Habitat segregation and interactive effects of multiple predators on a prey assemblage

1. In a field experiment we examined the interactive effects of two common predators of zooplankton, bluegill sunfish (Lepomis macrochirus) and Chaoborus spp. on the growth rate and habitat use of three congeneric prey species (Daphnia). Bluegill and Chaoborus both consume Daphnia, but bluegill also prey on Chaoborus. The prey species, Daphnia pulicaria, D. rosea and D. retrocurva, differed in body size and vertical distribution. We expected the largest species, D. pulicaria, to be most vulnerable to fish predation and the smallest species, D. retrocurva, to be most vulnerable to Chaoborus predation. 2. As we expected, the population growth rate of D. pulicaria was significantly reduced by fish. However, Chaoborus also significantly reduced the growth rate of this species. No significant interaction effect was detected, indicating that the effect of these predators was additive. The growth rates of D. rosea and D. retrocurva were significantly reduced by Chaoborus, but a significant interaction effect indicated that the effect of Chaoborus was stronger in the absence of fish than when fish were present. Therefore the impact of Chaoborus and fish on D. rosea and D. retrocurva was non-additive. The interactive effect of the two predators on D. retrocurva was greater in magnitude than on D. rosea. 3. In the absence of predators, the three Daphnia species showed no differences in mean habitat depth between day and night. Both predators significantly affected diel habitat use of D. pulicaria and D. rosea. Fish caused both of these Daphnia species to move deeper during the day, whereas Chaoborus caused Daphnia to move into shallower water at night. Daphnia retrocurva tended to migrate upwards at night in all predator treatments, but no significant differences in migration were observed among the predator treatments. The effects of predators on habitat use were not interactive for any prey species. 4. Our results suggest that body size, habitat use and the diel migratory response to predators are important factors mediating the interactive effects of multiple predator types on zooplankton.     

Optimising survival under predation: chemical cues modify curvature in Daphnia galeata

Morphological responses to the presence of predator info-chemicals havebeen described for many Daphnia (Cladocera) species, butD. galeata is generally considered to exhibit almost nomorphological changes that could increase its fitness under predation.Therefore, the aim of our study was to examine the nature and magnitude ofmorphological responses of D. galeata to their predatorsindetail and assess their potential role in decreasing the predation threat. Twoclones of Daphnia were exposed to predator info-chemicals(kairomones) from perch, a fish (Perca), and a phantommidge larvae (Chaoborus) an invertebrate, and a kairomone mixture fromboththese organisms. Laboratory life-table experiments were carried out and fiveparameters characterising the body shape of the daphnids were measured: helmetlength, head- and carapace width, eye diameter and body size. The last-namedthree parameters did not differ significantly between the clones or thetreatments. The differences found between the clones were significant for headwidth and helmet length, but only in combination with the treatment effects.Ourresults on genotype-dependent phenotypic plasticity indicated that, althoughphenotypic plasticity is present, the clonal composition of aDaphnia population can be altered by selection on themorphotype. This potential for a change in clonal frequencies is given by thedifferences measured between the two clones in head width and helmet length,altering the curvature of the Daphina body in response tokairomone presence.     

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.     

The effect of parthenogenic eggs in Daphnia magna on prey location by the bluegill sunfish (Lepomis macrochirus)

The distance at which the bluegill sunfish (Lepomis macrochirus) can locate Daphnia magna with parthenogenic eggs is shown to be significantly greater than the reactive distance for non-gravid Daphnia of the same size. This difference is due to greater visibility of the gravid prey and not to different locomotor behavior, since there was no correlation between the number of eggs carried by a Daphnia and the antennal beat frequency. Based on this experiment and other observations, an explanation is given for selective predation of polymorphic cladoceran populations.     

Changes in abundance of the icefish Neosalanx pseudotaihuensis Zhang (Salangidae) and the impact on the zooplankton community of Xujiahe Reservoir,central China

The icefish Neosalanx pseudotaihuensis Zhang is a small and commercially exploited zooplanktivorous fish in Xujiahe Reservoir (central China). Data of catch per unit effort (CPUE) from June 1995 to December 1997 show that the abundance of the icefish varied dramatically, which is reflected in the populations of the dominant cladocerans. The densities of Leptodora kindti, Daphnia galeata and Diaphanosoma dubia were the highest in 1997 when the abundance of the icefish was the lowest, and the reverse was the case for Boimina spp. These results document that the population of this annual fish is very variable and high biomass of the fish could exert a pronounced effect on the zooplankton community.     

