Larval fish-induced phenotypic plasticity of coexisting Daphnia: an enclosure experiment

1. The indirect effects of predators on lower trophic levels have been studied without much attention to phenotypically plastic traits of key food web components. Phenotypic plasticity among species creates phenotypic diversity over a changing environmental landscape. 2. We measured the indirect effects of planktivorous larval walleye (Stizostedion vitreum) on phytoplankton biomass through their effects on the dominant herbivore species, Daphnia pulicaria and D. mendotae. 3. Fish had no effect on phytoplankton biomass or overall Daphnia density. We observed a compensatory response to predation by functionally comparable species within a trophic level in the form of shifting dominance and coexistence of Daphnia species. We hypothesized that this phenotypically plastic response to predation decoupled a potential trophic cascade in this freshwater pelagic system. Daphnia pulicaria density decreased over time with fish predation, but D. mendotae density increased over time with fish predation. 4. Phenotypically plastic life history trait shifts and reproductive rates differed between species in fishless and fish enclosures, accounting for population trends. Daphnia pulicaria were also proportionally higher in walleye larvae stomachs than in the enclosures, indicating that walleye preferred to feed on D. pulcaria over D. mendotae. The resultant shift in dominance may partially explain the overall benign effect of fish on grazers and supports the hypothesis that trophic level diversity can decouple a trophic cascade.     

Trophic interactions and habitat segregation between competing Daphnia species

Summary Two commonly coexisting species of Daphnia segregate by habitat in many stratified lakes. Daphnia pulicaria is mostly found in the hypolimnion whereas D. galeata mendotae undergoes diel vertical migration between the hypolimnion and the epilimnion. I examined how habitat segregation between these two potentially competing species might be affected by trophic interactions with their resources and predators by performing a field experiment in deep enclosures in which I manipulated fish predation, nutrient levels, and the density of epilimnetic Daphnia. The results of the experiment indicate that habitat use by D. pulicaria can be jointly regulated by competition for food from epilimnetic Daphnia and predation by fishes. Patterns of habitat segregation between the two Daphnia species were determined by predation by fish but not by nutrient levels: The removal of epilimnetic fish predators resulted in higher zooplankton and lower epilimnetic phytoplankton densities and allowed D. pulicaria to expand its habitat distribution into the epilimnion. In contrast, increased resource productivity resulted in higher densities of both Daphnia species but did not affect phytoplankton levels or habitat use by Daphnia. The two species exhibit a trade-off in their ability to exploit resources and their susceptibility to predation by fish. D. g. mendotae (the less susceptible species) may thus restrict D. pulicaria (the better resource exploiter) from the epilimnion when fish are common due to lower minimum resource requirements than those needed by D. pulicaria to offset the higher mortality rate imposed by selective epilimnetic fish predators. D. g. mendotae does not appear to have this effect in the absence of fish.     

Contrasting patterns of body size for Daphnia species that segregate by habitat

Summary We investigate how body size of two coexisting Daphnia species varies among 7 lakes that represent a gradient of predation risk. The two species segregate vertically in stratified lakes; D. galeata mendotae is typically smaller and more eplimnetic than D. pulicaria. The extent of vertical habitat partitioning, however, varies seasonally within and among lakes in apparent response to predation intensity by epilimnetic planktivorous fishes. Daphnia pulicaria uses the epilimnion at low levels of fish predation but is restricted to the hypolimnion under high fish predation, whereas D. galaeta mendotae always utilizes the epilimnion. The species display contrasting patterns of genetic variation in neonate size and size at maturity. D. pulicaria is larger in lakes with higher fish and Chaoborus densities whereas D. galeata mendotae is smaller. This contrast in body size in lakes with high predation is associated with greater habitat segregation in those lakes. In lakes with low predation risk, the two species are similar in body size at birth and maturity.Authorship order alphabetical     

Formation of morphological defences in response to YOY perch and invertebrate predation in two <Emphasis Type="Italic">Daphnia</Emphasis> species coexisting in a mesotrophic lake

