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.     

Evaluation of a fluorescent microparticle technique for measuring filtering rates of Daphnia

The paper demonstrates the applicability of fluorescent microparticles (FMP) for the determination of cladoceran feeding rates. For this purpose we examined the relation between filtering rate, ingestion rate, gut passage time and body length of Daphnia pulex at different food concentrations in laboratory experiments, and diurnal changes of filtering rates of Daphnia longispina in in situ experiments, using FMPs as tracer particles. Good agreement of our results with published data based on cell counts and radiotracer methods proves the suitability of the FMP-method. Potentials and limitations of the method are discussed.     

Impact of an introduced Crustacean on the trophic webs of Mediterranean wetlands

Based on a review and our own data, we present an overview of the ecological impacts on the trophic web of Mediterranean wetlands by an introduced Decapod Crustacean, the red swamp crayfish (Procambarus clarkii). P. clarkii lacks efficient dispersal mechanisms but is very well adapted to the ecological conditions of Mediterranean wetlands (fluctuating hydroperiods with regular intervals of drought). As an opportunistic, omnivorous species, which adapts its ecology and life history characteristics, such as timing and size at reproduction to changing environmental conditions, it became readily established in most of the Mediterranean wetland environments. High reproductive output, short development time and a flexible feeding strategy are responsible for its success as an invader. Like most crayfish, it occupies a keystone position in the trophic web of the invaded system and interacts strongly with various trophic levels. It efficiently grazes on macrophytes and is one of the main factors, besides the impact of flamingos, cattle and introduced fish, of the change of many water bodies from a macrophyte dominated, clear water equilibrium to a phytoplankton driven turbid water balance. Juveniles feed on protein rich animal food with the corresponding impact on the macroinvertebrate community in competition with other crayfish or fish species. At the same time, it serves as a prey for mammals, birds and fish. Due to its predatory and grazing activity, it efficiently canalises energy pathways reducing food web complexity and structure. Feeding also on detritus it opens, especially in marshlands, the detritic food chain to higher trophic levels which results in an increase of crayfish predators. As a vector of diseases, it has a severe impact on the preservation and reintroduction of native crayfish. P. clarkii accumulates heavy metals and other pollutants in its organs and body tissues and transmits them to higher trophic levels. Due to the long history of its presence, the complex interactions it established within the invaded ecosystems and the socio-economic benefits it provides to humans, prevention and control seem the most promising management measures to reduce the negative impact of this crayfish species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Suspension feeding in Bithynia tentaculata (Prosobranchia,Bithyniidae), as affected by body size,food and temperature

The suspension feeding of Bithynia tentaculata was tested in laboratory experiments. The animals were fed in 1-1 aerated glass beakers, and filtration rates were calculated from changes in cell concentrations during the 6-h experiment. Temperature influenced the filtering rate, with minimum values of 5ml · ind^–1 · h^–1 at 5° C and maxima of 17.2 ml · ind^–1 · h^–1 at 18° C. Three food species of different size, motility and cell surface characteristics (Chlamydomonas reinhardii, Chlorella vulgaris and Chlorogonium elongatum) did not affect filtration rates. Suspension feeding increased with increasing food concentrations up to 12 nl · ml^–1, above which feeding rate was kept constant by lowering the filtering rates. Even the smallest animals tested (<4 mm body length) were found to be feeding on suspended food at a rate of 2.7 ml · ind^–1 · h^–1, and increasing rates up to 8.4 ml were found in the 6–7 mm size class. All size classes of Bithynia showed a circannual fluctuation of their filtration rates. The ecological consequences of Bithynia's ability to switch between two feeding modes, grazing and suspension feeding, are discussed.     

