Life history strategies of cladocerans: comparisons of tropical and temperate taxa

We review recent works on different life history variables of cladoceran taxa in tropical and temperate freshwater bodies, comparing the strategies that cladocerans have evolved to adapt to contrasting environmental conditions in the two geographical regions. These life-history parameters relate to age and size at maturity, survival, fecundity, life-expectancy at birth, lifespan, gross, and net reproductive rates, generation time, the rate of population increase, peak population density and day of peak abundance. We also discuss the role of photoperiod and temperature on some of these life history parameters. We found a general paucity of experimental work and field data in tropics on cladocerans. There is very limited information on the few Daphnia species found in the tropics. The misconception of low species diversity of cladocerans in the tropics arose due to several reasons including lack of extensive and intensive field collections. Higher water temperatures apparently promote permanent infestation of tropical waters with toxic cyanobacteria, which reduce the zooplankton diversity. In addition to higher temperatures in the tropics, the year-round high predation pressure of planktivorous fish probably causes the tropical species, particularly in pelagic habitats, to reach maturity earlier (< 3 days) than in temperate regions. Species of Daphnia in temperate regions are particularly adapted to living at food concentrations that are much lower and seasonably more variable than those for tropical genera such as Diaphanosoma. This is further corroborated by the more than an order of magnitude higher threshold food concentration (TFC) for tropical Cladocera than for their temperate counterparts. Fecundity patterns differ between tropical and temperate cladoceran taxa: cultured under optimal temperature regimes, tropical taxa have fewer eggs than temperate species of a comparable body size. Predation pressure may act differently depending on the size of the cladoceran neonates and thus on their population size structure. Global warming and climate changes seem to affect the behaviour (migration), distribution, and abundance of cladocerans. Apparently, in direct response to these changes, the possibility of encountering the tropical cladocerans in the northern, temperate hemisphere (bioinvasions) is on the rise.     

Genotypic characteristics of the Cladocera

Work on the genetics of cladocerans reproducing by cyclic parthenogenesis has indicated that populations usually include a large number of genotypes, whose frequencies are in close approximation to Hardy-Weinberg equilibrium. Less diverse genotypic arrays and pronounced instability in genotype frequencies occur only in permanent populations exposed to limited ephippial recruitment. Genetic diversification among local cladoceran populations is greater than in most other organisms as a consequence of the inefficiency of passive dispersal. Genotypic characteristics of cladocerans reproducing by obligate parthenogenesis are markedly different from those of cyclic parthenogens. Local populations include few clones, but genetic distances between them are often large and accompanied by significant ecological and morphological divergence. When considered over their entire range, cladoceran taxa reproducing by obligate asexuality are the most genotypically diverse asexual organisms known. This diversity has originated from the spread of a sex-limited meiosis suppressor through species that originally reproduced by cyclic parthenogenesis. The confused state of cladoceran taxonomy is partially a consequence of the presence of such obligately asexual groups, but also results from the occurrence of hybridization and sibling species. The genome size of cladocerans is exceptionally small and is associated with a large amount of endopolyploidy. Somatic tissues in adult cladocerans show a range in nuclear DNA content from 2–2048 c. DNA quantification studies have additionally revealed the frequent occurrence of polyploid clones in obligately asexual taxa.     

Selective predation by Lestes (Odoiiata, Lestidae) on littoral microcrustacea

• 1 Two experimental approaches were used to examine the predation risk of six littoral cladoceran and ostracod species to two size classes of the damselfly Lestes sp. Behavioural observations were conducted in a 2–1 arena and predation rate experiments in 1–1 jars in the laboratory.
• 2 Behavioural observations revealed that attack and capture efficiencies by Lestes were higher on smaller cladocerans (Polyphemus and Ceriodaphnia) than on other taxa. Small Lestes had little success ingesting the small ostracod Cypridopsis, whereas larger Lestes captured and ate it easily.
• 3 Predation-rate experiments showed that Lestes has a clear preference for smaller cladocerans over both a large cladoceran (Simocephatus) and all three ostracods (Cypricercus, Cyclocypris, and Cypridopsis). Most Lestes were unable to consume the larger Cypricercus.
• 4 Attack rates and predation rates were highly variable among individual predators.
• 5 These data indicate that damselfly larvae are size selective and consume all but very large cladocerans more readily than ostracods.

