Factors which affect the abundance of an invasive cladoceran, Daphnia lumholtzi, in U.S. reservoirs

1.  Daphnia lumholtzi is a subtropical, Old World species which is rapidly spreading throughout reservoirs in the southern U.S.A. It was first recorded in Lake Texoma (Oklahoma–Texas) in September 1991.
2. Southern U.S. reservoirs typically have strong spatial and temporal gradients in temperature, conductivity, turbidity and in the distribution of organisms. Therefore, the present experiments examined the reproductive and moulting rates, and survival of D . lumholtzi in relation to extremes of food concentration, temperature, conductivity and turbidity.
3. Increases in temperature (range = 15–29 °C) and decreases in turbidity (range = 0.6–470 NTU) increased reproductive and moulting rates, whereas food concentration only affected the rate of reproduction, while conductivity had no effects. Survival was affected only by temperature.
4.  Daphnia lumholtzi tolerates high temperatures (27–30 °C) at which other cladocerans in Lake Texoma disappear. Therefore, D . lumholtzi may exploit resources in midsummer, when there are few potential competitors.     

Competition Between Native and Exotic Daphnia: In situ Experiments

The recently introduced exotic cladoceran Daphnia lumholtzioffers an excellent opportunity to study the interactions betweenexotic and native species in invaded communities. Lake surveysin Missouri have indicated a seasonal succession between nativeDaphnia and D. lumholtzi. In the current study, we examinedcompetition between D. lumholtzi and the native Daphnia parvulaby conducting seasonal in situ field experiments in 1.6 l enclosures.Competition was assessed by comparing the rates of increase(r) and birth rates (b) of each species when grown alone versuswhen grown together in these enclosures. At high densities,D. lumholtzi suppressed D. parvula rates of increase duringthe late summer and fall experiments, but did not appear tosuppress D. parvula birth rates. The rates of increase of D.lumholtzi did not appear to be affected by the presence of D.parvula. The results of these experiments indicate that althoughcompetition between the two species occurs seasonally at highdensities, the effects are asymmetrical. The lack of competitiveeffects on D. lumholtzi byD. parvula suggests that factors otherthan competition are involved in explaining the absence ofD.lumholtzi in spring zooplankton assemblages.     

Differing effects of suspended sediments on the performance of native and exotic Daphnia

1. Although large cladocerans are usually uncommon in large rivers, Daphnia lumholtzi (an exotic species in North America) is widespread and occasionally abundant. We collected zooplankton on the Illinois River (Illinois, U.S.A.) in 1995 and 1996 and found that the peak density of D. lumholtzi (25 L⁻¹) typically exceeded that of all native species combined. Maximum density occurred during warm periods (up to 27 °C) when concentrations of inorganic suspended sediments were high (>50 mg L⁻¹).
2. Using a life table experiment, we examined the effects of variation in suspended sediment (0 and 80 mg L⁻¹) and food (10⁴ and 10⁵ Ankistrodesmus cells mL⁻¹) on fitness of D. lumholtzi and the native Daphnia parvula. Daphnia lumholtzi had greater survivorship than D. parvula in most treatments and higher life-time fertility than D. parvula in all treatments. Both species achieved their fastest intrinsic rates of growth in treatments with high food, but their responses to suspended solids differed. The growth rate of D. lumholtzi in high food was slightly increased by higher turbidity, whereas that of D. parvula was depressed.
3. Results suggest that the ability of D. lumholtzi to tolerate suspended solids is an important factor contributing to its success in invading North American rivers.     

Invasibility of a reservoir to exotic Daphnia lumholtzi: experimental assessment of diet selection and life history responses to cyanobacteria

