Long-term response of a shallow, moderately flushed lake to reduced external phosphorus and nitrogen loading

1. The responses of nutrient concentrations, plankton, macrophytes and macrozoobenthos to a reduction in external nutrient loading and to contemporary climatic change were studied in the shallow, moderately flushed Lake Müggelsee (Berlin, Germany). Weekly to biweekly data from 1979 to 2003 were compared with less frequently collected historical data. 2. A reduction of more than 50% in both total phosphorus (TP) and total nitrogen (TN) loading from the hypertrophic (1979–90) to the eutrophic period (1997–2003) was followed by an immediate decline in TN concentrations in the lake. TP concentrations only declined during winter and spring. During summer, phosphorus (P) release from the sediments was favoured by a drastic reduction in nitrate import. Therefore, Müggelsee acted as a net P source for 6 years after the external load reduction despite a mean water retention time of only 0.1–0.16 years. 3. Because of the likely limitation by P in spring and nitrogen (N) in summer, phytoplankton biovolume declined immediately after nutrient loading was reduced. The formerly dominant cyanobacteria (Oscillatoriales) Limnothrix redekei and Planktothrix agardhii disappeared, but the mean biovolume of the N₂‐fixing species Aphanizomenon flos‐aquae remained constant. 4. The abundance of Daphnia spp. in summer decreased by half, while that of cyclopoid copepod species increased. Abundances of benthic macroinvertebrates (mainly chironomids) decreased by about 80%. A resource control of both phytoplankton and zooplankton is indicated by significant positive correlations between nutrient concentrations and phytoplankton biovolume and between phytoplankton and zooplankton biomass. 5. Water transparency in spring increased after nutrient reduction and resulted in re‐colonisation of the lake by Potamogeton pectinatus. However, this process was severely hampered by periphyton shading and grazing by waterfowl and fish. 6. Water temperatures in Müggelsee have increased in winter, early spring and summer since 1979. The earlier development of the phytoplankton spring bloom was associated with shorter periods with ice cover, while direct temperature effects were responsible for the earlier development of the daphnid maximum in spring.     

Meteorological forcing of plankton dynamics in a large and deep continental European lake

The timing of various plankton successional events in Lake Constance was tightly coupled to a large-scale meteorological phenomenon, the North Atlantic Oscillation (NAO). A causal chain of meteorological, hydrological, and ecological processes connected the NAO as well as winter and early spring meteorological conditions to planktonic events in summer leading to a remarkable memory of climatic effects lasting over almost half a year. The response of Daphnia to meteorological forcing was most probably a direct effect of altered water temperatures on daphnid growth and was not mediated by changes in phytoplankton concentrations. High spring water temperatures during ”high-NAO years” enabled high population growth rates, resulting in a high daphnid biomass as early as May. Hence, a critical Daphnia biomass to suppress phytoplankton was reached earlier in high-NAO years yielding an early and longer-lasting clear-water phase. Finally, an earlier summer decline of Daphnia produced in a negative relationship between Daphnia biomass in July and the NAO. Meteorological forcing of the seasonal plankton dynamics in Lake Constance included simple temporal shifts of processes and successional events, but also complex changes in the relative importance of different mechanisms. Since Daphnia plays an important role in plankton succession, a thorough understanding of the regulation of its population dynamics provides the key for predictions of the response of freshwater planktonic food webs to global climate change. Received: 15 February 1999 / Accepted: 23 August 1999     

Recovery of Potamogeton pectinatus L. stands in a shallow eutrophic lake under extreme grazing pressure

In shallow lakes, submerged macrophytes contribute to the stabilization of the clear water state. If lost, a number of mechanisms prevent re-colonization. Lake Müggelsee (730 ha) lost its submerged vegetation due to increasing eutrophication and switched to phytoplankton dominance in 1970. After the reduction of nutrient loading in 1990, Potamogeton pectinatus L. started re-colonizing the lake. During the following years, it spread at a mean rate of 2.5 ha per year to all available areas <80 cm depth. Between 1993 and 1999, decreasing maximum biomass indicated hampered growth. Exclosure experiments revealed that herbivory reduced the aboveground biomass by more than 90%. Both waterfowl and fish were found to contribute to the grazing pressure despite a low abundance of the known herbivorous fish species and waterfowl in spring and summer. Protection of stands against grazing resulted in higher biomass of shoots, whereas shoot and tuber density did not change. Both shading by phytoplankton and periphyton, as well as grazing pressure, prevented the submerged vegetation of Lake Müggelsee from developing back to a dense zone that contributed to the reduction of turbidity.     

