Daphnia lumholtzi and Daphnia ambigua: population comparisons of an exotica and a native cladoceran in Lake Okeechobee, Florida

The population dynamics of an exotic cladoceran (Daphnia lumholtzi Sars) and a native cladoceran (Daphnia ambigua) were studied over a 12 month period in subtropical Lake Okeechobee, Florida (USA), to quantify the extent of invasion of the exotic species and compare ecological niches. Daphnia lumholtzi accounted for up to 70% of the Daphnia assemblage during the summer months (June-August), while D.ambigua accounted for up to 97% of the Daphnia assemblage from fall to spring (October-April). The densities of the two species were inversely corelated. The exotic species was most concentrated in the shallower, warmer, north and south ends of the lake during the summer. It also was present, but at much lower densities, in the central lake region during the fall. The native species displayed a ubiquitous distribution throughout the lake during spring and winter, but was concentrated in the deeper, cooler, central region during the summer. Relationships of the two species with environmental conditions indicate that water column temperature might affect the seasonal and spatial distribution of the two Daphnia species. The results also indicate that D.lumholtzi may be filling a 'vacant' seasonal or spatial niche when conditions are unfavorable for D.ambigua.      

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.     

Habitat partitioning of native and exotic Daphnia in gradients of temperature and food: mesocosm experiments

1. Invasion of tropical zooplankton into temperate lakes provides an interesting opportunity to explore habitat segregation in a thermal gradient. 2. We explored differing vertical positioning of native and exotic Daphnia (Daphnia mendotae and Daphnia lumholtzi) in a large indoor mesocosm system (Plön plankton towers) during 2 month‐long experiments. The two towers were manipulated to provide gradients of both temperature (15–29 °C) and algal food (0.05–0.58 mg C L^?1) and a day–night cycle. 3. Both juvenile and adult D. lumholtzi showed a ‘typical’ vertical migration pattern, with higher densities in the epilimnion at night than during the day. They avoided the food‐poor middle layer. In contrast, D. mendotae adults showed little tendency to migrate into the epilimnion at night, remaining in the cooler hypolimnion while juveniles migrated. The vertical distribution of D. mendotae appeared unaffected by the presence of D. lumholtzi. 4. The strong migration behaviour of D. lumholtzi in the absence of fish cues suggests that this behaviour may be a constitutive trait. Habitat partitioning of the two species is probably the result of different thermal tolerances, with D. mendotae constrained to remaining in deeper water by high temperatures in the epilimnion and the tropical D. lumholtzi able to use the warm epilimnion at night.     

Crowding‐induced changes in growth,reproduction and morphology of Daphnia

• 1 Daphnia may reach high population densities seasonally, or in patches, in lakes. To test the effects of chemicals released by high daphniid densities on their life‐history traits, nine species of Daphnia, D. magna, D. pulicaria, D. pulex, D. hyalina, D. galeata, D. laevis, D. lumholtzi, D. ambigua and D. cucullata, were grown in water from crowded Daphnia cultures in a flow‐through system in the presence of abundant food.
• 2 Water from Daphnia at 85 L^‐1 depressed growth rate, and lowered body size and clutch at first reproduction of six species of small‐bodied Daphnia (adult body length < 1.8 mm), but had no significant effects on larger species. Two clones of D. pulex differed in their growth rate in response to crowding, indicating that response patterns may vary within species.
• 3 Chemicals released by crowded D. magna reduced tail spine length in D. lumholtzi and D. cucullata by 37% and 11%, respectively, and induced changes in carapace morphology in D. lumholtzi and D. ambigua.
• 4 Chemicals released by crowded conspecifics may provide an additional, density‐dependent mechanism of population regulation; when large species of Daphnia coexist at a high population density with small species, these chemicals may reinforce the competitive advantage of large species.

     

Predator-Mediated Coexistence of Exotic and Native Crustaceans in a Freshwater Lake?

