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     

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.     

The possibility of predator avoidance by Lake Michigan zooplankton

Low densities of Diaptomus ashlandi, Diacyclops thomasi, and Daphnia galeata mendotae were measured at depths where Mysis relicta formed nighttime aggregations. Calculations suggest that mysid predation can not account for the rarity of prey animals at these depths, which further suggests that the prey avoided the mysids. Unlike D. galeata mendotae, Daphnia pulicaria was common in mysid aggregations. The somewhat larger size of D. pulicaria may reduce its vulnerability to mysid predation, and consequently may explain the vertical distribution differences between the two congeners. Vertical distributions of Limnocalanus macrurus and copepod nauplii showed no obvious relationships to the mysid distributions. These were the only two taxa with distributions that were correlated with chlorophyll a concentrations. All crustacean taxa were rare in the epilimnion at night when sonar recorded a dense fish school.     

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.     

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     

Phenotypic Plasticity in Life-history Traits and Feeding Appendages in Two Species of Daphnia Fed a Natural Phytoplankton Assemblage

The ability of two cladoceran species – Daphnia pulicaria and D. mendotae – to utilize a natural phytoplankton assemblage dominated by cyanobacteria was investigated experimentally. Reference animals were fed high quality green algae. The natural phytoplankton assemblage originated from Oneida Lake (NY, USA). Both cladoceran species occur in the lake, but their densities vary temporally. As expected, lake phytoplankton were a poor food source. Fecundity decreased, and age at first reproduction and offspring size increased. The magnitude of these effects varied between and within species. The increase in offspring size differs from studies using toxic cyanobacterial strains, but agrees with expectations on adaptive responses to low food availability. Both species performed equally on the lake phytoplankton, but D. pulicaria appeared to be the superior species in the high quality food treatment.     

Temperature and kairomone induced life history plasticity in coexisting Daphnia

We investigated the life history alterations of coexisting Daphnia species responding to environmental temperature and predator cues. In a laboratory experiment, we measured Daphnia life history plasticity under different predation risk and temperature treatments that simulate changing environmental conditions. Daphnia pulicaria abundance and size at first reproduction (SFR) declined, while ephippia (resting egg) formation increased at high temperatures. Daphnia mendotae abundance and clutch size increased with predation risk at high temperatures, but produced few ephippia. Thus, each species exhibited phenotypic plasticity, but responded in sharply different ways to the same environmental cues. In Glen Elder reservoir, Kansas USA, D. pulicaria dominance shifted to D. mendotae dominance as temperature and predation risk increased from March to June in both 1999 and 2000. Field estimates of life history shifts mirrored the laboratory experiment results, suggesting that similar phenotypic responses to seasonal cues contribute to seasonal Daphnia population trends. These results illustrate species-specific differences in life history plasticity among coexisting zooplankton taxa.     

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.     

Testing the ecological relevance of Daphnia species designations

1. Molecular approaches have increasingly revealed hidden genetic structure within ecologically important species, leading to the creation of sibling species whose ecological relevance is often unclear. A prime example is Daphnia galeata mendotae, which was split into D. dentifera and D. mendotae based on differences at two allozyme loci. 2. In a set of lake populations in Michigan USA, we test the geographical and temporal consistency of the genetic structure underlying this species split. We also test the morphological relevance of this molecular variation and its ecological significance in lakes. In essence, we ask: does recognition of these new species provide valuable information for plankton ecologists? 3. We found that D. dentifera and D. mendotae represent morphologically and ecologically distinct forms that are distributed among lakes in non‐random fashion, which were remarkably stable over 6 years. Key differences between the species concern their body and head shape, vertical habitat use within lakes and distribution among lakes of different size. We hypothesise that these differences represent specialisation to habitats that differ in risk of invertebrate predation. 4. Reproductive barriers alone are insufficient to explain the pattern of genetic structure; in some lakes complete introgression is apparent. However, parent species and hybrids exhibit a stable co‐existence in many lakes, which suggests that ecological specialisation reinforces divergence within this taxon.     

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.     

Competition in natural populations of Daphnia

I investigated the competitive relationships between two species of Daphnia, D. galeata and D. cucullata, and their interspecific hybrid. The term hemispecific competition was introduced to describe competition between parental species and hybrids. In eutrophic Tjeukemeer both parental species were found to compete with the hybrid, whereas competition between D. galeata and D. cucullata seemed limited. Although the effect of competition on life history traits of daphnids may be profound, the influence of the competitors on the seasonal dynamics of the Daphnia species seems limited.     