This study examined the formation of morphological defences by two coexisting Daphnia species, the large-sized D. pulicaria (2 mm) and the small-sized D. mendotae (1.4 mm), in response to the presence of young-of-the-year (YOY) yellow perch (Perca flavescens) and invertebrate predators (Chaoborus, Leptodora) during summer in a mesotrophic lake. We hypothesized that due to differential size-selective predation risk by YOY fish and invertebrates, the large-sized and the small-sized Daphnia species would show different morphological responses to predation threats. We followed changes in two morphological traits (relative length of the tail spine in D. pulicaria and of the helmet in D. mendotae) among different periods during summer according to YOY fish and invertebrate predation. We defined four YOY fish predation periods based on the presence of YOY perch in the pelagic zone of the lake and the relative abundance of Daphnia preys in their gut contents, and two invertebrate predation periods based on exclusive or mutual occurrence of the invertebrate predators. The large-sized (D. pulicaria) and the small-sized (D. mendotae) species showed different morphological responses to YOY fish and invertebrate predators, respectively. The tail spine ratio of the juveniles and adults of D. pulicaria did not change in response to YOY fish predation or to invertebrate predation. A gradual increase in the helmet ratio was observed in the small-sized D. mendotae over the summer period. This change was related to the co-occurrence of the invertebrate predators (Chaoborus and Leptodora) and to YOY fish predation. The warmer temperature cannot be accounted for helmet elongation since it was constant across depths, and not related with the co-occurrence of D. mendotae and YOY perch. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Cyclomorphosis inDaphnia galeata mendotae: variation and stability in phenotypic cycles

Phenotypic change is studied in a cyclomorphicDaphnia galeata mendotae population known from previous studies to be clonally diverse. Morphological analyses revealed cyclical changes in both adult and juvenile helmet length and tailspine length which were: 1) strongly correlated with mean water temperature; and 2) repeated annually during the 3-year study period. Field populations exhibited high (5% to 30%) coefficients of variation (CV) for both helmet length and tailspine length; the CV also exhibited seasonal fluctuations such that it was lowest in late summer. The period of highestDaphnia helmet development coincides with peak densities of the common invertebrate predatorsChaoborus andLeptodora. The fluctuations in CV may be due to both differential phenotypic expression of the coexisting clones and invertebrate predation.     

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.     

The effect of suspended sediments on Lake Texoma Daphnia: field distributions and in situ incubations

1. We measured the abundance and eggs per female of four Daphnia species in turbid and relatively clear regions of Lake Texoma (Oklahoma‐Texas, U.S.A.) on 12 dates over the course of 5 years. 2. Two species, Daphnia lumholtzi and Daphnia parvula, occurred and reproduced in turbid locations, but two other species, Daphnia mendotae and Daphnia pulicaria, occurred almost exclusively in relatively clear conditions. 3. To test the hypothesis that interference with foraging excludes clear‐water Daphnia species from turbid locations, we incubated adult D. mendotae at both a clear and a turbid site. In three successive experiments D. mendotae individuals incubated at the turbid site carried as many or more eggs than individuals incubated at the clear site.     

Variation in ephippial buoyancy in Daphnia pulicaria

1. Freshwater zooplankton often produce diapausing eggs to survive environmental stress. The diapausing eggs of Daphnia (Crustacea, Cladocera) are encased in an ephippium that either floats at the surface or sinks to the sediment. These two types of ephippia may represent different strategies between spatial (floaters) and temporal (sinkers) dispersal of offspring. 2. We observed floating and sinking characteristics of ephippia obtained from eight lakes. We then conducted an experiment with 26 Daphnia pulicaria clones obtained from six of these lakes and observed the production of buoyant and non‐buoyant ephippia under high and low food conditions. 3. Ephippia were more often non‐buoyant than buoyant both from females caught in nature and those reared in the laboratory. The experiment revealed that each clone was able to produce both types of ephippia, but that there was considerable among‐clone variation in the percentage of non‐buoyant ephippia produced. 4. We conclude that production of non‐buoyant versus buoyant ephippia may be driven by both genetic and environmental factors.     

Maternal and direct effects of natural-like changes of photoperiod and food condition manipulation on life history parameters in <Emphasis Type="Italic">Daphnia</Emphasis>

This study analyzed changes of life history parameters of a Daphnia pulicaria clone in response to increasing (spring conditions) versus decreasing (fall conditions) photoperiod and low and high food concentrations in maternal and offspring environments. One treatment increased the photoperiod from 12 to 16 h at a rate of 3 min per day, while the other treatment decreased the photoperiod from 16 to 12 h at the same rate. We also used two food concentrations: low (0.2 mg C/l) and high (0.5 mg C/l). Maturation time, first clutch size, survivorship, and somatic growth in offspring depended on the direction of photoperiodic changes for Daphnia grown under high food concentration. Natural-like changes of photoperiod on life history parameters and abrupt changes resulted in similar effects on life history parameters of Daphnia (see Alekseev and Lampert 2004 Hydrobilogia 526:225–230).     