The predatory effects of a Dipteran insect, Chaoborus, on the competition between exotic cladoceran Daphnia lumholtzi and two natives, D. catawba and D. pulex, were studied for a period of three years in a freshwater reservoir, Lake James, North Carolina (USA). D. lumholtzi was first encountered in September 1997 and it was present only between August and October when population densities of native species were low and that of Chaoborus sp. was high. The patterns observed in the population dynamics of the exotic D. lumholtzi and two natives, Chaoborus suggest that a predator mediated coexistence phenomenon might be taking place in Lake James. The strong positive correlation between Chaoborus and D. lumholtzi and the negative correlation between Chaoborus, D. catawba and D. pulex is supportive of this hypothesis.     

Hydrologic Connections and Overland Dispersal in An Exotic Freshwater Crustacean

Organisms living in lakes face the problem of dispersing through an uninhabitable matrix in order to reach suitable habitat. One possible mechanism for moving between lakes is by surface water connections. We used a seven-year data set to investigate the spread of the exotic cladoceran Daphnia lumholtzi among Missouri reservoirs with respect to stream connections. Reservoirs that were downstream of known populations of D. lumholtzi were more likely to become invaded than those that were not. However, invasion likelihood was only weakly related to the presence of upstream source populations, and reservoirs without potential upstream sources were colonized at a rate of 7.3% per year. The difference in invasion rate between lakes with and without upstream sources varied among years, and was significant in only two of the six years of study. In addition, the higher invasion rate of downstream lakes could be explained by their greater surface area. These patterns suggest that surface water connections may form one means of dispersal for D. lumholtzi, although overland movement is also important. The ability to utilize several modes of dispersal may explain the rapid expansion of this species' range since its arrival in North America.     

Linking biotic interactions and climate change to the success of exotic Daphnia lumholtzi

1. Warmer temperatures may increase cyanobacterial blooms in freshwater ecosystems, yet few ecological studies examine how increases in temperature and cyanobacterial blooms will alter the performance of non‐native species. We evaluated how competitive interactions and interactions between these two drivers of ecological change influence the performance of non‐native species using the native zooplankton Daphnia pulex and the non‐native zooplankton Daphnia lumholtzi as a model system. Based on the literature, we hypothesised that D. lumholtzi would perform best in warmer temperatures and in the presence of cyanobacteria. 2. Laboratory competition experiments showed that in the absence of competitors, growth rates of D. pulex (but not D. lumholtzi) were reduced at higher temperatures and with the cyanobacterial foods Anabaena flos‐aquae and Microcystis aeruginosa. In the presence of competitors, however, D. pulex emerged as the superior resource competitor at both temperatures with cyanobacterial food. We therefore predicted that, if competitive interactions are important to its establishment, D. lumholtzi would perform best in the absence of cyanobacteria in heated environments. 3. As predicted, when both species were introduced at low densities in field mesocosms, D. lumholtzi performed best at high temperatures without added cyanobacteria and worst at ambient temperatures with added cyanobacteria, indicating that competitive interactions are likely to be important for its establishment. 4. Taken together, these studies indicate that, while D. lumholtzi may benefit from increases in temperature, associated increased cyanobacterial blooms may hinder its performance. Thus, our findings underscore the importance of considering biotic interactions such as competition when predicting the future establishment of non‐native species in response to climate warming.     

Daphnid morphology deters fish predators

Spine and helmet production in zooplankton are thought to provide protection from invertebrate rather than vertebrate predators. We examined selectivity for Daphnia lumholtzi, a species that exhibits extreme cyclomorphosis with a large helmet and long tail spine (total length can exceed 5 mm), by juvenile bluegill (15–80 mm) in the laboratory and field. Bluegill consumed more D. pulex than D. lumholtzi when the species were presented alone. When the daphnids were offered together in equal numbers, bluegill selected against D. lumholtzi. Bluegill foraging behavior helped explain the observed nonrandom feeding. Bluegill capture efficiency foraging on D. pulex was high (85–100%) and handling times were low (usually too short to detect), whereas efficiencies were lower (40–96%) and handling times were longer (1–3 s) when foraging on D. lumholtzi, particularly for fish <50 mm. As they gained experience, bluegill <50 mm that oriented towards D. lumholtzi rejected them more often than striking. In addition, more D. lumholtzi were rejected and expelled than were D. pulex. From these experiments, we conclude that larger bluegill (>50 mm) are able to forage more successfully on D. lumholtzi than smaller fish. Selectivity by bluegill collected from a reservoir infested with D. lumholtzi verified our laboratory conclusions. Smaller bluegill selected against D. lumholtzi, whereas it was a preferred diet item for bluegill >50 mm. These results show that the morphology of D. lumholtzi interferes with predation by small planktivorous fish, posing foraging constraints for these fish more similar to those of piscivores, where handling time is important, than to those of planktivores, where prey density is of primary importance. Received: 13 August 1998 / Accepted 21 August 1998     