SUMMARY 1. In reservoirs of the south-central United States, the exotic cladoceran Daphnia lumholtzi is common during warm midsummer conditions, when cyanobacteria are abundant and native Daphnia are rare. In the current study, we employed surveys, field experiments, and a life table experiment to investigate the role of food quality in explaining the distribution and phenology of D. lumholtzi , relative to two native species ( Daphnia parvula and Daphnia mendotae ).
2. During May–September 2000 in eutrophic McDaniel Lake, Missouri U.S.A., cyanobacteria (primarily Oscillatoria ) first appeared at 6-m depth and then became abundant throughout the epilimnion.
3. During the May field experiment, D. lumholtzi , D. parvula and D. mendotae all consumed a similar diet of algae, showing positive selection for small greens (chlorophytes and cryptophytes <20 μm). During the July experiment, when the epilimnion exceeded 25 °C and cyanobacteria were common in the lake, D. lumholtzi consumed significantly more total algae and more cyanobacteria than the two native species. Although the Daphnia selected against cyanobacteria, all three species consumed about 25% of this food in their diet.
4. A life table experiment compared the responses of D. lumholtzi and D. parvula with variation in density of high-quality food ( Ankistrodesmus ) and concentration of a toxic strain of cyanobacteria ( Anabaena flos-aquae ). Both Daphnia species showed reduced survivorship, fertility and intrinsic rates of increase in response to elevated concentrations of cyanobacteria, particularly at the higher food level.
5. The results suggest that D. lumholtzi shows similar inhibition from cyanobacteria as does the native Daphnia . However, their continued high in situ feeding rates imply that D. lumholtzi is less affected by midsummer conditions in warm-water reservoirs than are native Daphnia .     

Trophic ecology of striped bass, Morone saxatilis, in a freshwater reservoir (Lake Texoma, U.S.A.)

Many vertebrate predators consume a wide variety of prey types, depending upon availability and vulnerability. In contrast, striped bass, Morone saxatilis , that have been introduced to Lake Texoma (Oklahoma-Texas, U.S.A.) use a very limited array of fish (mostly clupeids of the genus Dorosoma ) as prey. As a large, mobile predator, M. saxatilis should be capable of capturing and consuming numerous other species of fish that are available in the reservoir. However, examination of 1845 stomachs year-around over 5 years showed that the only marked 'switching' among prey was from Dorosoma to a diet including a high percentage of insects during spring–early summer, ignoring most other fish taxa that could have served as food. Even under essentially starvation conditions in late summer of years with scarce Dorosoma, M. saxatilis in Lake Texoma did not switch to other available fish as prey. Patterns of predation by M. saxatilis are trenchantly different from place to place: very narrow prey selectivity even under starvation conditions has been reported once previously for the species in a freshwater reservoir, but in its native marine and estuarine environment and in some other reservoirs the species is more catholic in its use of prey. Why this large predator shows fidelity in some environments to particular prey, even to the extent of starvation, remains an enigma.     

Invasibility of plankton food webs along a trophic state gradient

Biological invasions are becoming more common, yet the majority of introduced exotic species fail to establish viable populations in new environments. Current ecological research suggests that invasion success may be determined by properties of the native ecosystem, such as the supply rate of limiting nutrients (i.e. trophic state). We examined how trophic state influences invasion success by introducing an exotic zooplankter, Daphnia lumholtzi into native plankton communities in a series of experimental mesocosms exposed to a strong nutrient gradient. We predicted that the attributes of nutrient-enriched communities would increase the likelihood of a successful invasion attempt by D. lumholtzi . Contrary to our original predictions, we found that D. lumholtzi was often absent from mesocosms that developed under high nutrient supply rates. Instead, the presence of D. lumholtzi was associated with systems that had low nutrients, low zooplankton biomass, and high zooplankton species diversity. Using generalized estimating equations (GEE) and multivariate species data, we found that the presence–absence of D. lumholtzi could be explained by variations in zooplankton community structure, which was itself strongly influenced by nutrient supply rate. We argue that the apparent invasion success of D. lumholtzi was inhibited by the dominance of another cladoceran species, Chydorus sphaericus . These results suggest that the interaction between trophic state and species identity influenced the invasion success of introduced D. lumholtzi .     