     

Feeding behaviour of major 0 + fish species in a shallow, eutrophic lake (Tjeukemeer, The Netherlands)

The forage base and the food selectivity of 0+ representatives of six abundant freshwater fish species were studied in a shallow, eutrophic Dutch lake. Most species relied on the zooplankton; the size-selective predation in early summer was directed to the smaller copepods and in late summer to larger cladocerans and copepods than concurrently present in the lake. Daphnia spp. and cyclopoid copepods were the main zooplankton taxa for smelt, perch and pikeperch. Energetically, the large cladoceran, Leptodora kindtii, was especially important for pikeperch. Bream and roach preyed upon smaller zooplankton than the other fish species. The influence of the zooplankton predation by abundant 0+ fish was clear from a small mean Daphnia size in September; this size is to be used as an indicator in fishery management. Neomysis integer, the most important macrofauna species, was consumed by perch, pikeperch and ruffe; pikeperch was most size-selective in this respect. The 0+ ruffe was à typically benthivorous fish. Only the 0+ pikeperch became piscivorous, especially in years when smelt was abundant.     

Relative vulnerabilities of natural rotifer and ciliate communities to cladocerans: laboratory and field experiments

SUMMARY. 1. Two experiments with plankton communities from Storrs Pond (NH), one conducted in the laboratory and one in field enclosures, assessed the impact of different cladocerans on rotifers and ciliated protozoa.
2. The smallest cladoceran, Bosmina longirostris , did not depress rotifer or ciliate growth rates while the intermediate sized dadoceran, Daphnia galeata mendotae , reduced ciliate growth rates in the enclosure experiment but had only a marginal effect in the jar experiment. D. galeata mendotae had no effect on any of the rotifers in either experiment.
3. In both experiments the largest cladoceran, Daphniapulex , depressed the growth rates of ciliates and those rotifers known to be vulnerable to interference competition. Polyarthra vulgaris , previously shown to be resistant to cladoceran interference, was the only rotifer unaffected by D. pulex in the field experiment but was depressed by the much higher densities of this cladoceran in the laboratory experiment.
4. Cladocerans did not affect phytoplankton or bacterioplankton abundance in either experiment. Therefore the mechanism most likely to be responsible for the suppressive effect of cladocerans on rotifers and ciliates in these experiments is direct mechanical interference or predation, rather than exploitative competition.     

Variation in the neonate size of Daphnia pulex: the effects of predator exposure and clonal origin

Daphnia pulex clones originating from twelve small pond habitats were exposed to chemical cues from a size-selective predator, larvae of the phantom midge Chaoborus. Exposure delayed the onset of reproduction and increased the size at first reproduction. On the other hand, the neonates produced by these larger mothers were slightly smaller than the neonates produced by the smaller control mothers. In cladocerans, neonate size is usually positively correlated to the size of the mother. Thus exposure to Chaoborus kairomone apparently had direct effects on neonate size counterbalancing the maternal effects. Daphnia clones collected from Chaoborus-free and Chaoborus-rich environments exhibited different responses. In first adult instar, the clones from Chaoborus inhabited environments increased their offspring size under Chaoborus exposure whereas clones from Chaoborus-free environments did not. This may reflect clonal adaptation to the predation prehistory of their original habitat since larger neonates more quickly reach a size protected from the predator.     

Competition between rotifers and cladocerans of different body sizes

Summary We conducted laboratory experiments to test the hypothesis that rotifers could coexist with small (<1.2 mm) but not large (>1.2 mm) cladocerans. Keratella cochlearis was excluded in <8 days by the large cladocerans Daphnia pulex and D. magna, probably through both interference and exploitative competition. On the other hand, K. cochlearis persisted for 8 weeks with two small cladocerans (Bosmina longirostris and Ceriodaphnia dubia) and excluded a third small cladoceran (D. ambigua). Similarly, Synchaeta oblonga coexisted with B. longirostris for >7 weeks, and K. testudo coexisted with D. ambigua for >4 weeks. Coexistence of small cladocerans and rotifers was always accompanied by suppression of one or both populations, probably primarily if not exclusively by exploitative competition for limiting food resources. These results indicate that the competitive dominance of cladocerans over rotifers decreases markedly with cladoceran body size and that factors other than body size may determine the competitive outcome between rotifers and small cladocerans. Our study provides a mechanistic explanation for a commonly observed pattern in natural zooplankton communities: planktonic rotifers often are abundant when only small cladocerans occur but typically are rare when large cladocerans are present.     