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.     

Molecular Genetic Diagnostics of Some Daphnia Species (Crustacea,Cladocera) from the Volga River

Population genetic structure of the widespread Daphnia species from Kuibyshev and Saratov reservoirs was examined using RAPD–PCR technique with four mitochondrial DNA (16S rRNA) specific primers. One of the Daphnia clones examined from the Volga region appeared to be conspecific to the North American Daphnia galeata, while another clone was most likely a hybrid between D. galeata and D. cucullata.     

Diurnal periodicity in Daphnia reproduction

The diurnal periodicity of reproduction in a reservoir population of Daphnia galeata Sars was tested by two methods (egg age distribution analyses and direct observation of hatching). No significant diurnal synchronization of reproduction in the natural cladoceran population was found. A conspicuous shortage of older embryonic stages was parallely revealed by analyses of diurnal changes of egg age distribution. Suggested selective feeding of fish on gravid females of Daphnia brooding embryos with black eye pigmentation is discussed.     

Dynamics of Daphnia hyalina × galeata in Castelo-do-Bode Reservoir: The effect of food availability and flatworm predation

Daphnia hyalina × galeata in Castelo-do-Bode Reservoir was regulated mainly by food availability during the summers of 1993 and 1994. Because of high temperatures and low rainfall in 1994, the reservoir was, apparently, more eutrophic relatively to 1993. Higher chlorophyll-a concentrations and higher clutch size in 1994 suggest that Daphnia population benefited from better feeding conditions in 1994. The analysis of the contributions from fecundity and adult proportion to birth rate variation suggest that vertebrate predation and predation by copepods were negligible factors as population organisers. A general decrease in the abundance of large crustaceans during the periods of high density of typhloplanid flatworms, in both years, points to the importance of flatworm predation on zooplankton. A decrease in the number of individuals in all Daphnia size classes was observed during the periods of flatworms population increase.     

Climate-driven warming during spring destabilises a Daphnia population: a mechanistic food web approach

Temperature-driven changes in interactions between populations are crucial to the estimation of the impact of global warming on aquatic food webs. We analysed inter-annual variability in two data sets from Bautzen reservoir, Germany. In a long-term data set (1981–1999) we examined the pelagic phenology of Daphnia galeata, a keystone species, the invertebrate predator Leptodora kindtii, phytoplankton and Secchi depth in relation to water temperature and the North Atlantic Oscillation index. In a short-term data set (1995–1998) we examined food web relations, particularly the consumption of D. galeata by young-of-the-year (YOY) percids and L. kindtii and rates of population change of D. galeata (abundance, recruitment pattern and non-consumptive mortality). The start of the clear-water stage (CWS) was correlated with winter temperatures. It started 5.8 days earlier per degree warming after warm winters (mean January–March temperature ≥2.5°C) compared to cold winters (mean temperature <2.5°C). However, the end of the CWS remained relatively constant. Predation by L. kindtii and YOY percids on D. galeata started distinctly earlier, i.e. by 13.0 and 6.5 days per degree warming, respectively, in years when the average May temperature was high (≥14°C) compared to years when it was low (<14°C). Significant reductions of Daphnia abundance in midsummer occurred only in years in which the mean May temperature exceeded 14°C. This temperature regime resulted in a match of over-exploitation of food resources by Daphnia during the CWS and strong predation by YOY percids and L. kindtii. Consumptive mortality increased at higher rates with a rise in temperature than net recruitment, resulting in lower Daphnia densities at the end of the CWS. Our data suggest that even low warming by 1.7°C during a short, but critical seasonal period, resulting in the coincidence of two or more factors adversely affecting a keystone species, such as Daphnia, may induce changes in whole lake food webs and thus alter entire ecosystems. Priority programme of the German Research Foundation—contribution 9.     

Size-dependent habitat choice in Daphnia galeata Sars and size-structured interactions among zooplankton in a subarctic lake (lake Lombola, Norway)