The impact of food quantity on UV-B tolerance and recovery from UV-B damage in Daphnia pulex

UV-B (290 nm) tolerance of Daphnia pulex, conditioned to four different food levels (Chlorophyta), was tested under standardized conditions with an artificial radiation source. Parameters measured were survival, percentage of egg bearing Daphnia and the number of juveniles produced after irradiation. UV-B tolerance of Daphnia pulex was found to be significantly improved with increasing food concentrations at all three parameters. The impact of the four different food concentrations on the photoreactivation system was tested with simultanous UV-B and white-light irradiation of Daphnia. Survival rate improved significantly with increasing food levels compared to solely UV-B irradiation. Photoreactivation had no effect on the reproductive parameters.     

Phaeocystis and its interaction with viruses

Over the years, viruses have been shown to be mortality agents for a wide range of phytoplankton species, including species within the genus Phaeocystis (Prymnesiophyceae). With its polymorphic life cycle, its worldwide distribution, and the capacity of several of the Phaeocystis species to form dense blooms, this genus is a key player for our understanding of biogeochemical cycling of elements. This paper provides an overview of what is know to date about the ecological role of viruses in regulating Phaeocystis population dynamics. It explores which variables affect the algal host–virus interactions, and examines the impact of virally induced cell lysis of Phaeocystis on the function and structure of the pelagic food web as well as on the flow of organic carbon and nutrients.     

The perfect invader: a parthenogenic crayfish poses a new threat to Madagascar’s freshwater biodiversity

In 2007 an unusual crayfish found in food markets in the capital of Madagascar was preliminarily identified as Procambarus ‘Marmorkrebs’: a new world taxa and the only decapod known to reproduce by parthenogenesis. We present information on the identity, distribution and ecology of this recent invader and attempt to evaluate the threat it poses to Madagascar’s biodiversity and to livelihoods. The species appears to be currently limited to the area close to Antananarivo, but is being sold alive on major transport routes. We present molecular evidence of its taxonomic relationships and confirm that the Procambarus present in Madagascar is indeed the parthenogenic taxa. We investigate its reproductive ecology and find Procambarus ‘Marmorkrebs’ to have an extremely high fecundity; more than six times that of the native crayfish Astacoides. The limited evidence we have suggests that this species poses a serious threat to freshwater biodiversity and that it is likely to damage human livelihoods (through its impact on fishing and possibly rice agriculture). More research is urgently needed but in the meantime action is needed to reduce the rate of spread before it is too late.     

Xenopus laevis larvae (Amphibia,Anura) as model suspension feeders

Larvae of the South African clawed frog, Xenopus laevis (Daudin), are efficient, obligate suspension feeders. We examine the relationship between the ambient particle concentration offered these larvae as food and their filtering, ingestion, and buccal pumping rates. We demonstrate that: (i) the larvae can sense and respond to a broad range of particle concentrations, down to 0.2 mg 1^–1 (dry weight); (ii) their metabolic needs theoretically can be met by particle concentrations as low as 5 mg 1^–1; and (iii) their patterns of regulation of filtering and ingestion fit predictions from certain models used to describe zooplankton feeding dynamics. Two such models are discussed: the modified Monod (Michaelis-Menten) model, with a lower threshold below which the tadpoles do not feed, and an energy optimization model. Both the models and the observed behavior of the tadpoles allow for stability of populations of food organisms. Tadpole feeding dynamics apparently are compatible with both the predictions and assumptions of these models, suggesting similar regulation of feeding by tadpoles and zooplankton. However, the size, morphology, and behavior of X. laevis larvae make their feeding regulation uniquely accessible to direct observation.Contribution No. 223, Center for Great Lakes Studies, University of Wisconsin-Milwaukee, Milwaukee, Wisconsin.Contribution No. 223, Center for Great Lakes Studies, University of Wisconsin-Milwaukee, Milwaukee, Wisconsin.     

The effects of food and temperature regimes on life-history responses to fish kairomones in Daphnia hyalina × galeata*

Life-history responses to two concentrations of fish released info-chemicals at two temperature and food regimes were investigated for one clone of Daphnia hyalina × galeata. The presence of fish kairomones had a negative impact on size at maturity, carbon allocation to individual eggs and size of neonates in all treatments. Food concentration and temperature had positive effects on size of adult stages, independent of kairomone treatment. However, kairomone treatment were not found to interact with food or temperature. Age at maturity was positively influenced by increased temperature and food concentrations, whereas no direct kairomone effects were detected for this trait. Clutch size was not directly influenced by kairomone treatment, whereas both food concentration and temperature had strong, positive effects.     