Refuge availability and sequence of predators determine the seasonal succession of crustacean zooplankton in a clay-turbid lake

The contribution of predators and abiotic factors to the regulation of the biomass and seasonal succession of crustacean zooplankton was studied in Lake Rehtijärvi (southern Finland). Field data in combination with bioenergetics modeling indicated that invertebrate planktivory by Chaoborus depressed cladoceran populations during early summer. In particular, bosminids that generally form the spring biomass peak of cladocerans in stratified temperate lakes did not appear in the samples until July. In July, predation pressure by chaoborids was relaxed due to their emergence period and cladoceran population growth appeared to be limited by predation by planktivorous fish. The effect of fish predation was amplified by reduced refuge availability for cladocerans. The concentration of dissolved oxygen below the epilimnion was depleted, forcing cladocerans to move upward to less turbid and thus more dangerous water layers. The effect of size selective predation by fish resulted in reduced mean size of cladocerans during the period when refuge thickness (thickness of the water layer with oxygen concentration <1 mg l^?1 and water turbidity >30 NTU) was lowest. The results confirmed that in clay-turbid lakes, invertebrate predators could be the main regulators of herbivorous zooplankton even when cyprinid fish are abundant.     

轮虫和枝角类的种间竞争及其影响因子研究进展

轮虫和枝角类是浮游动物群落的重要组成部分和优势类群,它们之间的竞争互作是调节水生态系统结构和功能的主要动力之一。普遍的观点认为,轮虫和大型枝角类难以共存,往往被竞争排斥,而和小型枝角类可以共存;实际上,轮虫和枝角类的种间竞争结局存在一定的不确定性。介绍了轮虫和枝角类的种间竞争方式及其相对重要性,对影响轮虫和枝角类种间竞争结局的因素,包括温度、食物、相对起始密度、个体大小、食物临点、耐饥饿能力、捕食及竞争者和捕食者释放的化感物质等,进行了系统的梳理和分析,并提出了今后亟待解决的科学问题和研究切入点。     

Predation pressure induced by seasonal fishing moratorium changes the dynamics of subtropical Cladocera populations

Seasonal fishing moratorium with selective fish harvesting is a common means of increasing fish production in Asia. Such manipulation might be expected to lead to a seasonal variation in abundance and composition of fish and predation pressure on zooplankton, and consequently a seasonal change in the Cladocera populations. In this study, we examined the seasonal dynamics of cladocerans in a subtropical plateau lake in southwestern China, Lake Erh, in which a fishing moratorium was implemented in spring and early summer. Cladocerans showed a bimodal seasonal distribution linked to variations in predation pressure induced by the seasonal fishing moratorium. When predation was low in winter because of cool water temperatures and decreasing fish density, Daphnia galeata appeared in the water column and increased towards spring. The cladoceran peaked in the middle of the fishing moratorium, coinciding with a clear water phase. The feeding rate increased with temperature in the late moratorium, allowing planktivorous fish to strongly suppress large daphnids. Small cladocerans benefited from the reduced competition with large daphnids by increasing their numbers rapidly, which resulted in a second peak of cladocerans in the fishing season.     