Grazer control and nutrient limitation of phytoplankton biomass in two Australian reservoirs

1. Grazer and nutrient controls of phytoplankton biomass were tested on two reservoirs of different productivity to assess the potential for zooplankton grazing to affect chlorophyll/phosphorus regression models under Australian conditions. Experiments with zooplankton and nutrients manipulated in enclosures, laboratory feeding trials, and the analysis of in-lake plankton time series were performed. 2. Enclosures with water from the more productive Lake Hume (chlorophyll a = 3–17.5 μg l^–1), revealed significant zooplankton effects on chlorophyll a in 3/6, phosphorus limitation in 4/6 and nitrogen limitation in 1/6 of experiments conducted throughout the year. Enclosures with water from the less productive Lake Dartmouth (chlorophyll a = 0.8–3.5 μg l^–1), revealed significant zooplankton effects in 5/6, phosphorus limitation in 5/6 and nitrogen limitation in 2/6 of experiments. 3. While Lake Hume enclosure manipulations of the biomass of cladocerans (Daphnia and Diaphanosoma) and large copepods (Boeckella) had negative effects, small copepods (Mesocyclops and Calamoecia) could have positive effects on chlorophyll a. 4. In Lake Hume, total phytoplankton biovolume was negatively correlated with cladoceran biomass, positively with copepod biomass and was uncorrelated with total crustacean biomass. In Lake Dartmouth, total phytoplankton biovolume was negatively correlated with cladoceran biomass, copepod biomass and total crustacean biomass. 5. In both reservoirs, temporal variation in the biomass of Daphnia carinata alone could explain more than 50% of the observed variance in total phytoplankton biovolume. 6. During a period of low phytoplankton biovolume in Lake Hume in spring–summer 1993–94, a conservative estimate of cladoceran community grazing reached a maximum of 0.80 day^–1, suggesting that Cladocera made an important contribution to the development of the observed clear-water phase. 7. Enclosure experiments predicted significant grazing when the Cladocera/Phytoplankton biomass ratio was greater than 0.1; this threshold was consistently exceeded during clear water phase in Lake Hume. 8. Crustacean length had a significant effect on individual grazing rates in bottle experiments, with large Daphnia having highest rates. In both reservoirs, mean crustacean length was negatively correlated with phytoplankton biovolume. The observed upper limit of its variation was nearly twice as high compared to other world lakes.     

Interannual and between-site variability in the occurrence of clear water phases in two shallow Mediterranean lakes

It is widely accepted that clear water phases constitute a regular stage in the seasonal succession of plankton in dimictic lakes and reservoirs (i.e. PEG Model). The occurrence of such a phenomenon in Mediterranean shallow lakes is characterised by a marked interannual variability, which makes it difficult to establish reliable predictions on the dynamics and functioning of plankton in these ecosystems. In the present paper we analyse the factors influencing the occurrence of the clear water phases in the two shallow lakes of the Albufera of Adra, a coastal wetland region of south-eastern Spain: Lake Honda and Lake Nueva. Despite their geographical proximity, both lakes depicted large hydrological and limnological differences. Lake Honda is an epigenic and recharge lake that is strongly influenced by the hydrological conditions in its watershed, while Lake Nueva can be classified as a hypogenic and discharge lake and, as such, is less affected by the hydrological regime. In contrast, the morphometry, exposure and fetch of Lake Nueva make this ecosystem especially sensitive to wind forcing. Clear water phases in these shallow lakes were linked with periods of low thermal stability and the dominance of small-edible algae in the phytoplankton community, both of which allowed a Daphnia magna population to grow up and induce the algae collapse by grazing. In Lake Honda, those conditions were met during the spring of 2002 under the influence of intense rainfall-events, while in Lake Nueva the clear water phase was induced in the spring of 2003 by the occurrence of strong and frequent wind events. In both lakes, a relatively high water column thermal stability and the abundance of cyanobacteria early in the spring prevented the development of the Daphnia magna population and the occurrence of the clear water phase.     

