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.     

Notes on Cladocera and Copepoda from high altitude lakes in the Mount Everest Region (Nepal)

The paper deals with Cladocera and Copepoda species from eight high altitude lakes in the Khumbu area. In all lakes, an endemic known diaptomid of the palearctic genus Arctodiaptomus is found. The dark and large Daphnia tibetana occurs in five of the lakes sampled, characterized by the persistence of the spine in adults (parthenogenetic and ephippial females, males). Apparently, this is an unknown feature of this species. The literature on the diagnostic traits of the different morphs described is reviewed. In addition, a transparent and smaller-sized Daphnia species occurs in two lakes. This is a D. longispina characterized by the absence in adults of the carapace spine (var. aspina Weretschagin, 1911). The presence of these two species is discussed in relation to water transparency, colour, and vertical distribution. Two hypotheses on the evolution of cuticular pigmentation in Daphnia are examined. In addition to these mostly dominant species, a macrothricid also typical of high altitude lakes in the Alps was found (Macrothrix hirsuticornis) together with two cosmopolitan Chydoridae.     

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     

Differences in genetic structure and ecological diversity between parental forms and hybrids in a Daphnia species complex

Hybridization is a common phenomenon in Daphnia species complexes. Hybrids often dominate in Daphnia populations; therefore it is worthwhile to look for principal differences between parental and hybrid populations with respect to their genetic structure and clonal differentiation. We studied natural populations of members of the Daphnia galeata/hyalina/cucullata complex in three lakes. In one of these lakes, one parental species (D. galeata) and one hybrid (D. galeata × cucullata) were investigated more intensively. The frequency of sexual reproduction was higher in parental populations, whereas clonal diversity was higher in hybrid populations. Ecological differentiation among clonal groups was more pronounced in the D. galeata × cucullata hybrid compared to D. galeata, whereas selection intensity was weaker. These results are discussed with respect to stability of clonal groups, multiple hybridizations and selective constraints.     

Zooplankton seasonal dynamics in a recently filled mine pit lake: the effect of non-indigenous <Emphasis Type="Italic">Daphnia</Emphasis> establishment

We examined the temporal and vertical dynamics of zooplankton in Weavers Lake, New Zealand, between October 2004 and October 2005, at a time when it was colonised by a non-indigenous Daphnia species. Zooplankton community composition changed during the study from one of rotifer dominance (e.g. Asplanchna, Polyarthra, Brachionus and Keratella species) to cladoceran (Daphnia dentifera) dominance. Temporal changes in zooplankton community composition were strongly associated with a gradual increase in lake water clarity, and were attributable to the highly efficient filter feeding of D. dentifera. The corresponding reduction in rotifer densities may have resulted from the superior competitive abilities of the newly established Daphnia. As Daphnia were rare inhabitants of New Zealand lakes before 1990, the arrival and rapid spread of the non-indigenous D. dentifera has lead to widespread changes in both water clarity and zooplankton community composition. An apparent lack of mixing in the lake was facilitated by the lake’s extremely small surface area:depth ratio. However, we conclude that physical features of the lake had minimal influence on water clarity relative to the invasion of D. dentifera.     

Experimental interspecific hybridization in Daphnia

Hybridization is a common phenomenon in Daphnia (Cladocera; Anomopoda); interspecific hybrids have been found between several species and hybrids are found in many European lakes. Although much information on the morphology, ecology and genetics of hybrids is available, little is known about the level of reproductive isolation among species or about the relative fitness of hybrids and parental species. In order to facilitate studies on differentiation and speciation processes and comparative experimental studies on hybrids and recombinant genotypes, we present the first successful laboratory crossing experiments of two different Daphnia species, D. galeata and D. cucullata. Males and sexual females from two D. galeata and two D. cucullata clones were reciprocally crossed, juveniles hatched from resting eggs and reared until maturity. Hatching and juvenile survival rates of hybrids were relatively low (12.1% and 24%, respectively). D. galeata and D. cucullata clones vary in their level of successful interspecific matings and in the number of subsequent offspring. In general, hybrid crosses between D. cucullata females and D. galeata males were more successful than reciprocal crosses.     