Distribution and ecology of Demospongiae from the circalittoral of the islands of the Aegean Sea (Eastern Mediterranean)

Size-selective predation by fish is often considered to be a primary driver of seasonal declines in large-bodied Daphnia populations. However, large Daphnia commonly exhibit midsummer extinctions in ponds lacking planktivorous fish. A number of empirical and theoretical studies suggest that resource competition and its interaction with nutrient enrichment may determine variable dominance by large Daphnia. Low resource levels may favor competitive dominance by small-bodied taxa while large Daphnia may be favored under high resource conditions or following a nutrient/productivity pulse. Nutrient enrichment may also influence the strength of invertebrate predation on Daphnia by affecting how long vulnerable juveniles are exposed to predation. We investigated these hypotheses using an in situ mesocosm experiment in a permanent fishless pond that exhibited seasonal losses of Daphnia pulex. To explore the effects of nutrient enrichment, Daphnia plus a diverse assemblage of small-bodied zooplankton were exposed to three levels of enrichment (low, medium, and high). To explore the interaction between nutrient enrichment and invertebrate predation, we crossed the presence/absence of Notonecta undulata with low and high nutrient manipulations. We found no evidence of competitive reversals or shifts in dominance among nutrient levels, Daphnia performed poorly regardless of enrichment. This may have been due to shifts in algal composition to dominance by large filamentous green algae. Notonecta had significant negative effects on Daphnia alone, but no interaction with nutrient enrichment was detected. These results suggest that Daphnia are not invariably superior resource competitors compared to small taxa. Though predators can have negative effects, their presence is not necessary to explain poor Daphnia performance. Rather, abiotic conditions and/or resource-based effects are probably of greater importance.     

Reproduction strategies of <Emphasis Type="Italic">Daphnia pulicaria</Emphasis> population in a high mountain lake of Southern Spain

For three consecutive years, a population study of Daphnia pulicaria was undertaken in Río Seco Lake, a small high-mountain lake, in order to elucidate the reproductive strategies adopted by Daphnia in this system. Daphnia appears to colonize this lake every spring by hatching from ephippia and reproduce by means of subitaneous (non-diapausing) and ephippial (diapausing) eggs. D. pulicaria in this lake is an obligate parthenogenetic population. There is a short time period for subitaneous egg production and a much longer period for ephippial egg production. The contribution of subitaneous eggs to Daphnia population density and structure appears to be low. Diapause onset showed a high temporal synchronization in the three studied years in Río Seco Lake, and day-length emerged as the main cue triggering diapause onset and the main explanatory factor for the proportion of ephippial females observed. The development and reproduction of D. pulicaria in Río Seco Lake involves taking a gamble on resting forms to guarantee inter-annual Daphnia persistence in the lake, giving priority to investment in future generations. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Morphology of the Daphnia nervous system: A comparative study on Daphnia pulex,Daphnia lumholtzi,and Daphnia longicephala

The freshwater crustacean Daphnia is well known for its phenotypic plasticity, in which environmental cues are perceived by the nervous system and transformed into phenotypic adaptations beneficial under current conditions. Critical knowledge regarding the distribution and localization of neuronal antigens or neurotransmitters and differentially expressed proteins is sparse. Here, we applied immunohistochemical and histological‐staining methods in combination with confocal laser scanning microscopy on whole mount preparations in Daphnia pulex, Daphnia longicephala, and Daphnialumholtzi. We document the nervous system, elements of the sensory system, and cell clusters with secretory characteristics in the Daphnia head. This is the first report on the nervous system of the species D.longicephala and D.lumholtzi. The methods established in this investigation will help to foster research on specific neuronal mechanisms in this rapidly advancing model system of life science research. J. Morphol., 2012. © 2012 Wiley Periodicals, Inc.     