Stocking piscivores to improve fishing and water clarity: a synthesis of the Lake Mendota biomanipulation project

SUMMARYY 1. A total of 2.7 × 10⁶ walleye fingerlings and 1.7 × 10⁵ northern pike fingerlings were stocked during 1987–99 in eutrophic Lake Mendota. The objectives of the biomanipulation were to improve sport fishing and to increase piscivory to levels that would reduce planktivore biomass, increase Daphnia grazing and ultimately reduce algal densities in the lake. The combined biomass of the two piscivore species in the lake increased rapidly from < 1 kg ha^?1 and stabilised at 4–6 kg ha^?1 throughout the evaluation period. 2. Restrictive harvest regulations (i.e. increase in minimum size limit and reduction in bag limit) were implemented in 1988 to protect the stocked piscivores. Further restrictions were added in 1991 and 1996 for walleye and northern pike, respectively. These restrictions were essential because fishing pressure on both species (especially walleye) increased dramatically during biomanipulation. 3. Commencing in 1987 with a massive natural die‐off of cisco and declining yellow perch populations, total planktivore biomass dropped from about 300–600 kg ha^?1 prior to the die‐off and the fish stocking, to about 20–40 kg ha^?1 in subsequent years. These low planktivore biomasses lasted until a resurgence in the perch population in 1999. 4. During the period prior to biomanipulation when cisco were very abundant, the dominant Daphnia species was the smaller‐bodied D. galeata mendotae, which usually reached a biomass maximum in June and then crashed shortly thereafter. Beginning in 1988, the larger‐bodied D. pulicaria dominated, with relatively high biomasses occurring earlier in the spring and lasting well past mid‐summer of many years. 5. In many years dominated by D. pulicaria, Secchi disc readings were greater during the spring and summer months when compared with years dominated by D. galeata mendotae. During the biomanipulation evaluation period, phosphorus (P) levels also changed dramatically thus complicating our analysis. Earlier research on Lake Mendota had shown that Daphnia grazing increased summer Secchi disc readings, but P concentrations linked to agricultural and urban runoff and to climate‐controlled internal mixing processes were also important factors affecting summer readings. 6. The Lake Mendota biomanipulation project has been a success given that high densities of the large‐bodied D. pulicaria have continued to dominate for over a decade, and the diversity of fishing opportunities have improved for walleye, northern pike and, more recently, yellow perch. 7. Massive stocking coupled with very restrictive fishing regulations produced moderate increases in piscivore densities. Larger increases could be realised by more drastic restrictions on sport fishing, but these regulations would be very controversial to anglers. 8. If the lake's food web remains in a favourable biomanipulation state (i.e. high herbivory), further improvements in water clarity are possible with future reductions in P loadings from a recently initiated non‐point pollution abatement programme in the lake's drainage basin.     

Effects of Daphnia on microzooplankton communities

Intact phytoplankton and microzooplankton communities from eutrophicStar Lake were incubated for 4 days with and without Daphniapulex, Daphnia galeaia mendotae, or a natural assemblage ofDaphnia species. They were sampled at the onset and terminationof the experiment for bacterial, phytoplankton, ciliate, rotifer,copepod and cladoceran densities. The cladocerans had variedeffects on the rotifers, ranging from significant suppressionof most rotifer species (Keratella cochlearis, Polyarthra remata,Keratella crassa) in the D.pulex jars, to the suppression ofone (K.crassa) or no species in the D.galeata mendotae and StarLake Daphnia assemblage jars, respectively. Small ciliates (<30µm, longest dimension), such as Strobilidium sp. and Pseudo-cyclidiumsp., were adversely affected by most of the cladoceran treatments,while several larger ciliates (>81 µm) were unaffectedin all such treatments. Ciliates were not consistently morevulnerable to cladoceran suppression than similarly sized rotifers.The suppression of ciliates and rotifers was attributable toboth direct effects (predation, interference, or both) and indirecteffects (e.g. resource competition) of the cladocerans. ¹Present address: Department of Biology, University of Louisville,Louisville, KY 40292, USA     

Impact of fish predation on coexisting Daphnia taxa: a partial test of the temporal hybrid superiority hypothesis

Fish predation was tested as a factor mediating the coexistence of Daphnia taxa in the shallow, hypertrophic Lake Blankaart. Naturally co-occurring populations of D. galeata and the hybrid D. galeata x cucullata were subjected to different levels of fish predation in in situ enclosures. In control enclosures without fish, the largest taxon D. galeata rapidly became dominant over the intermediate sized D. galeata x cucullata, mainly as a result of higher birth rates. In enclosures with fish, population densities of D. galeata dropped relative to D. galeata x cucullata, due to higher mortality rates. These results are in concordance with the `temporal hybrid superiority hypothesis', and can be explained by a higher vulnerability of the large and more conspicuous D. galeata to the size selective predation exerted by visually hunting planktivorous fishes. After approximately one month, however, population growth rates of D. galeata and D. galeata x cucullata in the enclosures with fish converged, due to a relative reduction in the mortality rate of D. galeata. This suggests that, in the presence of fish, D. galeata may co-exist with hybrids due to a decrease in its relative vulnerability to visual predation with time. Indeed, both D. galeata and the hybrid showed strong reductions in adult body size in the enclosures with fish, but this size reduction tended to be stronger in D. galeata than in D. galeata x cucullata. In addition, turbidity increased in the enclosures with fish and may additionally have reduced the relative advantage of D. galeata x cucullata with regard to mortality caused by visual predation.     

