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1.
Mysis diluviana is a major component of prey fish diets in the Great Lakes, so annual production of M. diluviana is important for understanding and modeling energy flow through Great Lakes food webs. However, only three lake-wide measurements of M. diluviana annual production in Lake Ontario are currently available (1971, 1990, 1995). During 2013, lake-wide coverage of Lake Ontario was achieved during four periods from April to November. Annual mean density and biomass of M. diluviana in 2013 were 99?#/m2 (SE: 8) and 318?mg?dw/m2 (SE: 28) – approximately half of values observed in 1990s. M. diluviana comprised 13–30% of offshore zooplankton biomass in each period. Reproduction peaked in fall, with mean brood size of 32 embryos (range: 11–49), at least 10% larger than in 1990s. Generation time was two years from embryo to initial reproduction. Growth rates were 0.052?mm/d for the age-0 cohort and 0.027?mm/d for the age-1 cohort. Age-0 growth rate was significantly higher than in 1980s–90s (0.035?mm/d). Annual production in 2013 was 0.85?g?dw/m2/yr (SE: 0.03) which was 30–40% of values observed in 1990 and 1995 (2.23 and 2.53?g/m2/yr). Annual production to biomass ratio (P/B) in 2013 was 2.65?/yr which was 80–85% of values observed in 1990 and 1995 (3.24 and 3.11?/yr), but this difference was not statistically significant. Our results suggest that changes in annual production over time can be estimated using changes in biomass over time and a mean P/B ratio.  相似文献   

2.
Lake Michigan was invaded by zebra mussels (Dreissena polymorpha) in the late 1980s and then followed by quagga mussels (D. bugensis) around 1997. Through 2000, both species (herein Dreissena) were largely restricted to depths less than 50 m. Herein, we provide results of an annual lake-wide bottom trawl survey in Lake Michigan that reveal the relative biomass and depth distribution of Dreissena between 1999 and 2007 (although biomass estimates from a bottom trawl are biased low). Lake-wide mean biomass density (g/m2) and mean depth of collection revealed no trend between 1999 and 2003 (mean = 0.7 g/m2 and 37 m, respectively). Between 2004 and 2007, however, mean lake-wide biomass density increased from 0.8 g/m2 to 7.0 g/m2, because of increased density at depths between 30 and 110 m, and mean depth of collection increased from 42 to 77 m. This pattern was confirmed by a generalized additive model. Coincident with the Dreissena expansion that occurred beginning in 2004, fish biomass density (generally planktivores) declined 71% between 2003 and 2007. Current understanding of fish population dynamics, however, indicates that Dreissena expansion is not the primary explanation for the decline of fish, and we provide a species-specific account for more likely underlying factors. Nonetheless, future sampling and research may reveal a better understanding of the potential negative interactions between Dreissena and fish in Lake Michigan and elsewhere.  相似文献   

3.
In the 1990s, the Lake Ontario ecosystem was dramatically altered due to continued invasions of exotic species including dreissenid mussels and predatory cladocerans. We describe the diet and biomass of prey in the stomachs of adult (≥ 109 mm TL) and sub-adult (< 109 mm TL) alewife (Alosa pseudoharengus) in 2004 and 2005 across seasons and depths and compare our results to data from 1972 to 1988. During 2004 and 2005, adult alewife consumed primarily zooplankton prey at bottom depth zones < 70 m and primarily Mysis at bottom depth zones > 70 m. Mysis dominated the diets of adult alewife in all seasons except during the summer of 2004 when zooplankton dominated. Mysis dominated the diets of sub-adult alewife during early and late spring and zooplankton dominated the diets in summer and fall. Bythotrephes and Cercopagis were observed in the diets of both sub-adult and adult alewife. Diporeia was observed only rarely in adult alewife diets. The biomass of prey in alewife stomachs varied seasonally and increased with bottom depth for adult alewife. Alewife diets in 2004–2005 differed from those in 1972 and 1988 with an increase in the prevalence of Mysis, and a decline in the prevalence of zooplankton. The biomass of prey in adult alewife stomachs declined in 2004 and 2005 compared to 1972 and 1988, at bottom depth zones < 70 m but not at bottom depth zones > 70 m suggesting reduced food availability closer to shore. We hypothesize that consumption levels at the shallower depth zones, as indicated by very low biomass of prey in alewife stomachs, may not be sufficient to sustain alewife growth. The increased prevalence of Mysis and common occurrence of predatory cladocerans in the diet of alewife means that alewife have shifted to a higher trophic position.  相似文献   