Seasonal changes in vertical distribution of Daphnia galeata and other zooplankters were monitored in lake Lombola (69° 07&prime; N). Depth-habitat use, availability of edible algae and zooplankton densities were recorded to examine seasonal changes in intensity of competition between Daphnia and the other herbivores in the lake. Early in July, the exephippial generation of Daphnia aggregated near the surface, independently of body-size. In late July, when fish planktivory was expected to increase, the daphnids moved down during the day. In August, as intraspecific competition for food intensified, small and large Daphnia partitioned the water column, with larger individuals staying deeper. In September, Daphnia became dominated by large individuals, edible phytoplankton reached the seasonal minimum, and the vertical distribution of Daphnia gradually stretched out towards the surface. The observations on food availability and zooplankton densities suggest that interspecific competition intensified by the end of July. Species and stages that were most exposed to exploitative and interference competition by Daphnia were those staying deeper, because their vertical distribution overlapped more with the larger, competitively superior daphnids. These susceptible competitors included Keratella cochlearis and Synchaeta, among the rotifers, and nauplii and early copepodite stages of Cyclops scutifer. Depth-habitat use is discussed in relation to copepod development, zooplankton dynamics and predator-mediated coexistence.     

No Evidence for Adaptive Micro-Evolution to a Decrease in Phosphorus-Loading of a Daphnia Population Inhabiting A Pre-Alpine Lake

Many polluted lakes in Europe are being restored and phosphorus concentrations have dropped dramatically in these lakes. We studied the genetic structure of Daphnia galeata over the past 30 years in Lake Greifensee, Switzerland, a period during which the phosphorus concentration in the lake reduced dramatically. Distinct genotypes of D. galeata were hatched from diapausing eggs extracted from six different time horizons in dated sediments. We compared juvenile growth, size and egg production of D. galeata reared on high-P and low-P algae to investigate whether Daphnia have evolved to grow better on phosphorus-limited algae. Our results indicate that life histories of D. galeata differed significantly between both food types. We also found significant clone effects for size and egg number. But we found no significant interaction between the depth from which the clones were selected and food quality. This means that we found no evidence for adaptive micro-evolution in response to P reduction in the lake. We discuss our results in relation to other studies that found evidence for adaptive micro-evolution in comparable time frames.     

Feeding behaviour and morphology of filtering combs of Daphnia galeata

The population of Daphnia galeata Sars from the fish pond Velký Pálenec (Blatná, Czechoslovakia) living in high food conditions (7 mgC l^–1) was characterized by a small size of the filtering comb on the thoracic limb 3, measured as seta length, length of the base of the comb and number of setae (population 1). One month cultivation of this population in low food conditions (1.5 mgC l^–1) in the laboratory resulted in twofold increase in size of the filtering comb (population 2). Filtering and ingestion rates of both populations were measured at eight concentrations of food (approximately 0.025–3.2 mgC l^–1) using ¹⁴C labeled Scenedesmus acutus. The results show that size of the filtering combs influences considerably feeding behavior of Daphnia. The comparison of animals with the same body length suggests that the population with a large comb feeds at concentration of food below 0.4 mgC l^–1 more intensively and reaches the maximum of the filtering rate at a lower concentration than the population with a small comb. The situation is opposite at concentration above 0.4 mgC l^–1. The higher values of theoretical flow in population with a small projection of filtering area suggest that this population has to compensate disadvantage of a small comb with the higher appendages beat frequency.     

Competitive ability of Daphnia under dominance of non-toxic filamentous cyanobacteria

It is generally assumed that Daphnia is more susceptible to the inhibitory effects of filamentous cyanobacteria than small cladocerans since daphnids have a larger gape size and filtrate the filaments, whereas small cladocerans do not. This study addresses the question whether food limitation has the potential to modify this scenario of cladoceran response to dominance of non-toxic filamentous cyanobacteria. Daphnia galeatawas grown under limited (0.1 mg C l^–1) and unlimited concentrations (1.0 mg C l^–1) of high-quality food algae both in the absence/presence of non-toxic filamentous Aphanizomenon flexuosum. As the effects of these cyanobacteria on D. galeatawere positive under food limiting conditions and negative at the high food density, it was concluded that D. galeatawas mainly affected by nutritional quality due to its ability to ingest the filaments, while mechanical interference with food collection was not important. In competition experiments between D. galeataand Bosmina longirostris, D. galeatawas the dominant species at regular additions of food (1.0 mg C l^–1) in the absence of Aphanizomenon. In the presence of these cyanobacteria, D. galeatawas inhibited during the first days of the experimental period. However, the negative effect at the initially high food density was outweighed by nutrition at food limiting conditions and the outcome in competitive dominance was not changed. The results demonstrate that the ability of D. galeata to ingest large-sized non-toxic cyanobacteria can be considered as advantageous under food limiting conditions.     