Environmental correlates of the intrinsic rate of natural increase in primates

Morphological and life history traits of two clones of the cladoceran Daphnia pulex were measured in the presence and absence of size-selective insect predators, the midge larva Chaoborus flavicans, which preys on small Daphnia, and the water bug Notonecta glauca, which preys on large Daphnia. The aim was to detect predator-induced phenotypic changes, particularly the effect of simultaneous exposure to both types of predators. Other work has shown that in the presence of Chaoborus americanus, Daphnia pulex produce a socalled neck spine which may carry several teeth. The morphological modifications are supposed to serve as an anti-predator device. Furthermore, females exposed to Chaoborus often delay their maturation; this has been interpreted as a cost that balances the benefits of the neck teeth. In this investigation, females of both clones produced fewer but larger offspring than control animals when reared in the presence of Chaoborus flavicans. The offspring showed the typical neck spine and delayed first reproduction. In the presence of Notonecta glauca, one of the clones produced more and smaller offspring, and maturation occurred at earlier instars. The other clone also produced more offspring than the control but there was no size difference. When both predators were present, in most cases the reactions of the daphnids were similar to those in the Notonecta experiment. The response to Chaoborus appeared to be suppressed. The observed modifications are interpreted as evolved strategies that reduce the impact of size-selective predation. They are consistent with predictions of life-history theory.     

<Emphasis Type="Italic">Daphnia</Emphasis>species diversity in Kenya,and a key to the identification of their ephippia

The distribution of Daphniaspecies in tropical Africa is poorly known and understood. Daphniaare assumed rare in tropical regions, but systematic studies covering large areas are sparse. We sampled the active community (live zooplankton) and/or the dormant community (diapausing egg banks in the sediment) of 41 standing water bodies in Kenya in search for Daphnia.Overall the dormant communities yielded 11 species of Daphnia, a species richness more than twice the species richness found in the active communities. Dormant community species diversity better reflects the spatial, and particularly the temporal (multi-annual) variation in environmental conditions available to Daphniain these tropical standing waters. Hence, we suggest that the dormant community be taken into account when assessing local zooplankton diversity, especially in fluctuating tropical lake ecosystems, where the presence of each local Daphniaspecies in the active community may be strongly seasonal or erratic. Geographic distribution data from this study are supplemented with previous records of Daphniain East Africa to provide an overview of the known distribution of Daphniain Kenya and neighbouring countries. We also present a detailed key for morphological identification of the ephippia of the 11 Daphniaspecies encountered, complemented with photographs and drawings of diagnostic characters.     

Seasonal dynamics and interspecific competition in Oneida Lake Daphnia

I investigated the population dynamics and competitive interactions of two species of the suspension-feeding crustacean Daphnia in Oneida Lake, N.Y. Both species have persisted in the lake for decades, but their water-column densities are negatively correlated. The larger Daphnia pulicaria dominates in some years, the smaller D. galeata mendotae in others, and in some years one species replaces the other seasonally. Although this pattern results in part from annual variation in vertebrate predation pressure, predation alone cannot explain the irregular daphnid dynamics. In 1992–1995, I examined the water-column abundances, birth and death rates of both species. D. pulicaria dominated in two years, D. galeata mendotae was replaced by D. pulicaria in one year and in 1994, both species persisted in low numbers. To test the effect of temporal changes on the strength of intra- and interspecific competition on both juvenile and adult daphnids, I manipulated a series of field enclosures in 1994 and 1995. The outcome of competition varied within and between years, and its effects were most evident at the highest densities and lowest resource levels. For adults of both species, the effects of interspecific competition were detected more often than those of intraspecific competition. Lipid reserves (a metric of fitness) among juveniles were generally low, with those of D. galeata mendotae often being less than those of D. pulicaria. Contrary to the results of other studies examining competition in daphnids, spatial segregation and predictable within-year reversals in competitive dominance most likely do not play a large role in fostering coexistence of the Oneida Lake daphnids. Instead, coexistence of these competitors is promoted by interannual variation and long-lived diapausing eggs. Received: 20 July 1997 / Accepted: 21 November 1997     

Interclonal variation in diel horizontal migration behaviour of the water flea <Emphasis Type="Italic">Daphnia magna</Emphasis>—searching for a signature of adaptive evolution