4.
We conducted a probability-based net tow sampling of Lake Superior in 2006 and compared the zooplankton biomass estimate with an estimate from laser optical plankton counter (LOPC) surveys. The net survey consisted of 52 sites stratified across three depth zones (0–30, 30–150, > 150 m). The LOPC tow surveys were extensive and spatially covered much of Lake Superior (> 1300 km of towing). The LOPC was field calibrated to Lake Superior zooplankton samples collected across the years of 2004 to 2006. The volume-weighted lake-wide zooplankton biomass determined by traditional net tows to 100-m sample depth was 20.1 (± 7.8 SD n = 52) mg dry-weight m− 3. The estimates varied by depth zones within the lake, where nearshore (0–30 m) estimates were highest and highly variable. Net sites for the LOPC field calibration were removed to allow for LOPC validation with independent nets; the resulting net-based estimate 20.0 (± 9.3 SD n = 38) mg dry-weight m− 3 and LOPC lake-wide estimate 19.1 (± 3.3 SD) mg dry-weight m− 3 agreed well. Consistency across survey methods for lake-wide estimates suggested that LOPC survey data provides a comparable assessment tool to traditional nets for collecting zooplankton biomass data. We briefly compare our results with some observed historical patterns. Onshore–offshore trends in zooplankton biomass concentrations were similar to the last major lake-wide survey in 1973. The LOPC provided high resolution data on zooplankton biomass distribution. Using simultaneously collected in situ sensor data, the LOPC zooplankton biomass distributions over horizontal and vertical space can be modeled as a function of temperature and fluorescence.  相似文献   

5.
The amphipod Diporeia spp. has historically been an important component of the benthic food web of the Laurentian Great Lakes. The Great Lakes Water Quality Agreement included its population density as an indicator of ecological condition for Lake Superior, with target values of 220–320 m?2 in nearshore areas (≤100 m depth) and 30–160 m?2 in offshore areas (>100 m). To assess the status of Diporeia in Lake Superior, we used a probability-based lake-wide survey design to obtain estimates of Diporeia density and biomass in 2006, 2011 and 2016. A PONAR grab sampler was used to collect Diporeia at 50–53 sites each year, with approximately half in the nearshore (<100 m depth) region of the lake and half in the offshore. The mean area-weighted lake-wide density was 395 ± 56 (SE) m?2 in 2006, 756 ± 129 m?2 in 2011, and 502 ± 60 m?2 in 2016. For all years, both density and biomass were greater in the nearshore than in the offshore stratum. The densities for 2006–2016 were 3–5 times higher than those reported from a lake-wide survey conducted in 1973 by the Canada Centre for Inland Waters. The severe declines in Diporeia populations observed in the other Great Lakes during recent decades have apparently not occurred in Lake Superior. Further research is needed to understand spatial and temporal variability of Diporeia populations in Lake Superior to enhance the utility of Diporeia density as an indicator of benthic condition.  相似文献   

6.
The lipid content of Mysis diluviana in the offshore region of southeastern Lake Michigan was determined in 2009 and 2010. Total lipids (mg) increased with Mysis length, but percent lipids (% dry weight) did not. Increases in lipids for juveniles (< 10 mm) were related to increases in structural lipids (i.e., phospholipids and sterols), whereas increases in total lipids for adults were related to increases in storage lipids (i.e., triacylglycerols). Percent lipids varied seasonally, and were in general, highest in spring or late fall for juveniles and small adults (< 15 mm). For the largest adults (16+ mm), % lipid content did not vary much over the year in 2009, but in 2010, % lipids peaked in July. The rate of lipid retention differed among cohorts of juveniles, but it did not differ among cohorts of adults. In 2009 and 2010 respectively, 14% and 40% of Mysis > 13 mm had lipid content < 14%, a value associated with a high percentage of docosahexaenoic acid that may reflect starvation for this size of Mysis. Food availability (chl a levels or zooplankton biomass) was not correlated with Mysis lipid content for any size group of Mysis. Seasonal gradients in food availability might not be large enough to elicit strong responses in lipid content of Mysis.  相似文献   