The genusDaphnia in Cameroon,West Africa

Three species of the genusDaphnia are recorded from Cameroon.Daphnia obtusa was found in a single lake out of 37 sampled.Daphnia laevis was found in the same lake asD. obtusa and in two other lakes, whileD. rosea was found in three other lakes. All the lakes were at altitudes over 1 000 m. The records can be interpreted either as altitudinal survivals of continually dispersing species, or as relicts in an area whereDaphnia is usually very rare or absent.     

Four different head shapes in Daphnia hyalina (Leydig) induced by the presence of larvae of Chaoborus flavicans (Meigen)

In many species of Daphnia spines, neck teeth, and enlarged or reshaped helmets are well-known as defences against invertebrate predators. Until now, Daphnia hyalina (Leydig, 1860), a common species in many European lakes, has appeared to be an exception to this rule.Here, we provide evidence that the larvae of Chaoborus flavicans (Meigen) also can induce morphological changes in D. hyalina. Specimens react morphologically to the presence of larvae in three ways: (a) by changes in the shape of head shield, (b) by the formation of spine(s) on the head, and (c) by the elongation of the tail spine. The frequency and intensity of these morphological changes are correlated positively with midge larvae densities. The most pronounced reactions occurred in young D. hyalina.The predator kairomone also induced changes in Daphnia body size.     

Daphnia community analysis in shallow Kenyan lakes and ponds using dormant eggs in surface sediments

1. Water fleas of the genus Daphnia are considered rare in tropical regions, and information on species distribution and community ecology is scarce and anecdotal. This study presents the results of a survey of Daphnia species distribution and community composition in 40 standing waterbodies in southern Kenya. The study sites cover a wide range of tropical standing aquatic habitats, from small ephemeral pools to large permanent lakes between approximately 700 and 2800 m a.s.l. Our analysis combines data on Daphnia distribution and abundance from zooplankton samples and dormant eggs in surface sediments. 2. Nearly 70% (27 of 40) of the sampled waterbodies were inhabited by Daphnia. Although their abundance in the active community was often very low, this high incidence shows that Daphnia can be equally widespread in tropical regions as in temperate regions. 3. Analysis of local species assemblages from dormant eggs in surface sediments was more productive than snapshot sampling of zooplankton communities. Surface‐sediment samples yielded eight Daphnia species in total, and allowed the detection of Daphnia in 25 waterbodies; zooplankton samples revealed the presence of only four Daphnia species in 16 waterbodies. 4. Daphnia barbata, D. laevis, and D. pulex were the most frequently recorded and most abundant Daphnia species. Canonical correspondence analysis of species–environment relationships indicates that variation in the Daphnia community composition of Kenyan waters was best explained by fish presence, temperature, macrophyte cover and altitude. Daphnia barbata and D. pulex tended to co‐occur with each other and with fish. Both species tended to occur in relatively large (>10 ha) and deeper (>2 m) alkaline waters (pH 8.5). Daphnia laevis mainly occurred in cool and clear, macrophyte‐dominated lakes at high altitudes.     

Relative effects of Daphnia and Ceriodaphnia on phosphorus-chlorophyll relationships in small urban lakes