The temporal structure of the environment may influence range expansions during climate warming

Understanding the processes that influence range expansions during climate warming is paramount for predicting population extirpations and preparing for the arrival of non‐native species. While climate warming occurs over a background of variation due to cyclical processes and irregular events, the temporal structure of the thermal environment is largely ignored when forecasting the dynamics of non‐native species. Ecological theory predicts that high levels of temporal autocorrelation in the environment – relatedness between conditions occurring in close temporal proximity – will favor populations that would otherwise have an average negative growth rate by increasing the duration of favorable environmental periods. Here, we invoke such theory to explain the success of biological invasions and evaluate the hypothesis that sustained periods of high environmental temperature can act synergistically with increases in mean temperature to favor the establishment of non‐native species. We conduct a 60‐day field mesocosm experiment to measure the population dynamics of the non‐native cladoceran zooplankter Daphnia lumholtzi and a native congener Daphnia pulex in ambient temperature environments (control), warmed with recurrent periods of high environmental temperatures (uncorrelated‐warmed), or warmed with sustained periods of high environmental temperatures (autocorrelated‐warmed), such that both warmed treatments exhibited the same mean temperature but exhibited different temporal structures of their thermal environments. Maximum D. lumholtzi densities in the warmed‐autocorrelated treatment were threefold and eightfold higher relative to warmed‐uncorrelated and control treatments, respectively. Yet, D. lumholtzi performed poorly across all experimental treatments relative to D. pulex and were undetectable by the end of the experiment. Using mathematical models, we show that this increase in performance can occur alongside increasing temporal autocorrelation and should occur over a broad range of warming scenarios. These results provide both empirical and theoretical evidence that the temporal structure of the environment can influence the performance of species undergoing range expansions due to climate warming.     

Predator-induced phenotypic plasticity in the exotic cladoceran Daphnia lumholtzi

1. The exotic cladoceran Daphnia lumholtzi has recently invaded freshwater systems throughout the United States. Daphnia lumholtzi possesses extravagant head spines that are longer than those found on any other North American Daphnia. These spines are effective at reducing predation from many of the predators that are native to newly invaded habitats; however, they are plastic both in nature and in laboratory cultures. The purpose of this experiment was to better understand what environmental cues induce and maintain these effective predator‐deterrent spines. We conducted life‐table experiments on individual D. lumholtzi grown in water conditioned with an invertebrate insect predator, Chaoborus punctipennis, and water conditioned with a vertebrate fish predator, Lepomis macrochirus. 2. Daphnia lumholtzi exhibited morphological plasticity in response to kairomones released by both predators. However, direct exposure to predator kairomones during postembryonic development did not induce long spines in D. lumholtzi. In contrast, neonates produced from individuals exposed to Lepomis kairomones had significantly longer head and tail spines than neonates produced from control and Chaoborus individuals. These results suggest that there may be a maternal, or pre‐embryonic, effect of kairomone exposure on spine development in D. lumholtzi. 3. Independent of these morphological shifts, D. lumholtzi also exhibited plasticity in life history characteristics in response to predator kairomones. For example, D. lumholtzi exhibited delayed reproduction in response to Chaoborus kairomones, and significantly more individuals produced resting eggs, or ephippia, in the presence of Lepomis kairomones.     