7.
Diporeia hoyi and Mysis relicta are the most important prey items of slimy sculpins (Cottus cognatus) in the Great Lakes. Slimy sculpins were collected from dreissenid-infested bottoms off seven Lake Michigan ports at depths of 27–73 m in fall 2003 to study their lake-wide diets. Relatively large dreissenid biomass occurred at depths of 37- and 46-m. Quagga mussels (Dreissena bugnesis) composed at least 50% of dreissenid biomass at Manistique, Saugatuck, and Sturgeon Bay. Mysis accounted for 82% of the sculpin diet by dry weight at eastern Lake Michigan while Diporeia composed 54–69% of the diet at western Lake Michigan and dominated the diets of slimy sculpins at all sites deeper than 46 m. In northern Lake Michigan, this diet study in new sites showed that slimy sculpin consumed more prey with low energy contents, especially chironomids, than Mysis and Diporeia in shallow sites (depth <55 m). We recommend diet studies on sedentary benthic fishes to be conducted along perimeters of the Great Lakes to observe changes in their diets that may be impacted by changing benthic macroinvertebrate communities.  相似文献   

8.
With the large Diporeia declines in lakes Michigan, Huron, and Ontario, there is concern that a similar decline of Mysis diluviana related to oligotrophication and increased fish predation may occur. Mysis density and biomass were assessed from 2006 to 2016 using samples collected by the Great Lakes National Program Office's biomonitoring program in April and August in all five Great Lakes. Summer densities and biomasses were generally greater than spring values and both increased with bottom depth. There were no significant time trends during these 10–11 years in lakes Ontario, Michigan, or Huron, but there was a significant increase in Lake Superior. Density and biomass were highest in lakes Ontario and Superior, somewhat lower in Lake Michigan, and substantially lower in Lake Huron. A few Mysis were collected in eastern Lake Erie, indicating a small population in the deep basin of that lake. On average, mysids contributed 12–18% (spring-summer, Michigan), 18–14% (spring-summer, Superior), 30–13% (spring-summer, Ontario), and 3% (Huron) of the total open-water crustacean biomass. Size distributions consisted of two peaks, indicating a 2-year life cycle in all four of the deep lakes. Mysis were larger in Lake Ontario than in lakes Michigan, Superior, and Huron. Comparisons with available historic data indicated that mysid densities were higher in the 1960s–1990s (5 times higher in Huron, 2 times higher in Ontario, and around 40% higher in Michigan and Superior) than in 2006–2016.  相似文献   

9.
Zooplankton distributions are patchy due to multiple physical, chemical, and biological processes, including diel vertical migration (DVM) behavior. Heterogeneity in the offshore environment is difficult to study with net tows, but newer technologies measure finer-scale distributions. Here, we use laser optical plankton counter (LOPC) data, informed by net tows, to study distributions and DVM of zooplankton in offshore Lake Michigan during July and September 2015. Water column (5–60 m) zooplankton biomass varied by an order of magnitude among transects and a factor of two within individual transects (6–19 km distances); transect coefficients of variation (SD/mean) ranged from 7 to 22% (~0.5 km scale). Horizontal patterns in zooplankton biomass varied among size groups but were consistent from day to night, suggesting that processes driving heterogeneity persist for hours to days. Fine-scale LOPC data show that zooplankton often aggregate in thin layers (1–3 m) within the metalimnion, a feature undetectable by coarser net sampling. Although DVM was not consistently observed, some patterns emerged. Small zooplankton including copepodites, diaptomids (Leptodiaptomus ashlandi, L. minutus), and Diacyclops thomasi often migrated to surface waters at night, and large zooplankton (Limnocalanus macrurus) migrated upward at night in most cases. Beam attenuation coefficient (proxy for phytoplankton biomass) was a significant predictor for zooplankton mean depth (p < 0.001) although it explained more of the variation for night data (R2 = 0.72) than day data (R2 = 0.53). The heterogeneity observed in zooplankton distributions has implications for planktivorous fish feeding in the offshore zone, as prey density varies greatly with depth.  相似文献   