Empirical models based on zooplankton biomass were used to predict mean summer chlorophyll a (Chl a) and to examine how zooplankton influenced the total phosphorus (TP) - Chl a relationship. Four years of data were analyzed for three lakes having similar TP concentrations but varied abundances of Daphnia and Ceriodaphnia. Mean TP did not correlate significantly with mean Chl a during the study period, although mean Daphnia density was a good predictor of Chl a concentration (p > 0.001). Both residuals from the TP - Chl a relationship (p > 0.001) and Secchi depth (p > 0.007) were negatively correlated with Daphnia abundance. Ceriodaphnia abundance was positively correlated with Chl a (p > 0.002) and Secchi depth (p > 0.001). Mean size of Daphnia during spring was the best predictor of the Daphnia-Ceriodaphnia shift in mid-summer. Early establishment of a large-sized Daphnia cohort may prevent their summer elimination by Chaoborus and intensify competition with Ceriodaphnia. These results imply an important link between Daphnia and Ceriodaphnia thereby limiting the utility of Chl a - TP model predictions in these small, urban lakes. This linkage and the differential effect of these two zooplankton species on planktonic algae deserve further consideration in similar lakes where phytoplankton and zooplankton tend to be tightly coupled.     

<Emphasis Type="Italic">Daphnia</Emphasis> distribution in Andean Patagonian lakes: effect of low food quality and fish predation

Food quality in terms of carbon (C):phosphorus (P) ratio can constrain the success of highly demanding P herbivores as Daphnia. North Andean Patagonian lakes are ultraoligotrophic with low nutrient concentrations and well-developed euphotic zones. We investigated the distribution of the large Daphnia commutata in relation with food quality (sestonic C:P ratio) and predation risk in these lakes. The predation risk was estimated based on the fish species present and their relative eye diameter and transparency of the lake. The C:P ratios in the lakes were high, varying from 350 to >1,200. The lakes with D. commutata had significantly lower C:P ratio than those without these daphnids. On the other hand, those lakes where Daphnia is present have the lower predation risk than those were Daphnia is absent. In addition, we carried out growth experiments with neonates and natural seston of three lakes with different C:P ratio. The growth rates were inversely related with C:P of the food. Food quality and predation risk together determined the success or failure of large Daphnia populations in these Andean clear ultraoligotrophic lakes.     

Cyclomorphosis in Daphnia: an adaptation to avoid invertebrate predation

Seasonal morphological changes in three Daphnia species were followed over a two-year period in two lakes that differ in invertebrate and fish pressure. Whereas the morphology of D. hyalina, the biggest of the three species, varied little from season to season, D. cucullata, the smallest, exhibited the most pronounced seasonal changes in head height/carapace length ratio. The pattern of seasonal changes of body proportions was similar in all size classes and isometric growth of the head was reported for D. cucullata. Unlike the head, tail spine length/carapace length ratio almost did not vary seasonally. Strong negative allometry of tail spine growth was observed. These results are consistent with the hypothesis that helmets and tail spines provide protection against invertebrates in the two smallest, thus most endangered species.     

Species-specific population-density thresholds in cladocerans?

The population density of a Daphnia species seems more dependent on properties specific to the species than on those specific to the point in the season, location within a lake basin or the given lake itself. In spite of week-to-week fluctuations, the population density for each of two common European Daphniaspecies was found to be remarkably similar within single lakes (from station to station on a single date, and from date to date at a single station) as well as from lake to lake, regardless of trophic state. All lakes on all dates revealed densities in the range 10–50 ind l^–1 for the smaller-bodied D. cucullata and 1–5 ind l^–1 for the larger-bodied D. hyalina, in spite of different intensity of reproduction resulting from different food levels (chlorophyll a between 0.2 and 4.2 g/l). It can be asserted that the population density of each species remains far below the carrying capacity of the habitat K, and does not depend on food levels, which merely set the rate of population increase, while the population density reflects the species' vulnerability to predation by planktivorous fish. The reactive distance (the distance over which a foraging fish can see its prey) in 1+ roach, a dominant planktivore in the lakes studied, has been found to be twice as great for D. hyalina as for D. cucullata, whatever the light intensity. The relative reactive field volume was therefore an order of magnitude greater for the former than for the latter, showing that the densities of the two prey species would be assessed by a foraging roach as equal when, in reality, they differed by an order of magnitude, as they do in various lakes and in various seasons.The first of the two conclusions is that whatever the growth and reproduction in a population of a cladoceran such as Daphnia, its density would be fixed by mortality induced by fish predation. The second would be, that the difference between the bottom-up and top-down effects in freshwater is more than merely the upward or downward direction along the food web, since the bottom-up effects are about the flow control (the rate of net production, individual growth rate, rate of reproduction) and the top-down about the standing-crop control (biomass, individual body size, population density level).     