Clear,crashing, turbid and back – long‐term changes in macrophyte assemblages in a shallow lake

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Causes of Daphnia midsummer decline in two deep meromictic subalpine lakes

1. Daphnia are key organisms in pelagic food webs, acting as a food resource for fish and predatory zooplankton and regulating phytoplankton through grazing. Its population dynamic follows regular seasonal patterns, with spring peaks followed by summer population declines (midsummer declines, MSDs). Midsummer declines show high inter-annual variation, which has been attributed to different causes. However, the mechanisms controlling the MSD remain poorly understood, especially in deep stratified lakes.
2. We tried to disentangle the factors causing Daphnia MSDs in Lake Lugano and Lake Iseo (in Switzerland and Italy), two deep peri-alpine lakes with similar trophic status and vertical mixing dynamics, characterised by phosphorus accumulation in the hypolimnion and variable mixing during late-winter turnovers.
3. Specifically, we assessed the effects of three different hypothetical pathways according to which: (1) winter air temperature controls MSDs by influencing mixing depth during turnovers and epilimnetic phosphorus replenishment; (2) vernal air temperature influences MSD by accelerating the timing of spring population peak; and (3) summer temperature influences MSDs by increasing fish predation. We assessed the relative strength of these pathways using structural equation modelling on long-term datasets for the two lakes (29 years for Lake Lugano and 19 years for Lake Iseo).
4. Between the hypothesised pathways, the one driven by winter air temperature (through P replenishment) influenced Daphnia abundance in spring in both lakes, but the effects propagated to summer Daphnia abundance only in Lake Lugano. Additionally, summer Daphnia abundance was influenced by the summer air temperature through a positive (although weak) effect. By comparison, vernal air temperature had no detectable effects on summer Daphnia abundance.
5. The results revealed marked differences between the meromictic study lakes and the shallow hypertrophic water bodies that were the focus of previous research on Daphnia MSD, and also between the two study lakes. The influence of epilimnetic P replenishment on the summer Daphnia abundance in Lake Lugano, which was recovering from past eutrophication, may have reflected the greater susceptibility of deep, stratified lakes to P depletion after spring compared to shallow hypertrophic lakes or reservoirs. This effect might not have been detected in Lake Iseo because P was more consistently depleted during the study period (i.e. variance in the predictor was too low to detect an effect).
6. This study highlighted the complexity of the effects of climate variability on Daphnia MSD in deep lakes, showing that the responses can differ even between two neighbouring lakes with similar vertical mixing dynamics and trophic status. At the same time, the results suggest that future increases in winter air temperature, caused by global warming, may cause critically low densities of Daphnia during spring and summer and compromise the ability of zooplankton to control phytoplankton biomass.

     

Uniform Temperature Dependency in the Phenology of a Keystone Herbivore in Lakes of the Northern Hemisphere

Spring phenologies are advancing in many ecosystems associated with climate warming causing unpredictable changes in ecosystem functioning. Here we establish a phenological model for Daphnia, an aquatic keystone herbivore based on decadal data on water temperatures and the timing of Daphnia population maxima from Lake Constance, a large European lake. We tested this model with long-term time-series data from two lakes (Müggelsee, Germany; Lake Washington, USA), and with observations from a diverse set of 49 lakes/sites distributed widely across the Northern Hemisphere (NH). The model successfully captured the observed temporal variation of Daphnia phenology in the two case study sites (r² = 0.25 and 0.39 for Müggelsee and Lake Washington, respectively) and large-scale spatial variation in the NH (R² = 0.57). These results suggest that Daphnia phenology follows a uniform temperature dependency in NH lakes. Our approach – based on temperature phenologies – has large potential to study and predict phenologies of animal and plant populations across large latitudinal gradients in other ecosystems.     