Influence of Temperature on Exotic Daphnia lumholtzi and Implications for Invasion Success

We investigated the effects of water temperature on the exoticcladoceran Daphnia lumholtzi in a eutrophic Kansas reservoir(USA) and under laboratory conditions. Daphnia lumholtzi demonstrateda distinct late summer appearance in the reservoir at temperaturesbetween 26 and 31°C, which corresponded with steep declinesin the densities of native Daphnia spp. Laboratory life-tableexperiments confirmed that D. lumholtzi performs well at elevatedwater temperatures. The intrinsic rate of increase (r), netreproductive rate (R_o), age at first reproduction, survivorship(l_x) and molting rates all demonstrate that D. lumholtzi hasa high temperature optimum between 20 and 30°C. A comparisonof literature-reported r-values indicates that the reproductiverate of D. lumholtzi is comparable with other Daphnia spp. between20 and 25°C, but also implies that D. lumholtzi may out-performsome Daphnia spp. at temperatures >25°C. Collectively,these results suggest that D. lumholtzi may be taking advantageof a late summer thermal niche, and that this invader may continueto colonize lakes and reservoirs in the southern US. However,life table data also indicate that D. lumholtzi performs poorlyat temperatures < 10°C, which may inhibit the range expansionof this invader into northern waters.     

Implications of an invertebrate predator's ( Bythotrephes cederstroemi) atypical effects on a pelagic zooplankton community

The traditional view of predaceous zooplankton is that they prefer small-bodied prey, are hindered by morphological anti-predator defenses, and have a minor influence on zooplankton communities when fish are present. We performed a series of experiments with the large-bodied onychopod (cladoceran) Bythotrephes cederstroemi, in which we incubated this predator with known prey to determine prey preference and predation rates. We also performed an allozyme analysis of prey tissue in the gut of B. cederstroemi collected from several stations around Lake Michigan to determine what prey types are chosen in the field. We found that B. cederstroemi does not fit the standard invertebrate predator mold: adult B. cederstroemi prefer large (>2.0 mm) Daphnia pulicaria over smaller individuals; the elongated tailspine and helmet of Daphnia galeata mendotae are not effective deterrents to B. cederstroemi predation; and B. cederstroemi is a generalist predator with the potential to consume a significant portion of cladoceran production in Lake Michigan. This revised version was published online in July 2006 with corrections to the Cover Date.     

The distribution and variation of Daphnia lumholtzi (Crustacea: Cladocera) in relation to fish predation in Lake Albert, East Africa

In Lake Albert Daphnia lumholtzi is found in two forms. One has a pointed anterior prolongation, or helmet, on the head. The other has a shorter rounded head, and was originally described as a separate species, D. monacha . The latter form dominates the zooplankton in the middle of the lake where planktivorous fish are rare or absent. The helmeted form becomes commoner near the margins of the lake and reaches its greatest abundance in Ndaiga Lagoon, where planktivorous fish are common. The possession of a helmet is associated with a reduction in the size of the carapace compared to the round headed form. The carapace with its contained eggs is the most conspicuous part of a cladoceran, so that the helmeted forms are at an advantage in the presence of planktivorous fish which locate their prey by sight. The mid-lake monacha forms are larger than specimens of the same form in Ndaiga Lagoon, where it is shown that Alestes baremose feeds selectively on the larger specimens of the monacha form.
The helmeted form produces more, but smaller eggs than the monacha form. The total brood volume (= mean egg volume x mean number eggs per female) is greatest in the midlake monacha forms. The selective advantages of variations in egg size and the possession of a helmet are discussed. It is concluded that the data from Lake Albert support the hypothesis of Brooks (1965) concerning the adaptive significance of helmet development in Daphnia .     

The distribution and variation of Daphnia lumholtzi (Crustacea: Cladocera) in relation to fish predation in Lake Albert, East Africa

In Lake Albert Daphnia lumholtzi is found in two forms. One has a pointed anterior prolongation, or helmet, on the head. The other has a shorter rounded head, and was originally described as a separate species, D. monacha . The latter form dominates the zooplankton in the middle of the lake where planktivorous fish are rare or absent. The helmeted form becomes commoner near the margins of the lake and reaches its greatest abundance in Ndaiga Lagoon, where planktivorousfish are common. The possession of a helmet is associated with a reduction in the size of the carapace compared to the round headed form. The carapace with its contained eggs is the most conspicuous part of a cladoceran, so that the helmeted forms are at an advantage in the presence of planktivorous fish which locate their prey by sight. The mid-lake monacha forms are larger than specimens of the same form in Ndaiga Lagoon, where it is shown that Alestes baremose feeds selectively on the larger specimens of the monacha form.
The helmeted form produces more, but smaller eggs than the monacha form. The total brood volume (= mean egg volume x mean number eggs per female) is greatest in the midlake monacha forms. The selective advantages of variations in egg size and the possession of a helmet are discussed. It is concluded that the data from Lake Albert support the hypothesis of Brooks (1965) concerning the adaptive significance of helmet development in Daphnia .     