In shallow temperate lakes, zooplankton populations may exhibit diel horizontal migration (DHM) and move towards macrophytes during the day to avoid fish. Using a natural Daphnia magna population, we undertook an experimental investigation aimed to describe the genetic variation for DHM and to study whether an adaptive micro-evolutionary response occurred to changes in macrophyte coverage and fish predation pressure through time. Twenty-seven D. magna clones were hatched from ephippia in the sediment of shallow Lake Ring, Denmark. This lake was eutrophied during the 20th century and was subject to restoration measures in the 1970s. The DHM behaviour of the clones was observed both in the presence and absence of fish kairomone. Significant interclonal variation in DHM behaviour occurred in both treatments. To study the micro-evolutionary response of the Lake Ring D. magna population, two approaches were used. First, we compared the DHM behaviour of clones derived from ephippia collected at different depths. A comparison was conducted between clones resurrected from the period of eutrophication (1960–1980) and from the period of recovery (1986–2000). A significant treatment (presence and absence of fish kairomone) × period interaction effect was identified, suggesting a significant micro-evolutionary response for DHM behaviour. The D. magna clones exhibited a significantly stronger horizontal migration response during the period of eutrophication than in the recovery phase. Second, clonal means, representing the influence of the genotype on the trait, were correlated with environmental conditions (macrophyte cover, fish predation pressure and Secchi depth). The results of this analysis also suggest that a micro-evolutionary response by Daphnia has occurred in reaction to changes in fish predation pressure. In periods with high fish predation pressure, Daphnia migrated more strongly towards the plants. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Influence of two Daphnia species on summer phytoplankton assemblages from eutrophic lakes

We conducted grazing experiments to test whether larger-bodiedDaphnia pulicaria have a different effect from smaller-bodiedDaphnia galeata mendotae on the composition of summer algalassemblages in eutrophic lakes. Three separate cubitainer experimentswere run for 5 days in a replicated factorial design utilizingtwo algal community types and the two Daphnia species. Inorganicphosphorus and nitrogen were added to prevent nutrient limitationof the algae. Both edible and inedible size fractions of chlorophylla increased in cubitainers without Daphnia spp. Grazer additionusually resulted in a reduction in edible chlorophyll; reductionswere greater in D.pulicaria cubitainers. Grazing by Daphniaspp. on presumed inedible chlorophyll was variable. Algal sizewas not always a good predictor of grazeability. The resultsof this study indicate that D.pulicaria, because of its greaterfiltration potential and ability to ingest larger particles,provides a stronger control on inedible-sized algae when comparedto equal numerical densities of D.g.mendotae. However, Aphanizomenonincreased as a response to heavy grazing pressure by D.pulicariaon other algal species. This suggests that biomanipulation effortsthat promote large-bodied Daphnia may not produce desirableresults if nutrient inputs remain high.     

Inter-genotype variation in reproductive response to crowding among Daphnia pulex

Crowding is known to have a major influence on reproduction in the freshwater microcrustacean Daphnia pulex. We analyzed reproductive output of six different D. pulex genotypes under two different density regimes in the laboratory. Four of these genotypes reproduce via obligate parthenogenesis, allowing thorough analysis of the life history strategies of some asexual lines. Among 30,109 neonate offspring and 1041 resting egg ephippia collected, several trends were evident. Crowding induced increased resting egg production and reduced neonate offspring production among all genotypes. Offspring sex ratios grew more male-biased with maternal age. The extent, but not direction, of each of these trends varied among genotypes. Offspring sex ratios, and the very direction in which they changed in response to crowding, differed significantly among genotypes with some genotypes producing more and others fewer males in response to crowding. Obligately parthenogenetic genotypes seemed to respond to the crowding stimulus in similar ways as the facultatively parthenogenetic genotypes, as expected from the sexual origins of their genomes. The inter-genotype variation in life-history traits observed in this and other investigations calls into question the common practice of extrapolating results from a single Daphnia genotype to an entire species. Our findings are considered in the context of other research in the field of environmental influences on Daphnia reproduction with a review of representative literature.     

Influence of different sized Daphnia species on chlorophyll concentration and summer phytoplankton community structure in eutrophic Wisconsin lakes

We conducted field observations on 13 eutrophic Wisconsin lakesdominated by either the larger bodied Daphnia pulicaria or thesmaller bodied Daphnia galeata mendotae. While Daphnia numericaldensities were not significantly different between groups oflakes, pulicaria lakes had much higher Daphnia biomasses andfiltration potentials than galeata lakes. Although we foundsignificant differences in chlorophyll (Chl) a between bothgroups of lakes during June, on a seasonal basis populationsof different sized Daphnia were not associated with significantdifferences in Chl a. Filtration potential per se was the majordeterminant of Chl a, regardless of which Daphnia species dominated.However, in pulicaria lakes, the clear-water phase started earlier,lasted longer, and was usually characterized by greater Secchidisc readings than in galeata lakes. For large blue-green algaesuch as Aphanizomenon, D.pulicaria appeared to delay bloom conditions,but ultimately did not prevent the alga from growing. Our resultssuggest that high densities of large-sized Daphnia are a desirablegoal of biomanipulation because they can attain filtration potentialshigh enough to increase summer water clarity in eutrophic lakes.     

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.