Temperature elevation reduces the sensitivity of invasive cladoceran Daphnia lumholtzi to filamentous cyanobacterium Raphidiopsis raciborskii

1. Exotic cladoceran Daphnia lumholtzi is a highly invasive species in the north and south American continents and can potentially also invade European freshwaters and outcompete native Daphnia populations. However, European waterbodies are frequently dominated by less edible filamentous cyanobacteria including also invaders such as Raphidiopsis raciborskii, which might affect the fitness of D. lumholtzi. Furthermore, temperature may influence the sensitivity of D. lumholtzi to R. raciborskii filaments.
2. In this study, we determined whether the presence of R. raciborskii could obstruct the invasion of Europe by D. lumholtzi, through reducing its fitness, and whether this depends on temperature. We compared the population growth rate (r) and the somatic growth rate of D. lumholtzi maintained at two temperatures (20 or 26°C) and fed with two diets: green microalgae alone or green microalgae mixed with filaments of R. raciborskii. Three clones of D. lumholtzi were used to evaluate potential variation in response to the treatments among different clones.
3. At 20°C, the population growth rate of D. lumholtzi fed with cyanobacterial filaments declined sharply. This reduction was caused by increased egg abortion, egg degeneration, and mortality of newborn daphnids. At 26°C, R. raciborskii lost its harmful effect on the population growth of D. lumholtzi. The presence of cyanobacteria did reduce the somatic growth rate of D. lumholtzi at both temperatures and in all three clones except for one that had a similar somatic growth rate on both diets at 26°C.
4. The presence of filamentous cyanobacteria does affect growth in D. lumholtzi and may thereby substantially reduce its invasive potential, but only at lower temperatures. Therefore, the presence of filamentous cyanobacteria may not present an obstacle to the invasion of Europe by D. lumholtzi in a warmer future climate.

     

Food web structure provides biotic resistance against plankton invasion attempts

It is generally accepted that native communities provide resistance against invaders through biotic interactions. However, much remains uncertain about the types of ecological processes and community attributes that contribute to biotic resistance. We used experimental mesocosms to examine how zooplankton community structure, invertebrate predation, and nutrient supply jointly affected the establishment of the exotic Daphnia lumholtzi. We predicted that establishment would increase with declining biomass and diversity of native zooplankton communities and that an invertebrate predator (IP) would indirectly facilitate the establishment of D. lumholtzi due to its relatively long predator-deterring spines. Furthermore, we hypothesized that elevated nutrient supply would increase algal food availability and facilitate establishment. Only when the biomass and diversity of native zooplankton were significantly reduced, was D.␣lumholtzi able to successfully invade mesocosms. Although invertebrate predation and resource supply modified attributes of native zooplankton communities, they did not influence the establishment of D. lumholtzi. Overall, our␣results are consistent with observed population dynamics in invaded reservoirs where D.␣lumholtzi tends to be present only during the late summer, coinciding with historic mid-summer declines in native zooplankton populations. Lakes and reservoirs may be more susceptible to invasion not only by D. lumholtzi, but also by other planktonic species, in the late summer when native communities exhibit characteristics associated with lower levels of biotic resistance.     

Is competition important to arctic zooplankton community structure?

1. Daphnia pulex and Daphnia middendorffiana are commonly found in the Toolik Lake region of arctic Alaska. These two species are very similar morphologically, although their natural distributions differ markedly: D. pulex is restricted to shallow ponds, while D. middendorffiana is widely distributed and found in a variety of ponds and lakes. We compared the reproductive capabilities of D. pulex and D. middendorffiana grown under similar resource conditions and in the absence of the invertebrate predator Heterocope septentrionalis. In situ life table and mesocosm experiments were conducted in Toolik Lake and Dam Pond, habitats that have historically contained natural populations of D. middendorffiana, but never D. pulex. 2. Daphnia pulex exhibited a significantly higher net growth rate than D. middendorffiana in both life table and mesocosm experiments although D. pulex has never been found in either Toolik Lake or Dam Pond. Daphnia middendorffiana exhibited a negative net growth rate in Dam Pond, which had lower resource levels then Toolik Lake. Therefore, the smaller D. pulex appears to have a lower food threshold concentration than the larger D. middendorffiana. 3. Our results indicate that D. pulex is a superior resource competitor in the Toolik Lake region. These results combined with distributional patterns suggest that the restricted distribution of D. pulex in these arctic lakes and ponds cannot be explained by resource competition alone. We suggest that in the presence of H. septentrionalis, predation is an important factor structuring arctic zooplankton communities in the Toolik Lake region.     