Abiotic drivers of interannual phytoplankton variability and a 1999–2000 regime shift in the North Sea examined by multivariate statistics

The Dutch coastal zone is a region of the North Sea with a marked interannual and long‐term abiotic and phytoplankton variability. To investigate the relationship between abiotic variability and phytoplankton composition, two routine water monitoring data sets (1991–2005) were examined. Multivariate statistics revealed two significant partitions in the data. The first consisted of interannual abiotic fluctuations that were correlated to Rhine discharge that affected the abundance of summer and autumn diatom species. The second partition was caused by a shift in the abiotic data from 1998 to 1999 that was followed by a shift in phytoplankton composition from 1999 to 2000. Important factors in the abiotic shift were decreases in suspended matter (SPM) and phosphate (DIP) concentrations, as well as in pH. The decrease in SPM was caused by a reduction in wind speed. The increase in water column daily irradiance from the decrease in SPM led to increases in the abundance of winter–spring species, notably the prymnesiophyte Phaeocystis globosa. Because wind speed is related to the North Atlantic Oscillation (NAO) index it was possible to correlate NAO index and P. globosa abundance. Only five abiotic variables representing interannual and long‐term variability, including Rhine discharge and NAO index, were needed to model the observed partitions in phytoplankton composition. It was concluded that interannual variability in the coastal phytoplankton composition was related to year‐to‐year changes in river discharge while the long‐term shift was caused by an alternating large‐scale meteorological phenomenon.     

Can overwintering versus diapausing strategy in <Emphasis Type="Italic">Daphnia</Emphasis> determine match–mismatch events in zooplankton–algae interactions?

Mismatches between predator and prey due to climate change have now been documented for a number of systems. Ultimately, a mismatch may have far-reaching consequences for ecosystem functioning as decoupling of trophic relationships results in trophic cascades. Here, we examine the potential for climate change induced mismatches between zooplankton and algae during spring succession, with a focus on Daphnia and its algal food. Whereas the development of an overwintering population of daphnids may parallel shifts in phytoplankton phenology due to climate warming, changes in the photoperiod–temperature interaction may cause the emerging population of daphnids to hatch too late and mismatch their phytoplankton prey. A decoupling of the trophic relationship between the keystone herbivore Daphnia and its algal prey can result in the absence of a spring clear water phase. We extended an existing minimal model of seasonal dynamics of Daphnia and algae and varied the way the Daphnia population is started in spring, i.e., from free swimming individuals or from hatching resting eggs. Our model results show that temperature affects the timing of peak abundance in Daphnia and algae, and subsequently the timing of the clear water phase. When a population is started from a small inoculum of hatching resting eggs, extreme climate warming (+6°C) results in a decoupling of trophic relationships and the clear water phase fails to occur. In the other scenarios, the trophic relationships between Daphnia and its algal food source remain intact. Analysis of 36 temperate lakes showed that shallow lakes have a higher potential for climate induced match–mismatches, as the probability of active overwintering daphnids decreases with lake depth. Future research should point out whether lake depth is a direct causal factor in determining the presence of active overwintering daphnids or merely indicative for underlying causal factors such as fish predation and macrophyte cover. Priority program of the German Research Foundation—contribution 5.     

Role of hydrology in development of a vernal clear water phase in an urban impoundment

1. Grazer control of phytoplankton and water transparency was responsive to interannual variations in river discharge and corresponding water retention time in Ford Lake, Michigan, U.S.A. 2. A simple mathematical model was developed to separate the effects of washout from in situ zooplankton population dynamics. The threshold reservoir flushing time for development of a large Daphnia population and corresponding clear water phase during May was about 20 days. Predation by Leptodora was insufficient to cause the subsequent decline of the Daphnia population. 3. Recruitment success by the zebra mussel Dreissena polymorpha appeared responsive to flushing time indirectly, through hydrological effects on the magnitude of the spring diatom crop.     