Zooplankton community structure driven by vertebrate and invertebrate predators

Summary A zooplankton community was established in outdoor experimental ponds, into which a vertebrate predator (topmouth gudgeon: Pseudorasbora parva) and/or an invertebrate predator (phantom midge larva: Chaoborus flavicans) were introduced and their predation effects on the zooplankton community structure were evaluated. In the ponds which had Chaoborus but not fish, small- and medium-sized cladocerans and calanoid copepods were eliminated while rotifers became abundant. A large-sized cladoceran Daphnia longispina, whose juveniles had high helmets and long tailspines as anti-predator devices, escaped from Chaoborus predation and increased. In the ponds which had fish but not Chaoborus, the large-sized Daphnia was selectively predated by the fish while small-and medium-sized cladocerans and calanoid copepods predominated. In the ponds containing both Chaoborus and fish, the fish reduced the late instar larvae (III and IV) of Chaoborus but increased the early instar larvae (I and II). Small- and large-sized cladocerans were scarcely found. The former might have been eliminated by predation of the early instar larvae of Chaoborus, while the latter was probably predated by fish. Consequently, the medium-sized cladocerans, which may have succeeded in escaping from both types of predator, appeared abundantly. The results suggest that various combinations of vertebrate and invertebrate predators are able to drive various kinds of zooplankton community structure.     

Long‐term dynamics of a cladoceran community from an early stage of lake formation in Lake Fukami‐ike,Japan

An increase in nutrient levels due to eutrophication has considerable effects on lake ecosystems. Cladocerans are intermediate consumers in lake ecosystems; thus, they are influenced by both the bottom‐up and top‐down effects that occur as eutrophication progresses. The long‐term community succession of cladocerans and the effects cladocerans experience through the various eutrophication stages have rarely been investigated from the perspective of the early‐stage cladoceran community assemblage during lake formation. In our research, long‐term cladoceran community succession was examined via paleolimnological analysis in the currently eutrophic Lake Fukami‐ike, Japan. We measured the concentration of total phosphorus and phytoplankton pigments and counted cladoceran and other invertebrate subfossils in all layers of collected sediment cores, and then assessed changes in the factors controlling the cladoceran community over a 354‐year period from lake formation to the present. The cladoceran community consisted only of benthic taxa at the time of lake formation. When rapid eutrophication occurred and phytoplankton increased, the benthic community was replaced by a pelagic community. After further eutrophication, large Daphnia and high‐order consumers became established. The statistical analysis suggested that bottom‐up effects mainly controlled the cladoceran community in the lake''s early stages, and the importance of top‐down effects increased after eutrophication occurred. Total phosphorus and phytoplankton pigments had positive effects on pelagic Bosmina, leading to the replacement of the benthic cladoceran community by the pelagic one. In contrast, the taxa established posteutrophication were affected more by predators than by nutrient levels. A decrease in planktivorous fish possibly allowed large Daphnia to establish, and the subsequent increase in planktivorous fish reduced the body size of the cladoceran community.     

Summer zooplankton dynamics in the limnetic and littoral zones of a humic acid lake

The limnetic and littoral zooplankton were studied during summer 1989 in Triangle Lake, a humic acid (pH 4.9) bog lake in Ohio, USA. The limnetic zooplankton showed low species richness and biomass, and dominance by the rotifers Kellicottia bostoniensis and Polyarthra vulgaris. In the littoral, species richness and biomass were markedly higher, and the crustaceans Alona guttata, Ceriodaphnia quadrangula, Chydorus sphaericus, Simocephalus serrulatus and Tropocyclops prasinus, and the rotifer Ascomorpha ecaudis were the dominants. The extreme rotifer dominance and lack of cladocerans in the limnetic zone were likely the result of Chaoborus predation. A pronounced mid-summer decline in cladoceran biomass in the littoral was likely due to predation by T. prasinus and Utricularia (bladderwort).     

Applicability of planktonic biomanipulation for managing eutrophication in the subtropics

Although large-bodied cladocerans such asDaphnia can reduce algal biomass significantly in temperate lakes if freed from fish predation, the applicability of such biomanipulation techniques for eutrophication management in the subtropics and tropics has been examined only recently. Subtropical cladoceran assemblages differ from those of temperate lakes by their low species richness, early summer gameogenesis, and greatly reduced body size. Eutrophic Florida lakes are dominated by pump-filter feeding fish rather than by size selective planktivores as a temperate lakes. Cladocerans in Florida lakes can increase in abundance significantly if freed from fish but fail to have an impact on algal biomass or composition. The greatest potential for using biomanipulation to manage phytoplankton-dominated lakes in the subtropics and tropics lies with phytophagous fish. Future research should concentrate on defining the role of individual fish taxa on phytoplankton composition and community structure, nutrient cycling, and planktonic productivity before embarking on whole lake manipulation projects.     