Feeding dynamics of the copepod Diacyclops thomasi before, during and following filamentous cyanobacteria blooms in a large, shallow temperate lake

Cyanobacteria blooms are an increasing problem in temperate freshwater lakes, leading to reduced water quality and in some cases harmful effects from toxic cyanobacteria species. To better understand the role of zooplankton in modulating cyanobacteria blooms, from 2008 to 2010 we measured water quality and plankton abundance, and measured feeding rates and prey selectivity of the copepod Diacyclops thomasi before, during and following summertime cyanobacteria blooms in a shallow, eutrophic lake (Vancouver Lake, Washington, USA). We used a combined field and experimental approach to specifically test the hypothesis that copepod grazing was a significant factor in establishing the timing of cyanobacteria bloom initiation and eventual decline in Vancouver Lake. There was a consistent annual succession of zooplankton taxa, with cyclopoid copepods (D. thomasi) dominant in spring, followed by small cladocerans (Eubosmina sp.). Before each cyanobacteria bloom, large cladocerans (Daphnia retrocurva, Daphnia laevis) peaked in abundance but quickly disappeared, followed by brief increases in rotifers. During the cyanobacteria blooms, D. thomasi was again dominant, with small cladocerans abundant in autumn. Before the cyanobacteria blooms, D. thomasi substantially consumed ciliates and dinoflagellates (up to 100% of prey biomass per day), which likely allowed diatoms to flourish. A shift in copepod grazing toward diatoms before the blooms may have then helped to facilitate the rapid increase in cyanobacteria. Copepod grazing impact was the highest during the cyanobacteria blooms both years, but focused on non-cyanobacteria prey; copepod grazing was minimal as the cyanobacteria blooms waned. We conclude that cyclopoid copepods may have an indirect role (via trophic cascades) in modulating cyanobacteria bloom initiation, but do not directly contribute to cyanobacteria bloom decline.     

Trophic relationships and seasonal utilization of salt-marsh creeks by zooplanktivorous fishes

Synopsis In a high salinity estuary at North Inlet, South Carolina, co-occurrence and possible competition among adults of four dominant zooplanktivorous fishes were minimized by seasonal adjustments in lateral and vertical distributions as well as in dietary preferences. In winter, Atlantic silversides, Menidia menidia, occupied the entire water column while other planktivores were rare or absent from the estuary, and they consumed large prey such as mysid shrimps and fish larvae. An immigration of bay anchovies, Anchoa mitchilli, in the spring resulted in a redistribution of species with Atlantic silversides shifting to the surface waters and bay anchovies dominating the lower half of the water column. Both fishes consumed mostly copepods in the spring, but each favored a different species. There was little similarity in the large prey items consumed by the two fishes. Striped anchovies, Anchoa hepsetus, arrived in mid-summer and were most abundant at the surface while bay anchovies continued to dominate the bottom waters. Atlantic silversides were rare in all summer collections. The diets of the two anchovies were similar, but vertical separation during the period of maximum zooplankton abundance probably minimized competition. Rough silversides, Membras martinica, which were obligate surface dwellers, shared the upper water column with striped anchovies, but the two species had very different diets during their period of co-occurrence. Although seasonal changes in fish diets reflected shifts in zooplankton composition and all fishes consumed a variety of prey types, preferences for some prey taxa and total avoidance of others were indicated. Electivity indices indicated an especially strong selection for fiddler crab megalopae by all fishes in the summer and fall. All fishes, except rough silversides, which fed almost exclusively on copepods and crab zoeae, consumed large prey items when they were available. Fine scale partitioning of the food resources was apparent in the selection of different copepod and insect species by the fishes. Spatial and temporal separation in the distribution and/or dietary preferences of the zooplanktivores fishes probably reduces the potential for resource competition. Given the high abundances and selectivity of the planktivores, significant impacts on some zooplankton populations probably result.     