Status of Daphnia Resting Egg Banks in Bohemian Forest Lakes Affected by Acidification

We studied the egg banks of Daphnia gr. longispina in four acidified lakes in the Bohemian Forest (Czech Republic, Europe). Daphnia had become extinct in three of the studied lakes due to anthropogenic acidification and we assessed the possibility of autochthonous recovery of the population after recent amelioration of the lake environment. We determined the ephippia distribution in eleven cores 10–30 cm long, and the state of Daphnia resting eggs in over 13 000 ephippia from eight cores. Apparently well-preserved eggs were used for hatching experiments and for DNA amplification. Vertical profiles of ephippia densities in several cores did not agree with historical data on the Daphnia presence in the lakes, as the sediment had been repeatedly disturbed by human activities in the past. Ephippia are present near the surface of the sediment in all lakes, and they might therefore receive hatching cues. We were not able, however, to prove that viable eggs are present in the ephippia banks of the lakes. The percentage of the well-preserved eggs in all but one core was below 1%. In addition, no egg hatched in the hatching experiments and we were not able to amplify DNA from the preserved eggs of lakes where Daphnia is extinct, although amplification from relatively young eggs from the fourth lake was successful in 60% of the cases. We conclude that the recovery of Daphnia populations in studied lakes from autochthonous sources is unlikely.     

Using morphological characters of subfossil daphniid postabdominal claws to improve taxonomic resolution within species complexes

Daphnia subfossils from lake sediments are useful for exploring the impacts of environmental stressors on aquatic ecosystems. Unfortunately, taxonomic resolution of Daphnia remains is coarse, as only a small portion of the animal is preserved, and so the identification of daphniid subfossils typically relies upon postabdominal claws. Daphniid claws can be assigned to one of two species complexes: D. longispina or D. pulex. Both complexes contain species with differing environmental optima, and therefore improved taxonomic resolution of subfossil daphniid claws would aid paleolimnological analyses. To identify morphological features that may be used to help differentiate between species within complexes, we used species presence/absence data from net tows to select lakes in central Ontario (Canada) containing only a single species from a particular complex, then used remains preserved in surface sediments of these lakes to isolate four Daphnia species: D. ambigua and D. mendotae from the D. longispina complex, and D. pulicaria and D. catawba from the D. pulex complex. Our analyses demonstrate that, within the D. longispina complex, postabdominal claw length (PCL) and spinule length can be used to distinguish D. mendotae from D. ambigua. In addition, within the D. pulex complex, there are differences between D. pulicaria and D. catawba in the relative lengths of the proximal and middle combs on the postabdominal claw. However, the number of stout spines on the middle comb is an unreliable character for differentiating species. Overall, our data demonstrate that greater resolution within Daphnia species complexes is possible using postabdominal claws; however, the process is arduous, and applicability will likely decrease with the number of taxa present.     

Competition or niche segregation between Holopedium and Daphnia; empirical light on abiotic key parameters

A survey of 346 Norwegian lakes revealed some consistent patterns with regard to the occurrence of Holopedium gibberum versus Daphnia spp. Both biomass and frequency of Holopedium were negatively correlated with pH and Ca-concentration, and this species rarely occurred at pH > 8, or Ca-concentrations > 10 mg l^–1. On the contrary, the various Daphnia species, although showing different susceptibilities to fish predation and different preferences for lake productivity, were all positively correlated with Ca. Statistical analysis as well as laboratory experiments and biochemical considerations, suggest that Holopedium may be a superior competitor at very low Ca-concentrations, but is replaced by Daphnia at higher pH and increasing Ca-content. High pH in itself is possibly disadvantageous for Holopedium by preventing formation of the muco-polysaccaride mantle.     