Trait sorting in Daphnia colonising man‐made lakes

1. We used a zooplankton metacommunity to ask how dispersal, genetic drift and selection act to determine the local and regional distributions of trait variation. Since the formation of the lakes 80 years ago, cladoceran species have sorted into local assemblages that cluster by lake depth. Given this species sorting, we ask whether intraspecific variation in an ecologically important phenotypic trait – body size – has sorted as well. 2. We quantified changes in body size through time by measuring ephippia from D. pulicaria, D. dentifera and D. ambigua recovered from sediment cores from two lakes. We then estimated mean body size of contemporary populations of two competing species, Daphnia pulicaria and D. dentifera, in a laboratory common garden experiment. Finally, we used microsatellite loci to characterise genetic diversity and gene flow among local sites in the metacommunity. 3. Body size was variable both within and among years for the three species of Daphnia examined using sediment cores. For two lakes where we examined body size distributions through time, we observed a significant shift in body size of the first species to arrive after colonisation by other Daphnia species, which suggests selection has occurred historically. 4. Despite heritable variation in body size in the laboratory, evidence for trait sorting was only found for D. pulicaria, which was larger in deeper lakes. Mean body size varied among lakes, but did not sort relative to depth for D. dentifera. 5. Microsatellite data indicated that neutral genetic diversity was low in the region; only 27% of the individuals assayed were unique multi‐locus genotypes. We also found significant patterns of isolation by distance for both species. However, population structure was stronger in D. dentifera than in D. pulicaria. Hence, we conclude that a limited number of colonists have successfully invaded this metacommunity, and those genotypes arriving in this new region have experienced significant dispersal limitation among local sites. 6. Overall, while dispersal and selection have clearly led to the development of predictable community assemblages related to depth in this metacommunity, the distribution of phenotypic traits within species can differ substantially even between two trophically similar species. Our results highlight the complex roles of colonisation history, dispersal, selection and stochasticity in determining inter‐ and intra‐specific patterns in metacommunities.     

Zooplankton–phytoplankton relationships in shallow subtropical versus temperate lakes Apopka (Florida,USA) and Trasimeno (Umbria,Italy)

This study compares and contrasts the dynamics of phytoplankton, zooplankton, and nutrients in two of the largest shallow lakes in the USA (Lake Apopka, Florida) and Europe (Lago Trasimeno, Umbria, Italy) and considers particularly the biomass ratio of zooplankton to phytoplankton (BZ:BP) in relation to nutrient levels and in the context of data from other subtropical and temperate lakes. Lake Apopka is hypereutrophic with higher concentrations of total phosphorus (TP), nitrogen (TN), and nearly an order of magnitude higher BP than Lago Trasimeno. However, combined data from the two lakes can be fit to a single log–log regression model that explains 72% of the variability in BP based on TP. In contrast, BZ has a significant positive log–log relationship with TP only for Lago Trasimeno, and is much lower than expected based on the TP concentrations observed in Lake Apopka. Lake Apopka has a fish assemblage that includes high densities of gizzard shad (Dorosoma cepedianum) and threadfin shad (D. petenense), similar to other eutrophic Florida lakes that also have extreme low BZ. The ratio BZ:BP is below 0.01 in Lake Apopka, 10-fold lower than in Trasimeno and among the lowest values reported in the literature. Although stress of high water temperature and a greater proportion of inedible cyanobacteria may be contributing factors, the collective results support an emerging view that fish predation limits the biomass of crustacean zooplankton in subtropical lakes. Handling editor: S. I. Dodson     

Biological Invasions Across Spatial Scales: Intercontinental, Regional, and Local Dispersal of Cladoceran Zooplankton