10.
There have been few direct measurements of phytoplankton production made in Lake Michigan since invasive dreissenid mussels became established in the lake. Here we report the results of 64 measurements of phytoplankton primary production made in Lake Michigan during 2016 and 2017. We conducted two lake-wide surveys, one in the spring 2016 isothermal period and one after summer stratification in 2017 and examined seasonal production with bi-weekly sampling between May and November 2017 at an offshore station in the southwestern part of the lake. We assessed nearshore-offshore gradients by sampling at three transect locations on three occasions in 2017. Spring 2016 production and production:biomass (P:B) ratios (reflective of growth rates) were similar across the lake and were higher than those reported before dreissenid mussels became established, suggesting that despite decreases in phytoplankton biomass, growth rates remain high. Summer 2017 production and growth rates increased from south to north. Areal production in 2017 peaked in late summer. Mean 2017 summer production (499 ± 129 mg C m?2 day?1) was lower than values reported prior to the mussel invasion, and the fraction of total production occurring in the deep chlorophyll layer was about half that measured pre-mussels. At the offshore site picoplankton accounted for almost 50 % of the chlorophyll. As spring P:B ratios have increased and summer P:B and seston carbon:phosphorus ratios have not changed, we conclude that the decrease in phytoplankton production in Lake Michigan is due primarily to grazing by mussels rather than to stronger nutrient limitation.  相似文献   

11.
Adequate densities of zooplankton prey are critical for growth and survival of larvae of many fish species. Little information exists on the density of zooplankton in Great Lakes inshore areas during early spring, when larvae of important fishes rely on zooplankton. Reduced age-0 walleye recruitment and the absence of data on zooplankton availability for larval walleyes in northern Green Bay, Lake Michigan, led us to assess zooplankton densities during this critical spring period. We conducted biweekly vertical plankton tows in 2014–2016 near reefs and river plumes used by spawning walleyes for periods when larval walleyes were expected to be relying on zooplankton prey. Densities of zooplankton were well below literature values identified for good growth and survival of larval walleyes, averaging 1.5 individuals L−1 for all taxa and 0.12 individuals L−1 for large-bodied taxa across all sites and sampling dates. Various factors could contribute to the low density of zooplankton observed. We found low but significantly higher densities of cyclopoid copepods, nauplii, Bosmina, and total zooplankton at river mouth sites compared to open water sites. These results suggest that food availability for larval walleye in our study area was severely limiting which is consistent with the paucity of strong year classes observed since 2000. We suspect northern Green Bay has limited potential for producing strong year classes of walleyes under such conditions. Fishery managers working in unproductive waters should consider assessing the zooplankton community during critical periods to identify potential bottlenecks to reproductive success and larval fish survival.  相似文献   

12.
We examined three decades of changes in dreissenid populations in Lake Ontario and predation by round goby (Neogobius melanostomus). Dreissenids (almost exclusively quagga mussels, Dreissena rostriformis bugensis) peaked in 2003, 13 years after arrival, and then declined at depths <90 m but continued to increase deeper through 2018. Lake-wide density also increased from 2008 to 2018 along with average mussel lengths and lake-wide biomass, which reached an all-time high in 2018 (25.2 ± 3.3 g AFTDW/m2). Round goby densities were estimated at 4.2 fish/m2 using videography at 10 to 35 m depth range in 2018. This density should impact mussel populations based on feeding rates, as indicated in the literature. While the abundance of 0–5 mm mussels appears to be high in all three years with measured length distributions (2008, 2013, 2018), the abundance of 5 to 12 mm dreissenids, the size range most commonly consumed by round goby, was low except at >90 m depths. Although the size distributions indicate that round goby is affecting mussel recruitment, we did not find a decline in dreissenid density in the nearshore and mid-depth ranges where goby have been abundant since 2005. The lake-wide densities and biomass of quagga mussels have increased over time, due to both the growth of individual mussels in the shallower depths, and a continuing increase in density at >90 m. Thus, the ecological effects of quagga mussels in Lake Ontario are likely to continue into the foreseeable future.  相似文献   