Seasonal succession of zooplankton in the north basin of Lake Biwa

To examine the seasonal succession of the entire zooplankton community in Lake Biwa, zooplankton biomass (on an areal basis) and its distribution patterns among crustaceans, rotifers and ciliates were studied in the north basin from April 1997 to June 1998. Seasonal changes in phytoplankton and population dynamics of Daphnia galeata were also examined to assess food condition and predation pressure by fish. From March to November, crustaceans dominated zooplankton biomass, but rotifers and ciliates were dominant from December to February. Among crustaceans, Eodiaptomus japonicus was the most abundant species, followed by D. galeata. Zooplankton biomass increased from January to a peak in early April, just before the spring bloom of phytoplankton, then decreased in mid-April when mortality rate of D. galeata increased. From mid-June, zooplankton increased and maintained a high level until the beginning of November. During this period, both birth and mortality rates of D. galeata were relatively high and a number of rotifer and crustacean species were observed. However, their abundances were very limited except for E. japonicus which likely preys on ciliates and rotifers. In Lake Biwa, food sources other than phytoplankton, such as resuspended organic matter from the sediments, seems to play a crucial role in zooplankton succession from winter to early spring, while zooplankton community seems to be regulated mainly by fish predation from summer to fall.     

Utilization of the exotic cladoceran Daphnia lumholtzi by juvenile fishes in an Illinois River floodplain lake

Daphnia lumholtzi comprises a substantial component of the zooplankton community during mid‐ to late‐summer in Lake Chautauqua, a floodplain lake along the Illinois River near Havana, Illinois. In order to quantify the utilization of D. lumholtzi by juvenile fishes, diet analyses were conducted for seven juvenile fish species collected from Lake Chautauqua during the 2001 annual drawdown period. Freshwater drum Aplodinotus grunniens and emerald shiner Notropis atherinoides demonstrated negative selectivity for D. lumholtzi relative to native zooplankton species whereas four species of fish (bluegill Lepomis macrochirus, white bass Morone chrysops, white crappie Pomoxis annularis and black crappie Pomoxis nigromaculatus) consumed substantial amounts of D. lumholtzi. Although selectivity values for D. lumholtzi varied among these fish species, positive selection for D. lumholtzi increased similarly among larger size classes of each fish species, and corresponded with ontogenetic shifts in diet. Mean body length of D. lumholtzi consumed by 20–69 mm L_T juvenile fishes ranged from 0·75 to 0·99 mm with a calculated total length range of 2·0–2·6 mm. Results from this study provide evidence that high abundances of D. lumholtzi in mid‐ to late‐summer provide an additional food source for several juvenile fish species during a time when abundances of large native cladoceran species (i.e. Daphnia) are low, and juvenile fishes are searching for larger prey associated with ontogenetic shifts from zooplankton to macroinvertebrates and fishes. Because zooplankton production is typically lower in rivers than in lakes, survivorship of juvenile fishes produced in floodplain lakes may be higher in riverine systems if they are not reliant on zooplankton as a primary food resource. Therefore, high abundances of D. lumholtzi may benefit juvenile fishes in managed floodplain lakes, such as Lake Chautauqua, by increasing growth and facilitating the transition from zooplanktivory to insectivory or piscivory.     

Diet composition and prey preference of tench, Tinca tinca (L.), common bream, Abramis brama (L.), perch, Perca fluviatilis L. and roach, Rutilus rutilus (L.), in two contrasting gravel pit lakes: potential trophic overlap with wildfowl

The abundance of benthic macroinvertebrate taxa and the relative abundance of zooplankton taxa were compared with the diet of bream, tench. perch and roach from two gravel pit lakes during May-July 1986 and 1987. Significant food preferences were demonstrated between species and between lakes. Chironomid pupae dominated the macroinvertebrate diet of perch, roach, ate predominantly either Spirogyra sp. (St Peters Lake) or Daphniu hyalina (Main Lake).
The Main Lake, bream fed largely upon chironomid larvae and the bivalve Sphuerium but in St Peters Lake they positively selected a variety of less abundant benthic invertebrate taxa. Bream switched from benthos to zooplankton in the Main Lake in 1986. Tench ate large numbers of Aselhs and showed positive selection of various macroinvertebrate prey in St Peters but ate D. hjulim in Main Lake. Tropic overlap for chironomids and other macroinvertebrate prey was demonstrated between perch, bream and tench, and potentially with wildfowl which used the gravel pits for breeding and wintering.     