Costs of Predator Avoidance Reduce Competitive Ability of Daphnia

Large-sized Daphnia hyalina×galeata hybrid and small-sized Ceriodaphnia reticulata were raised together for 3 weeks in thermally stratified flow-through tubes ('plankton organ'), in the presence and the absence of fish kairomone, and their relative performance was monitored, as indicated by the populations' growth rates (r). In the kairomone-free medium, D. hyalina grew faster than C. reticulata; the presence of kairomone reversed this ratio, producing relatively higher r in C. reticulata. The simulated fish presence reduced growth in D. hyalina population also when it was separated from the competitor, while growth in C. reticulata was altered by the kairomone only in the presence of Daphnia. It is suggested that costs of surface avoidance in Daphnia can reduce its competitive superiority to small-sized cladocerans, permanently dwelling at the surface.     

Density and size distribution of Daphnia populations at different fish predation levels

Density and size structure changes of natural daphniids populations were studied in enclosures with a different level of fish predation. Daphnia pulicaria was totaly removed in high predation variants, and the differences of the mean body length both in adults and juveniles are apparent between low predation enclosure and enclosure without fish. Daphnia galeata was replaced by D. magna and D. pulicaria in the enclosure without fish. The decline of densities and the substantial (30–50%) and fast (during 12 days) shift of mean body length both in adults and juveniles of D. galeata was induced by the fish (carp fry) introduction to the high predation enclosures.     

Life history patterns of parental and hybrid Daphnia differ between lakes

1. We investigated whether Daphnia galeata × hyalina hybrids of Lake Constance and Lake Greifensee show the same pattern of life history parameters as previously reported for D. galeata × cucullata hybrids and whether such a pattern is consistent between Daphnia populations from those two lakes. 2. Hybrids in Lake Constance were intermediate in size compared with the parental species. Hybrids in Lake Greifensee were smaller than D. galeata. The intrinsic growth rate (r) of hybrids from Lake Constance was not significantly different from the faster growing parental taxon D. galeata. However, r of hybrids from Lake Greifensee was significantly lower than that of D. galeata. 3. The observed juvenile body length differences between the taxa varied with the clutch number. The first clutch juvenile lengths of the three taxa did not differ for Lake Constance. First clutch juveniles of Lake Greifensee D. galeata were smaller than hybrid first clutch juveniles. The third clutch juvenile length did not differ between taxa from Lake Greifensee, but D. galeata juveniles from Lake Constance were bigger than those of D. hyalina. 4. The life history pattern found in Lake Constance corresponds to previous findings from other studies. The hybrids in this lake combine the faster population growth of one parental species with a relatively small size. In the case of Lake Greifensee hybrids, the relatively large size of first clutch juveniles and the small size of the adults could be interpreted as dual adaptations to invertebrate and fish predation. We speculate that the lower population growth rate of the hybrids is a trade‐off for this twofold protection.     

Fitness optimization of Daphnia in a trade-off between food and temperature

In thermally stratified lakes with a deep chlorophyll maximum (DCM), Daphnia face a trade-off between food availability and optimum development temperatures. We hypothesize that Daphnia optimize their fitness by allocating the time spent in the different vertical habitats depending on the distribution of algal resources and the temperature gradient. We used the plankton towers (large indoor mesocosms) to study the vertical distribution of a population of Daphnia hyalina×galeata in three different temperature gradients with a DCM. Additionally, we determined the fitness of Daphnia in the epilimnion and hypolimnion by transferring water from these layers into flow-through systems where we raised Daphnia and assessed their juvenile growth rate as a measure of fitness. The fitness distribution was correlated with the vertical distribution. The vertical distribution most likely reflected the proportions of time Daphnia allocated to dwelling in the two vertical habitats.     

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.     

The possible origin ofDaphnia cucullata procurva Poppe 1887 in the lakes of the Pomeranian Lakeland by hybridization in the past

Besides the genusBosmina, the genusDaphnia offers considerable taxonomic difficulties due to the existence of a very great number of different morphotypes and their intergrades. Especially the morphotypes clustering aroundDaphnia galeata Sars, 1864,D. hyalina Leydig, 1860, andD. cucullata Sars, 1862 offer a bewildering picture. In the last years an attempt has been made to understand this phenomenon mainly as the result of hybridization and colonization of new biotopes by populations with hybridogenetic polymorphism and genotypic diversity (founder effect). Now it will be suggested on a morphological base thatD. cucullata procurva Poppe, 1887, a unique and aberrant formtype restricted to few lakes along the river Brda in Poland, is probably an old and stabilized hybrid originating from interspecific hybridization betweenD. galeata andD. cucullata.