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.     

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.     

Coexistence of plant species in a biodiversity hotspot is stabilized by competition but not by seed predation

Understanding the mechanisms of species coexistence is a key task for ecology. Recent theory predicts that both competition and predation (which causes apparent competition among prey) can either promote or limit species coexistence. Both mechanisms cause negative interactions between individuals, and each mechanism promotes stable coexistence if it causes negative interactions to be stronger between conspecifics than between heterospecifics. However, the relative importance of competition and predation for coexistence in natural communities is poorly known. Here, we study how competition and apparent competition via pre‐dispersal seed predators affect the long‐term fecundity of Protea shrubs in the fire‐prone Fynbos biome (South Africa). These shrubs store all viable seeds produced since the last fire in fire‐proof cones. Competitive effects on cone number and pre‐dispersal seed predation reduce their fecundity and can thus limit recruitment after the next fire. In 27 communities comprising 49 990 shrubs of 22 Protea species, we measured cone number and per‐cone seed predation rate of 2154 and 1755 focal individuals, respectively. Neighbourhood analyses related these measures to individual‐based community maps. We found that conspecific neighbours had stronger competitive effects on cone number than heterospecific neighbours. In contrast, apparent competition via seed predators was comparable between conspecifics and heterospecifics. This indicates that competition stabilizes coexistence of Protea species, whereas pre‐dispersal seed predation does not. Larger neighbours had stronger competitive effects and neighbours with large seed crops exerted stronger apparent competition. For 97% of the focal plants, competition reduced fecundity more than apparent competition. Our results show that even in communities of closely related and ecologically similar species, intraspecific competition can be stronger than interspecific competition. On the other hand, apparent competition through seed predators need not have such a stabilizing effect. These findings illustrate the potential of ‘community demography’, the demographic study of multiple interacting species, for understanding plant coexistence.     

Phylogenetics and Evolution of theDaphnia longispinaGroup (Crustacea) Based on 12S rDNA Sequence and Allozyme Variation

Although members of the crustacean genusDaphniahave been the target of much research, there is little understanding of the group's evolutionary history. We addressed this gap by inferring a phylogeny for one of the major species groups (longispina) using nucleotide sequence variation of a 525-bp segment of the mitochondrial 12S rDNA and allozyme variation at 21 loci. We identified the major lineages and their relationships, assessed the phylogenetic utility of the few morphological characters in the group, and examinedDaphniaphylogeography. Nuclear and mtDNA phylogenies were generally concordant in recognizing the same four species complexes. An exception was the position ofDaphnia galeata mendotae.The allozyme tree paired this species with theDaphnia rosealineage, whereas the mtDNA trees groupedD. g. mendotaewithDaphnia galeata galeata.This discordance was consistent with the reticulate evolution of nuclear genes supporting the hypothesis thatD. g. mendotaerepresents a case of homoploid hybrid speciation. Striking morphological stasis in thelongispinagroup was evidenced by its very limited morphological divergence over an estimated 100 MY, and by the unusual transitional saturation of the conservative 12S rRNA gene within a species group. Phylogenetic inference also provided evidence that similarities in cephalic crest shape likely resulted from convergent or parallel evolution among species. Endemism at the continental level was indicated for previously cosmopolitan species, but the estimated times of these divisions were inconsistent with vicariance events suggesting recent dispersal among continents. A significant role for divergent selection in new habitats during speciation was suggested by the neighboringly sympatric distributions of four sister species pairs over broad geographic areas.     

Contrasting migration behaviour of Daphnia pulicaria and D. galeata x hyalina, in avoidance of predation by 0+perch (Perca fluviatilis)

In spring and early summer, a small population of the large-bodied Daphnia pulicaria coexists with a much larger population of the medium-sized hybrid Daphnia galeata × hyalina in the epilimnion of Lake Maarsseveen (The Netherlands). When large shoals of juvenile perch (Perca fluviatilis) appear in the open water, both species start to migrate vertically. Since D. pulicaria has a larger body-size than D. galeata × hyalina, and is therefore competitive dominant over the hybrid, it is unlikely that both species interact via their common food resource, but they react both to fish predation. However, since they differ in size, and therefore in vulnerability for fish predation, both species adopt different strategies. The smaller bodied, and less vulnerable D. galeata × hyalina exhibits diel vertical migration ascending to the surface at dusk, and staying there during the night. In this way, benefiting from the higher temperatures of the surface layers. In contrast, the large-bodied, and more vulnerable D. pulicaria selects the deep cold hypolimnion water layers as refuge against fish predation. In this way it benefits from a safe habitat, free from fish predators, but on the other hand suffers from low water temperatures, which decrease its fitness. It is likely that the relatively higher temperature in the upper water layers serves as a proximate factor for the downward migration of D. pulicaria.