Two new species of Keratella (Rotifera: Monogononta: Brachionidae) from Inner Mongolia,P.R. China

The relationship of species abundance to eight environmental variables was tested for 24 common species of crustacean zooplankton collected in the Atchafalaya River Basin during the summer of 1994. Stepwise regressions (α = 0.05) revealed significant relationships between zooplankton abundance and at least one environmental variable for 18 species ( R2 = 0.14-0.61, p < 0.0435-0.0001). The majority of these species' peak abundances were correlated with variables indicative of seasonal changes in floodplain habitat, as the Atchafalaya river receded, water temperature increased, and/or phytoplanktonic photosynthesis increased. Surface water temperature and the percent saturation of dissolved oxygen showed the most significant relationships, but specific conductance, current velocity, and Secchi disk depths were also related to abundance patterns of certain taxa. A principal components analysis of species abundances provided further insight into the partitioning of temporally-distinct zooplanton assemblages, showing that several species ( Bosmina longirostris, Daphnia parvula, Eurytemora affinis, and Ceriodaphnia quadrangula) predominated during early summer, and were supplanted by a distinct late-summer assemblage ( Diaphanosoma birgei, Moina micrura, Mesocyclops edax, and Daphnia lumholtzi) as time progressed. The transitional assemblage was dominated by Simocephalus serrulatus, Macrocyclops albidus, Microcyclops rubellus, and Thermocyclops inversus, all of which were most abundant in the hypoxic conditions characteristic of the latter stages of the Atchafalaya River flood-pulse. This revised version was published online in July 2006 with corrections to the Cover Date.     

Interactions between Nile tilapia (Oreochromis niloticus) and cladocerans in ponds (Khartoum,Sudan)

Juvenile Nile tilapia (Oreochromis niloticus) are omnivorous, and the question asked in this study is how they affect on their environment? Do they mainly act as predators on the cladoceran zooplankton or do they compete with the cladocerans for phytoplankton? This problem was studied in three ponds with and three ponds without small tilapia (3–5 cm). The fish growth rate, the succession of plankton species and the changes in abiotic conditions, were monitored over a period of 67 days. The fish biomass was kept low and the mean was approximately constant (12.6 g m^?2) during the experiment. Phosphate was added to avoid phytoplankton nutrient limitation. Although the diet of Nile tilapia contained both phytoplankton and zooplankton, the fish affected the ecosystem in a similar way as zooplanktivorous fish. The fish ponds got more phytoplankton due to increase of Chlorophyta. Effects on the other phytoplankton groups Euglenophyta, Bacillariophyta, Cryptophyta and Cyanophyta could not be registered. The ponds without fish had higher densities of Daphnia lumholtzi and D. barbata. The other Cladocerans seemed less influenced by fish presence. The relative fish growth rate was most positively correlated with the density of Daphnia lumholtzi, Diaphanosmoa excisum and Bosmina longirostris. Tilapia seemes to have two feeding modes: (1) preying on large zooplankton and (2) unselective filtration of small planktonic organisms such as phytoplankton. In our experiment the first feeding mode affected the ecosystem more than the second.     