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.

     

Planktivory by bluegill (Lepomis macrochirus) on Leptodora kindti in a small North American lake

Leptodora kindti (Crustacea: Cladocera) is a large species of zooplankton (2–18 mm length) that is exceptionally transparent. This transparency is believed to be a means by which it successfully coexists in lakes with planktivorous fishes. We investigated the gut remains of bluegill (Lepomis macrochirus) that had been feeding on L. kindti and Daphnia (D. galeata and D. retrocurva) in the wild (Lake Zurich, Illinois) and found that bluegill readily preyed on L. kindti as small as 3–5 mm length, and strongly selected L. kindti over Daphnia galeata and Daphnia retrocurva. The large compound eye of L. kindti is one half to one complete order of magnitude larger than Daphnia's eye, consistent with the hypothesis that eye area is an important visual cue for fishes. Moreover, the slope of the relationship between eye area and body length is an order of magnitude shallower in L. kindti than Daphnia, suggesting that eye area has been under stronger negative selection in L. kindti. Results suggest that L. kindti's large and dark eye compromises the transparent nature of its body.     

Differences in allozyme patterns between Diaphanosoma brachyurum and Diaphanosoma mongolianum,as revealed in Central European populations

Recent taxonomic revisions in the genus Diaphanosoma have made the uncertainty in species discrimination with morphological characters obvious. Therefore species characterization on a genetic basis seems to be required.Here we examined the genetic structure of two Diaphanosoma mongolianum populations and three Diaphanosoma brachyurum populations in Central Europe by allozyme electrophoresis. A genetic differentiation between both species was evident. 5 out of 9 tested loci carried diagnostic alleles. Both species differed in their habitat choice: D. mongolianum was adapted to higher trophic levels than D. brachyurum. Co-occurrence was observed in a eutrophic lake. Populations sampled from their preferred habitats showed high genotype diversities.Most genotype frequency distributions were in agreement with Hardy-Weinberg expectations and temporal (between years) changes in genotypic compositions were not common. This indicated annually occurring bouts of sexual reproduction, similar to intermittent Daphnia populations. Genetic differentiation among lakes suggested restricted gene flow.     

Patterns and sources of variation in Daphnia phosphorus content in nature

It has recently been shown that Daphnia can vary in the phosphorus (P)-content of their body tissues, but the relative importance of genetic versus environmental causes for this variation is unexplored. We measured variation in P-content (as % body mass) of Daphnia from eight lakes and conducted experiments to contrast three sources of variation: interspecific variation, clonal variation and phenotypic plasticity. Daphnia P-content decreased with increasing seston C:P ratio across lakes. This relationship reflected both inter- and intraspecific variation. Daphnia parvula and D. dubia exhibited high P-content and were found in shallow lakes with low C:P seston, whereas D. pulicaria had low P-content and was found in deep, stratified lakes having high C:P seston. Populations of D. dentifera spanned this lake gradient and exhibited P-content that was negatively related to seston C:P. Evidence for phenotypic plasticity came from experiments with D. pulicaria and D. dentifera collected from a lake with P-deficient seston and fed a P-sufficient diet in the laboratory. In addition, populations of D. dentifera differed in P-content even after 7 d of feeding on P-sufficient resources, suggesting within-species clonal variation. However, mesocosm experiments revealed broad and surprisingly continuous variation in the P-content of individual clones of D. pulex (range 1.54–1.05%) and D. mendotae (1.51–1.07%) over a gradient in dietary C:P. The broad range in P-content exhibited by individual clones, acclimated for generations, suggests that variation in Daphnia P-content from laboratory experiments needs to be interpreted with caution. These results also show that phenotypic variation in response to environment can be a larger source of variation in P-content than genetic differences within or among species.