CHAOBORUS PREDATION AND LIFE-HISTORY EVOLUTION IN DAPHNIA PULEX: TEMPORAL PATTERN OF POPULATION DIVERSITY,FITNESS, AND MEAN LIFE HISTORY

The effect of predation by the aquatic dipteran larva Chaoborus americanus on genetic diversity and life-history evolution in the cladoceran Daphnia pulex was investigated in large replicate laboratory populations. Instantaneous daily loss rates of clonal diversity and genetic variance for fitness indicate that 93–99% of initial genetic diversity can be removed from populations during the 8–12 generations of clonal reproduction that occur each year in natural populations. In the absence of predation, the principal evolved changes in mean population life history were smaller immature body size and increased and earlier fecundity. In the presence of size-selective Chaoborus predation, populations evolved toward larger body size and increased and earlier reproduction. The difference between these two trajectories is an estimate of the direct additive effect of Chaoborus predation. This effect was manifested as evolution toward larger body size with a trend toward earlier and increased reproduction.     

Cladocera assemblages from the surface sediments of south-central Ontario (Canada) lakes and their relationships to measured environmental variables

Fossil cladoceran remains preserved in surface sediment samples from 44 oligotrophic lakes in south-central Ontario were examined to evaluate the relationships between species assemblages and measured environmental variables. Differences in cladoceran assemblages were related to physical and chemical variables using multivariate techniques. Redundancy Analysis (RDA) identified five environmental variables as significantly influencing assemblage composition: sulphate (SO₄²⁻), calcium (Ca²⁺), pH, maximum lake depth (Z _max) and dissolved organic carbon (DOC). There was a distinct separation of lakes and taxa along the ion gradient based on SO₄, Ca and pH. Additionally, cladoceran communities in coloured, shallow lakes had relatively higher abundances of littoral chydorid species and the pelagic taxa Holopedium spp., and the Daphnia pulex complex. Deep, clear lakes had relatively higher abundances of other pelagic taxa. Predation by fish (measured as presence–absence) and Chaoborus (measured as density) were less significant than some of the physico-chemical variables in influencing cladoceran assemblage structure. However, this could be due to the limited resolution of the predation data that was available at the time of this study. The distribution of cladocerans in the surface sediment, and their relation to these important environmental variables, suggests that there is considerable potential for the use of sedimentary cladoceran remains as environmental indicators in south-central Ontario lakes. Handling editor: J. Saros     

Prey selectivities of threespine sticklebacks (Gasterosteus aculeatus) and phantom midge larvae (Chaoborus spp.) in Newfoundland lakes

SUMMARY. 1. Threespine stickleback ( Gasterosteus aculeatus ) and Chaoborus ( C. punctipennis and C. trivittatus ) planktivore populations were sampled from several oligotrophic lakes on the Avalon Peninsula of Newfoundland, Canada. Pelagic zooplankton communities in these lakes consist of Daphnia catawba, Holopedium gibberum, Eubosmina longispina, Leptodiaptorma minutus, Epischura nordenskioldi, Cyclops scutifer and rotifers ( Conochilus unicornis, Keratella and Kellicottia spp.).
2. Comparison of gut contents of G. aculeatus and crop contents of later instar Chaoborus larvae with lake zooplankton communities indicated that the vertebrate and invertebrate plank tivores show contrasting species and size selectivities in their choice of zooplankton prey. Prey selectivities were quantified by means of the selectivity index α.
3. G. aculeatus showed positive selectivity for larger zooplankton (greater than 0.76 mm in length), typically the cladoceran D. catawba , while Chaoborus larvae showed positive selectivity for smaller zooplankton (less than 0.50 mm in length), typically the cladoceran E. longispina . While rotifers never featured in the diet of the sticklebacks, these zooplankters were a common component of the chaoborid diet. H. gibberum , with its gelatinous sheath, was rarely found in the gut contents of G. aculeatus and was never found in the crop contents of Chaoborus larvae.
4. Selective predation by these planklivores in the relatively shallow and well-mixed oligotrophic Avalon lakes may influence species composition and size structure of the pelagic zooplankton communities.     