Ecological aspects,explosive range extension and impact of a mass invader,Corophium curvispinum Sars, 1895 (Crustacea: Amphipoda), in the Lower Rhine (The Netherlands)

A few years after it invaded, the amphipod Corophium curvispinum Sars appeared to be the most numerous macroinvertebrate species in the River Rhine. From 1987 to 1991 the densities of this species on the stones of groins in the Lower Rhine at a depth of 0.5 m increased from 2 to 200000 specimens per m². In the Lower Rhine and its branches the densities of C. curvispinum increased with increasing current velocities and with increasing water depths. So far, a maximum population density of 750000 specimens per m² has been found in the Lower Rhine, which is many times the densities recorded elsewhere. Population parameters, densities and distribution of C. curvispinum were studied in the Lower Rhine and its branches, using artificial substrates and sampling stones from groins. The success of this immigrant is related to its competitive strategy, which shows several aspects of a r-strategy. In addition, the heavily eutrophicated Lower Rhine provides abundant food (phytoplankton, suspended organic matter) for this opportunistic filter-feeder. The increased salinity and water temperatures in the Lower Rhine resulting from industrial discharges have contributed to the current success of this southern species originating in brackish waters. The very high densities of C. curvispinum might have an enormous impact on the river ecosystem by changing food webs.     

Cross-generic Food Sharing in Tamarins

Adult-infant food-sharing behavior is a major component of the infant care strategies of callitrichids (marmosets and tamarins). It is particularly well-developed in cotton-top tamarins (Saguinus Oedipus) and lion tamarins (Leontopithecus spp), which show frequent adult-initiated food offering, as well as sharing of food in response to begging by infants. This report documents a case of cross-generic food sharing, in which a male golden-headed lion tamarin (Leontopithecus chrysomelas) shared food with an infant cotton-top tamarin. The lion tamarin provided more food to the infant than its mother did. This emphasizes the importance of this behavior in the reproductive strategies of the communally-rearing Callitrichidae and raises questions about mechanisms that underly it.     

Invasion of an exotic forb impacts reproductive success and site fidelity of a migratory songbird

Although exotic plant invasions threaten natural systems worldwide, we know little about the specific ecological impacts of invaders, including the magnitude of effects and underlying mechanisms. Exotic plants are likely to impact higher trophic levels when they overrun native plant communities, affecting habitat quality for breeding songbirds by altering food availability and/or nest predation levels. We studied chipping sparrows (Spizella passerina) breeding in savannas that were either dominated by native vegetation or invaded by spotted knapweed (Centaurea maculosa), an exotic forb that substantially reduces diversity and abundance of native herbaceous plant species. Chipping sparrows primarily nest in trees but forage on the ground, consuming seeds and arthropods. We found that predation rates did not differ between nests at knapweed and native sites. However, initiation of first nests was delayed at knapweed versus native sites, an effect frequently associated with low food availability. Our seasonal fecundity model indicated that breeding delays could translate to diminished fecundity, including dramatic declines in the incidence of double brooding. Site fidelity of breeding adults was also substantially reduced in knapweed compared to native habitats, as measured by return rates and shifts in territory locations between years. Declines in reproductive success and site fidelity were greater for yearling versus older birds, and knapweed invasion appeared to exacerbate differences between age classes. In addition, grasshoppers, which represent an important prey resource, were substantially reduced in knapweed versus native habitats. Our results strongly suggest that knapweed invasion can impact chipping sparrow populations by reducing food availability. Food chain effects may be an important mechanism by which strong plant invaders impact songbirds and other consumers.     