The frequency of dispersal of invertebrates among lakes depends upon perspective and spatial scale. Effective passive dispersal requires both the transport of propagules and the establishment of populations large enough to be detected. At a global scale, biogeographic patterns of cladoceran zooplankton species suggest that effective dispersal among continents was originally rare, but greatly increased in the past century with expanded commerce. Genetic analysis allows some reconstruction of past dispersal events. Allozyme and mitochondrial DNA comparisons among New World and Old-World populations of several exotic cladocerans have provided estimates for likely source populations of colonists, their dispersal corridors, and timing of earlier dispersal events. Detecting the Old-World tropical exotic Daphnia lumholtzi early in its invasion of North America has allowed detailed analysis of its spatial spread. Twelve years of collection records indicate a rapid invasion of reservoirs in the United States, by both regional spread and long-distance jumps to new regions. Combining landscape features with zooplankton surveys from south-central US reservoirs revealed higher colonization rates of D. lumholtzi at lower landscape positions, a result which can be explained by either greater propagule load or by higher susceptibility of these downstream reservoirs. Because invaded reservoirs provide a source of propagules for nearby floodplain ponds, the rarity of this species in ponds suggests limitation by local environments. Such analyses of invading species over multiple spatial scales allow a better understanding of ecological processes governing invasion dynamics.     

Seasonality and Fecundity of Daphnia Lumholtzi Sars in Lake Phewa,Nepal

Monthly sampling of zooplankton in Lake Phewa, Nepal revealed 18 species of zooplankton excluding protozoans. The seasonality and fecundity of the largest common cladoceran Daphnia lumholtzi Sars was studied. Two distinct peaks of abundance were noted. No active stages were found during May and June. Egg ratio varied from zero to 0.88 in mature females. From the presence of only helmeted individuals of this species it is concluded that the predation pressure by fish was high.     

Overcrowding,food and phosphorus limitation effects on ephipphia production and population dynamics in the invasive species <Emphasis Type="Italic">Daphnia lumholtzi</Emphasis>

Daphnia lumholtzi has been very successful in colonizing North America since its appearance in Texas in 1990. Although previous studies have sought to link its success as an invasive species with various aspects of its population biology, there is little experimental data linking the invasion success of D. lumholtzi with its autecology, specifically its reproduction strategy. In this study we sought to link food quality and quantity to diapause in D. lumholtzi through a variation in phosphorus (P) content of algae, food quantity, and light level. We also assessed the effect of Daphnia peak population densities on reproductive rates and production of resting eggs. We found that when food is abundant, per capita ephippia production may be limited by P, but under food limitation conditions, there is no significant effect of food quality on ephippia production. Our results suggest that a combination of food quality/quantity and population density may work together to induce the production of resting eggs in this invasive species. Handling editor: S. Declerk     

Ecological consequences of intraspecific variation in lake Daphnia

1. Although populations harbour considerable diversity, most ecological studies still assume they are homogeneous. However, mounting evidence suggests that intraspecific diversity is not only common, but also important for interactions with community members. Here, intraspecific variation in Daphnia dentifera in haemoglobin content is shown to be a marker of hypolimnion use. 2. Hypolimnion use differed substantially within and among D. dentifera populations. Daphnia dentifera with haemoglobin resided primarily in the hypolimnion, while D. dentifera lacking haemoglobin migrated vertically. These ‘deep’ and ‘migratory’D. dentifera had different seasonal phenologies and dynamics. 3. Deep and migratory D. dentifera had qualitatively different relationships with an important competitor, Daphnia pulicaria. Deep D. dentifera density was negatively correlated with D. pulicaria density, whereas migratory density was not correlated with D. pulicaria density. 4. Given that D. pulicaria tends to reside in the hypolimnion, this negative correlation probably reflects competition between D. pulicaria and the deep D. dentifera. This pattern would have been missed if only the relationship between the overall lake populations of D. dentifera and D. pulicaria had been studied. 5. Abundances of deep D. dentifera and D. pulicaria were both correlated with the size of the hypolimnetic refuge from fish predation, but in opposite directions. Lakes with large refuges generally had high D. pulicaria and low deep D. dentifera densities.