environmental variables. Large numbers of plank-
Despite the central role of colonization to invasion biology, little is known about the early dynamics of founder populations because this time period is often unobserved and short-lived. This study documents with high resolution the early population ecology of an invading freshwater zooplankton, Bythotrephes longimanus (Cladocera : Cercopagidae), by measuring accumulation rates of its caudal spine in sediments. Using dated ( 210 Pb and 137 Cs) sediment cores from four spatially distinct sites in Island Lake Reservoir (Minnesota, USA), we describe its first presence, early distribution, growth trajectory, and early impacts on prey. The sediment record shows that B. longimanus first appeared and was widely distributed in the lake in 1982 (6 2 yr, standard deviation), 8 yr before its first detection in the water, making it one of the earliest documented invasions in North America and suggesting that ecosystems may serve as dispersal hubs for years prior to detection. Logistic growth models describing spine accumulation rates show that B. longimanus required about two decades to achieve an annual equilibrium (K). Prolonged buildup to K may owe to several factors, including accumulation of a sufficiently large bank of resting eggs, the obligate overwintering life stage. Early exponential growth was incongruent with the presence of a lag phase. Post invasion, Daphnia mendotae became proportionally the most abundant daphniid in the lake, but the timing of the switch in prey species composition coincided more with the proliferation of B. longimanus density and its attainment of K than with its arrival to the lake.
1. Stocking of lakes with rainbow trout is a common practice that presents a potential conflict for lake managers who must balance the interests of anglers with those concerned that zooplanktivory by trout may trigger a trophic cascade and result in decreased water clarity. 2. This study examined how the timing of trout stocking (autumn versus spring) in a Minnesota (U.S.A.) lake affected (i) the population dynamics of their zooplankton food supply (Daphnia pulicaria), (ii) phytoplankton biomass and water clarity and (iii) trout survival. Sizes of both Daphnia and trout populations were estimated acoustically with high-frequency (192 kHz) sonar. 3. Daphnia were nearly eliminated from the lake during winters after trout were stocked in autumn. In both of these years (1996 and 1997), the Daphnia population was small in the spring, and grew during the summer and into the autumn as the trout population diminished. 4. The lake was then stocked in spring for 2 years (1998 and 1999). This fisheries manipulation alleviated predation over the winter, but increased predation on D. pulicaria during the spring, summer and autumn. However, the high mortality caused by the spring-stocked trout was offset by even higher rates of reproduction by the relatively large populations of fecund Daphnia that survived the winter in 1998 and 1999. 5. Grazing by these dense populations of Daphnia produced clear-water phases during May and June that were inhibited in autumn stocking years. In addition, the large Daphnia populations present during the spring and early summer of 1998 and 1999 provided abundant forage for trout. 6. This fisheries manipulation achieved seemingly mutually exclusive management objectives: a robust planktivorous sport fishery, and clear water for other forms of recreation.
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