Range expansion and potential dispersal mechanisms of the exotic cladoceran Daphnia lumholtzi

. Recently the exotic cladoceran Daphnia lumholtzi has invadedfreshwater systems throughout the southern and midwestern UnitedStates. We conducted regional surveys of eastern Kansas reservoirsto document the range expansion of D.lumholtzi. Daphnia lumholtziwas found in five of 35 reservoirs sampled in 1994, and 11 ofthe 35 reservoirs when re-sampled in 1997. In addition, we sampled40 small ponds inaccessible to recreational boats, within thewatershed of an invaded reservoir. We did not find D.lumholtziin any of these ponds, suggesting that non-human dispersal mechanismsplay an insignificant role in the range expansion of D.lumholtzithroughout the United States. Further experimentation, however,is needed to determine if the absence of D.lumholtzi from thesesmall ponds is due to insufficient dispersal mechanisms or theinability of this invader to successfully colonize followingarrival. Daphnia lumholtzi has broad limnological tolerances.However, invaded reservoirs tend to be larger in area, havehigher Secchi disk depths, and lower total phosphorus, totalnitrogen and chlorophyll a levels relative to non-invaded reservoirs.Analyses of pre-invasion zooplankton communities indicate thatD.lumholtzi may be invading reservoirs in which native Daphniaspecies are rare. While the long-term effects of the invasionof D.lumholtzi are unknown, it has the potential to dominatelate summer zooplankton communities in eastern Kansas reservoirs.Therefore, we need to continue to survey and monitor invadedreservoirs to document the range expansion of D.lumholtzi anddetermine the long-term implications of the introduction ofthis invader.     

How well do laboratory experiments explain field patterns of zooplankton emergence?

1. We conducted a laboratory experiment to explore potential mechanisms driving variation in zooplankton emergence from diapausing eggs observed in Oneida Lake, NY, U.S.A. We hypothesized that variation in timing of ice-out (date of thawing of ice) between 1994 and 1995, which resulted in variation in photoperiod–temperature cues, contributed to the differences in the observed field patterns. Environmental chambers were used to establish weekly photoperiod–temperature combinations that reproduced natural conditions in Oneida Lake in 1994 and 1995. In addition, a third treatment ('dark') exposed eggs only to the changes in temperature. We recorded zooplankton emergence for 2.5 simulated ice-free seasons.
2. Nine cladoceran taxa were found to hatch, but only Daphnia pulicaria in large numbers. Hatching of D. pulicaria was recorded throughout the season in the two light treatments and sporadically in the dark treatment. The early ice-out treatment had the highest emergence, followed by the late ice-out and dark treatments. Among taxa, there was temporal segregation with five hatching in the early weeks of sampling and two taxa hatching during the middle weeks. Alona hatched late in the first year, but earlier in the second year.
3. We compared our laboratory results of D. pulicaria hatching with the field data obtained by Cáceres (1998) . Hatching was continuous in the laboratory, whereas a synchronous spring emergence was found in the field. However, in both the laboratory and the field, more D. pulicaria hatched under conditions reflecting ice-out occurring in March as opposed to April. Because differences in rates and timing of emergence can affect the population and community dynamics of pelagic systems, we suggest caution when applying laboratory results to field populations.     

Responses of zooplankton and zoobenthos to experimental acidification in a high-elevation lake (Sierra Nevada, California, U.S.A.)

SUMMARY. 1. During the summer of 1987 we conducted an acidification experiment using large enclosure at Emerald Lake, a dilute, high-elevation lake in the Sierra Nevada, California, U.S.A. The experiment was designed to examine the effects of acidification on the zooplankton and zoobenthos assemblages of Sierran lakes.
2. Treatments consisted of a control (pH 6.3) and pH levels of 5.8, 5.4, 5.3, 5.0 and 4.7; each treatment was run in triplicate. The experiment lasted 35 days.
3. The zooplankton assemblage was sensitive to acidification. Daphnia rosea Sars emend. Richard and Diaptomns signicauda Lilljeborg decreased in abundance below pH 5.5–5.8, and virtually disappeared below pH 5.0. Bosmina longirostris (Müller) and Keratella taurocephala Ahlstrom became more abundant with decreasing pH. although B. longirostris was rare in the pH 4.7 treatment. These species might serve as reliable indicators of early acidification in lakes such as Emerald Lake.
4. The elimination of D. rosea in acidified treatments probably allowed the more acid-tolerant taxa to increase in abundance because interspecific competition was reduced. Even slight acidification can therefore alter the structure of the zooplankton assemblage.
5. In contrast to the zooplankton, there was no evidence that the zoobenthos in the enclosures was affected by acidification.