The role of environmental parameters in the structure of phytoplankton assemblages and cyanobacteria toxins in two hypereutrophic lakes

We evaluated the variability of cyanotoxins, water chemistry, and cyanobacteria communities in two hypereutrophic drowned river mouth lakes (Spring Lake and Mona Lake; summer 2006) in west Michigan, USA. Even with considerable geographical and watershed similarity, local variations in nutrient concentrations and environmental factors were found to influence the differences observed in cyanobacteria assemblages and cyanotoxins levels between the two lakes. Limnothrix sp. dominated the phytoplankton community in Spring Lake (82% of biovolume) and was negatively correlated with total phosphorus (TP) concentrations. Although Spring Lake was treated with alum during the previous year, Limnothrix sp. was able to bloom in the lower P environment. In contrast, the N₂-fixing cyanobacterium, Anabaena flos-aquae, dominated the phytoplankton in Mona Lake (64% of biovolume). N₂-fixing cyanobacteria dominance in Mona Lake was correlated with higher TP lower dissolved nitrogen levels. Cylindrospermopsis raciborskii was found in both systems; however, the toxin-producing polyketide synthetase gene was not present in either population. The higher TP in Mona Lake appeared to account for the 3-fold increase in cyanobacteria biovolume. Restoration plans for both lakes should include assessments of internal loading and continued phytoplankton monitoring to track the temporal distribution of cyanobacteria species and cyanotoxin concentrations.     

Climatic effects on the phenology of lake processes

Populations living in seasonal environments are exposed to systematic changes in physical conditions that restrict the growth and reproduction of many species to only a short time window of the annual cycle. Several studies have shown that climate changes over the latter part of the 20th century affected the phenology and population dynamics of single species. However, the key limitation to forecasting the effects of changing climate on ecosystems lies in understanding how it will affect interactions among species. We investigated the effects of climatic and biotic drivers on physical and biological lake processes, using a historical dataset of 40 years from Lake Washington, USA, and dynamic time‐series models to explain changes in the phenological patterns among physical and biological components of pelagic ecosystems. Long‐term climate warming and variability because of large‐scale climatic patterns like Pacific decadal oscillation (PDO) and El Niño–southern oscillation (ENSO) extended the duration of the stratification period by 25 days over the last 40 years. This change was due mainly to earlier spring stratification (16 days) and less to later stratification termination in fall (9 days). The phytoplankton spring bloom advanced roughly in parallel to stratification onset and in 2002 it occurred about 19 days earlier than it did in 1962, indicating the tight connection of spring phytoplankton growth to turbulent conditions. In contrast, the timing of the clear‐water phase showed high variability and was mainly driven by biotic factors. Among the zooplankton species, the timing of spring peaks in the rotifer Keratella advanced strongly, whereas Leptodiaptomus and Daphnia showed slight or no changes. These changes have generated a growing time lag between the spring phytoplankton peak and zooplankton peak, which can be especially critical for the cladoceran Daphnia. Water temperature, PDO, and food availability affected the timing of the spring peak in zooplankton. Overall, the impact of PDO on the phenological processes were stronger compared with ENSO. Our results highlight that climate affects physical and biological processes differently, which can interrupt energy flow among trophic levels, making ecosystem responses to climate change difficult to forecast.     

Earlier onset of the spring phytoplankton bloom in lakes of the temperate zone in a warmer climate

The decoupling of trophic interactions is potentially one of the most severe consequences of climate warming. In lakes and oceans the timing of phytoplankton blooms affects competition within the plankton community as well as food–web interactions with zooplankton and fish. Using Upper Lake Constance as an example, we present a model‐based analysis that predicts that in a future warmer climate, the onset of the spring phytoplankton bloom will occur earlier in the year than it does at present. This is a result of the earlier occurrence of the transition from strong to weak vertical mixing in spring, and of the associated earlier onset of stratification. According to our simulations a shift in the timing of phytoplankton growth resulting from a consistently warmer climate will exceed that resulting from a single unusually warm year. The numerical simulations are complemented by a statistical analysis of long‐term data from Upper Lake Constance which demonstrates that oligotrophication has a negligible effect on the timing of phytoplankton growth in spring and that an early onset of the spring phytoplankton bloom is associated with high air temperatures and low wind speeds.     