13.
Zooplankton in a nearshore area in southeastern Lake Michigan displayed strong seasonal and depth-related trends in abundance (#/m3) during the period 1971 to 1977. Depth-related assemblages defined by principal component analysis varied with season. From mid-spring to mid-autumn, zooplankton assemblages were strongly related to water depth. In general, zooplankton were least abundant in the inner region (5–10 m) where small organisms such as nauplii, Asplanchna spp., and Bosmina longirostris and epibenthic organisms such as Eurytemora affinis dominated. Zooplankton were most abundant between 20 and 30 m, with a peak July 1972–1977 mean density of 87,000/m3. Larger zooplankton, including Cyclops spp., Diaptomus spp., Daphnia spp., and Eubosmina coregoni dominated the outer region (20–50 m). During autumn, the inner region was dominated by cladocerans while the outer region was dominated by copepods. After thermal stratification disappeared in late autumn and before the thermal bar moved beyond the survey area in early spring, zooplankton tended to occur in higher densities in the shallow depths of the survey area. Also, in contrast to the mid-spring to mid-autumn period, zooplankton occurred in similar compositions over the survey grid. Factors implicated in producing seasonal differences in abundance patterns include temperature regimes, phytoplankton standing stocks, and predation by fish and invertebrates.  相似文献   

14.
The meiobenthos of nearshore southeastern Lake Michigan was quantified by taking cores from three depths (11,17, and 23 m) at monthly intervals from May to November 1976–79. Total meiobenthic abundance ranged from 69,700/m2 to 1,300,000/m2 and total biomass ranged from 0.03 to 0.87 g/m2. Nematodes accounted for 80% of all individuals and 66% of the biomass. Most of the major groups peaked in late spring/summer, but some peaked in early spring and fall. With the exception of nematodes, mean annual densities varied from two-fold to twelve-fold at a given station. Harpacticoids, tardigrades, and ostracods tended to be more abundant at the deeper depths, while cyclopoids and cladocerans were less abundant. There was no consistent relationship between sampling depth and the abundance of nematodes and rotifers. Temporal and spatial variation in many of the groups appeared related to changes in the amount of bottom detritus. The overall macrobenthos:meiobenthos biomass ratio was 15:1 and ranged from 5:1 to 45:1 on an annual basis.  相似文献   

15.
Pontoporeia hoyi was the major food consumed by most sizes of deepwater sculpins (Myoxocephalus thompsoni) during spring and summer, 1981–82, at a 100-m station in southeastern Lake Michigan. There was no apparent seasonality in choices of food or volumes consumed. Mysis relicta made up less than 9% of the total food volume and averaged second to Pontoporeia volumetrically—a result which differed from previous investigations for Lake Michigan. Reduced consumption of Mysis in our study compared to previous work may be a function of lower prey density or station depth; mysids may have been farther above the lake bottom than at previously studied, more shallow stations. Ostracods were important in 6- and 7-cm fish diets. Mysis, chironomids, and fish eggs were only found in sculpins larger than 8 cm. Chironomid larvae, which typically burrow into sediments, and copepod and cladoceran zooplankton constituted a small percentage of the diet of all sculpins. These results indicated that the deepwater sculpins examined were primarily epibenthic predators at the sediment-water interface and apparently did not burrow for food or feed higher up in the water column.  相似文献   

16.
We examined seasonal dynamics of zooplankton at an offshore station in Lake Michigan from 1994 to 2003 and 2007 to 2008. This period saw variable weather, declines in planktivorous fish abundance, the introduction and expansion of dreissenid mussels, and a slow decline in total phosphorus concentrations. After the major expansion of mussels into deep water (2007–2008), chlorophyll in spring declined sharply, Secchi depth increased markedly in all seasons, and planktivorous fish biomass declined to record-low levels. Overlaying these dramatic ecosystem-level changes, the zooplankton community exhibited complex seasonal dynamics between 1994–2003 and 2007–2008. Phenology of the zooplankton maximum was affected by onset of thermal stratification, but there was no other discernable effect due to temperature. Interannual variability in zooplankton biomass during 1994 and 2003 was strongly driven by planktivorous fish abundance, particularly age-0 and age-1 alewives. In 2007–2008, there were large decreases in Diacyclops thomasi and Daphnia mendotae possibly caused by food limitation as well as increased predation and indirect negative effects from increases in Bythotrephes longimanus abundance and in foraging efficiency associated with increased light penetration. The Bythotrephes increase was likely driven in part by decreased predation from yearling and older alewife. While there was a major decrease in epilimnetic–metalimnetic herbivorous cladocerans in 2007–2008, there was an increase in large omnivorous and predacious calanoid copepods, especially those in the hypolimnion. Thus, changes to the zooplankton community are the result of cascading, synergistic interactions, including a shift from vertebrate to invertebrate planktivory and mussel ecosystem impacts on light climate and chlorophyll.  相似文献   