Zooplankton populations and the diets of three Chaoborus species (Diptera, Chaoboridae) in a tropical lake

SUMMARY. 1. Zooplankton populations in a small, natural, tropical lake are dominated by a few, small-sized taxa including the copepod Thermo- cyclops consimilis , the cladoceran Moina micrura and several rotifer species.
2. Moina micrura and adults of Thermocyclops consimilis undergo diel vertical migrations within the water column. Population densities of T. consimilis show marked intra-annual variations which may to some extent be related to variations in rainfall and to lunar periodicities In predator abundance.
3. The diets of Chaoborus lavae include other Chaoborus , Cladocera, Copepoda. Rotifera and the dinoflagellate Peridinium. The diets of late instars of the largest species, edulis , were dominated by Crustacea, while those of the two smaller species. C, ceratopogones and C. anomalus , were dominated by Rotifera and the dinoflagellate alga Peridinium , as were the diets of early instars of all species. Algae have not been previously reported to be a large component of the diet of Chaoborus populations in nature.
4. Some of the dietary differences among Chaoborus instars and species are related to the size of each prey species in relation to the mouth gape of each instar. However, there are also important differences in electivity among instars of different species of the same size.     

Fish-mediated indirect effects in a littoral food web

The strength of the direct effect by scraping cladocerans and the indirect effect of nutrient regeneration by filtering herbivorous cladocerans on periphyton growth was investigated in a littoral food web. Ten enclosures were erected in a lake in an area with artificial vegetation. Five enclosures were stocked with juvenile perch ( Perca fluviatilis ) and five lacked fish. In addition, a reference area in the artificial vegetation was sampled. The mesh size of the net surrounding the cages was chosen to allow an inflow of phytoplankton into the cages from the surrounding water. The periphyton and filtering herbivorous cladoceran biomasses were highest in the fish-free treatment. There was no difference in phytoplankton biomass between treatments despite the large difference in filtering herbivorous cladoceran biomass, suggesting that the inflow of phytoplankton into enclosures completely compensated for the loss due to filtering. The reference area and the enclosure with fish showed the same patterns in developments with respect to filtering herbivorous cladocerans and periphyton. Scraping cladoceran biomass was higher in the fish-free treatment resulting in a positive correlation between scraping cladoceran and periphyton biomass. Our results suggest that the positive indirect effect of filtering herbivorous cladoceran nutrient regeneration on periphyton was stronger than the negative direct effect of grazing by scraping cladocerans on periphyton in this semi-open system, and that pelagic production by phytoplankton may foster periphyton growth in the littoral habitat via filtering herbivorous cladocerans. Furthermore, heterogeneity within trophic levels involving primary producers of different growth forms such as phytoplankton and periphyton may enhance the potential for compensatory responses via nutrient recycling.     

A quantitative test of the size efficiency hypothesis by means of a physiologically structured model

According to the size‐efficiency hypothesis (SEH) larger bodied cladocerans are better competitors for food than small bodied species. In environments with fish, however, the higher losses of the large bodied species due to size‐selective predation may shift the balance in favor of the small bodied species. Here we present a theoretical framework for the analysis of the competitive abilities of zooplankton species that takes both competition and predation into account in one coherent analysis. By applying the conceptually well‐understood framework of physiologically structured population models we were able to predict the relative difference in predation rates necessary to cause a shift in dominance of the large‐bodied species (Daphnia pulicaria) to the small‐bodied species (D. galeata). These predictions depend only on seven easily interpretable parameters per species: size at birth, size at maturity and maximum size, age at maturity, maximal clutch size, egg development time and finally the half‐saturation constant for food. The critical equilibrium mortality of D. pulicaria was 0.16 d^?1 at food concentrations close to the critical food concentration of D. galeata, i.e. D. pulicaria will win the competition as long as its mortality rate is below 0.16 d^?1. At higher food concentrations the differential mortality curve (plotting equilibrium mortalities of both species against each other) approached a linear function with a slope of one and an intercept equal to the difference in maximal population birth rates. The prediction of critical predation rates was independent of the ingestion rate of the cladocerans and the algal carrying capacity and food regeneration rate of the environment although the mechanism works through competition for a shared algal food resource. We interpret these findings in terms of the relative predation risk large and small‐bodied cladocerans will face in various freshwater ecosystems.