Resilience to changes in lake trophic state: Nutrient allocation into Daphnia resting eggs

During past decades, many lakes underwent drastic human‐caused changes in trophic state with strong implications for population dynamics and food web processes. We investigated the influence of trophic state on nutrient allocation into Daphnia resting eggs. The production of resting eggs is an important survival strategy, allowing Daphnia to cope with unfavorable environmental conditions. Allocation of essential nutrients into resting eggs may crucially influence embryonic development and offspring survival and thus is of great ecological and evolutionary interest. The capacity of Daphnia to adjust the allocation of nutrients into resting eggs may depend on the dietary nutrient supply, which may vary with trophic state‐related changes in the phytoplankton community composition. Resting eggs were isolated from sediment cores taken from Lake Constance, a large prealpine lake with a distinct eutrophication and reoligotrophication history, and analyzed for elemental (carbon, nitrogen, and phosphorus) and biochemical (sterols and fatty acids) nutrients. Carbon allocation into Daphnia resting eggs continuously decreased over time, irrespective of changes in trophic state. The allocation of nitrogen into Daphnia resting eggs followed the changes in trophic state, that is, nitrogen concentrations in resting eggs increased with eutrophication and decreased again with reoligotrophication. The allocation of phosphorus, sterols and long‐chain polyunsaturated fatty acids, such as eicosapentaenoic acid, into Daphnia resting eggs did not change significantly over time. Changes in trophic state strikingly influenced all trophic levels in Lake Constance. However, nutrient allocation into Daphnia resting eggs was mostly resilient to changes in lake trophic state.     

Timing of predation by rainbow trout controls Daphnia demography and the trophic status of a Minnesota 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.     

Zooplankton biomass dynamics in oligotrophic versus eutrophic conditions: a test of the PEG model

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Pelagic zonation of water quality and phytoplankton in the Great Lakes

Analysis of aquatic ecosystem data collected from large water bodies must consider spatial variations. A suite of pelagic survey stations exists for the Laurentian Great Lakes, but little is known about their redundancy. We present a strategy to delineate the lakes into zones based on water quality and phytoplankton biovolume. Water samples were collected from 72 sites in two seasons (spring and summer) from 2007 to 2010 in all five lakes. Integrated samples were analyzed for phytoplankton biovolume and nine water quality parameters. We conducted cluster analysis, principal components analysis and non-metric multidimensional scaling methods for water quality and phytoplankton taxon-specific biovolume for the Great Lakes basin and for each lake separately. There were significant lake-to-lake differences, and based on lake-specific analyses, Lake Superior, Lake Michigan and Lake Erie were each divided into three zones; Lake Huron and Lake Ontario were each grouped into two zones. The zones identified by water quality and phytoplankton provide an understanding of spatial distributions for evaluating monitoring data.     

Ecological Characteristics of Autotrophic Picoplankton in a Prealpine Lake

The seasonal distribution of autotrophic picoplankton in Lake Constance was investigated over four consecutive years. Cell numbers varied seasonally and vertically over four orders of magnitude (10² to 10⁶ cells ml⁻¹). A horizontal variation by a factor of 3 in abundance and biomass across the different parts of the lake was found during summer stratification. Picoplankton peaks occurred during the phytoplankton spring bloom and in late summer. Low values were characteristic for the clear-water phase in early summer and for autumn-winter. This seasonal pattern differed from that of larger phytoplankton in Lake Constance and from the seasonal distribution of picoplankton known from other lakes and marine environments. Picoplankton was predominated by chroococcoid cyanobacteria of about 0.6 μ³ biovolume. The average cell size increased from winter until early summer. Using HPLC pigment analysis, we identified zeaxanthin and β-carotene as typical picoplankton pigments. Results of the pigment analyses suggest that algae others than picocyano-bacteria may be more prominent in the picoplankton size class than derived from routine epifluorescence counting.