17.
Burrowing mayfly nymphs (Hexagenia spp.) were sampled monthly, September through October 1995 and April through August 1996, with a standard Ponar grab (538 cm2 jaw opening) at 16 stations in U.S. waters of Lake St. Clair. Annual production (production, P) was 0 to 477 mg dry weight/m2 at three stations where pollution and sediment grain-size distribution limited the population, and was 738 to 5,255 mg dry weight/m2 at the other 13 stations. The highest production value measured for Hexagenia in Lake St. Clair was about three times higher than the highest value reported for other areas in the northern United States and Canada (39° to 53° North latitude). The production-mean annual biomass (biomass, B) ratio (P/B) for Hexagenia in Lake St. Clair in 1995–96 was described by the straight line P = 2.4 B (R2 = 0.94). Adding published P/B data for other North American populations changed the relation only slightly to P = 2.5B (R2 = 0.96). A P/B ratio of 2.5 is consistent with the expected value for an aquatic insect with a 2-year life cycle and overlapping cohorts, and these data suggest this relation has general applicability for estimating production of Hexagenia in the northern United States and Canada. Size-class and seasonal partitioning of Hexagenia biomass and production were evident in the data. Both biomass and production were highest among nymphs 16.0 mm and larger, and biomass was highest in October and again in June, immediately before the annual emergence of subimagos. The large size of the mature nymphs and the concentration of biomass and production among the larger nymphs in the population is consistent with their importance in the diets of many fishes in the northern United States and Canada.  相似文献   

18.
To better understand zooplankton dynamics in Lake Ontario’s Toronto Harbour and adjacent coastal area (CA), we sampled zooplankton, phytoplankton, nutrients and physical parameters on six dates in 2016. Despite higher levels of nutrients, chlorophyll and primary production in the inner harbor (IH), the areas supported similar May to November zooplankton biomass (IH = 32 ± 7 and CA = 42 ± 10 mg/m3). IH values were much lower than other nutrient-enriched embayments in Lake Ontario, yet CA biomass was twice that of nearshore sites away from Toronto. Small zooplankton such as rotifers and Bosmina dominated IH; and large taxa (Daphnia, calanoids and predatory cladocerans) were more important in the CA. Daphnia, Bosmina, cyclopoids and calanoids were larger in the CA, and adult cyclopoids had higher egg ratios. This led to low annual IH production estimates for both cyclopoid and calanoid copepods. Total phosphorus and chlorophyll did not appear to regulate zooplankton biomass, but positive relationships were found with bacterial biomass in the IH and with temperature in the cool season. Atypically high fish planktivory rates likely suppressed larger IH zooplankton in 2016, allowing small, resilient Bosmina to flourish and contribute 84% of total production in the IH. Comparing 2016 data to previous zooplankton surveys revealed considerable inter-annual variation in proportions of Daphnia, Bosmina and predatory cladocerans over the 1994 to 2016 period, and the strong top-down controls observed in 2016 were not typical. Elevated microbial production may serve as an important alternate trophic pathway supporting cladoceran populations in Toronto Harbour.  相似文献   

19.
20.
In 1999, a long-term monitoring program in southwestern Lake Michigan captured the invasion of Cercopagis pengoi (fish hook flea), a predatory zooplankton native to the Ponto-Caspian region. This invasion provided an opportunity to examine both the responses of a population following establishment in a new area and the immediate response of the native community. We compared seasonal dynamics and several morphological and life history traits of C. pengoi both among and within the 4 years following its invasion into Lake Michigan. Samples collected from five nearshore sites in southwestern Lake Michigan from 1999–2002 indicated that the average density doubled from 75 individuals/ m3 in 1999 to 150 individuals/m3 in 2002. We found no evidence for rapid changes in body size or average clutch size as the population established. We did, however observe a shift in seasonal phenology, with the population occurring in the water column earlier each year. By 2002, Cercopagis pengoi appeared in the water column by July and persisted until early fall. We also compared the average monthly densities of several potential prey items in August in the years before (1998–1999) and after (2000–2003) establishment. Although we found a significant reduction in the average number of rotifers, the general trend was a decline in all zooplankton species. Given the number of previously established exotic species in this system, it may prove difficult to quantify the impact of this most recent addition on what is left of the native community.  相似文献   

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