Moulting diving ducks and their food supply

Numbers of moulting diving ducks at Myvatn, north Iceland, were monitored over a period of 25 years; aquatic insects (Chironomidae and Simuliidae), a major food resource, were monitored with window traps for 23 years. The response of the duck populations to changes in the food situation during the moulting period varied between species. Histrionicus histrionicus and Melanitta nigra invariably go to the sea to moult. Three species (Aythya fuligula, Clangula hyemalis and Mergus serrator) have shown long term variation in moult migration habits. Aythya marila and Bucephala islandica stay for moulting, their numbers supplemented by birds from elsewhere. A. marila is a generalist feeder and therefore at low risk of starving, the strategy of Bucephala islandica is to move within the water system where the food situation tends to alternate between the lake and the river. Numbers of Bucephala islandica males moulting on Lake Myvatn were strongly and positively correlated with chironomid numbers and those moulting on the river Laxá with Simulium vittatum, the main food resource there. We did not find convincing evidence that numbers of moulting A. marila and C. hyemalis responded to variation in the food supply during the study period. This applied also to A. fuligula in the South Basin of Myvatn, but numbers in the North Basin were positively associated with chironomid numbers. M. serrator moulting on Myvatn showed negative correlations with the chironomids, perhaps reflecting a negative association between chironomids and its main food, Gasterosteus aculeatus. Apart from safety considerations for a flightless bird, the choice of a moulting site is apparently influenced by the local food conditions on one hand and by the opportunities and risks involved in migrating to distant moulting sites with an unknown food situation on the other hand.     

Zooplankton feeding on algae and bacteria under ice in Lake Druzhby,East Antarctica

The feeding of the cladoceran Daphniopsis studeri on algae and bacteria was investigated under ice in an ultra-oligotrophic Antarctic lake from late autumn (May) to early spring (October) in 2004. D. studeri fed on both algae and bacteria with estimated filtering rates of 0.048 and 0.061 l ind⁻¹ day⁻¹), respectively. Algae seemed to be the major food resource for the D. studeri population, however at times of low algal densities the bacterioplankton represented an important alternative food resource. The D. studeri grazing impact on the algal and bacterial standing stock was in general low (0.6–4.6% removed per day), but during the winter period this organism can remove up to 34% of the bacterial production (BP). At times D. studeri grazing can temporarily have a significant impact on the BP rates, though their impact was relatively low when compared to viral-induced bacterial mortality in the lake.     

Morphological adaptations in filtering screens of Daphnia galeata to food quantity and food quality

We reared clones of the waterflea Daphnia galeata, a common grazer in many types of lakes, under several food regimes to study adaptations to feeding conditions in filter screen morphology and life history. As food regimes, we used low and high concentrations of the green alga Scenedesmus, a high concentration of the filamentous cyanobacterium Oscillatoria, and a mixture of Scenedesmus and clay. Natural seston from Lake IJsselmeer was also tested. Clones from two contrasting habitats (mesotrophic versus hypertrophic lake) did not differ in either population growth rate, r, or filtering structures. However, all the clones exhibited phenotypic plasticity when reared under low and high food concentrations. In relation to their body length, daphnids grew larger filter screens with longer setae in response to low food concentrations, thus enhancing filtering efficiency. Supplementing the low food concentration with inert clay particles had no effect on either the growth performance or filter screen morphology. In both cases, the growth and reproduction were low and filtering areas large. Filter screens were of intermediate size when daphnids were supplied with food supporting intermediate growth and reproduction. From these laboratory results, we concluded that the nutritional status of Daphnia is a more important cue for the phenotypic response of the filter screen morphology than particle concentration.      

Type-3 functional response in limnetic suspension-feeders,as demonstrated by in situ grazing rates

Field-measured grazing rates (ml/animal/d) of cladocerans (mostly daphniids) and diaptomids were assembled from various published studies and plotted as a function of corresponding phytoplankton concentration (μg l⁻¹ f.w.). Filtering rates of both zooplankton groups initially increased with seston concentration until maximal grazing rates were observed at approximately 4 × 10² and 1 × 10² μg l⁻¹ for cladocerans and copepods, respectively; at higher algal concentrations, filtering rates of both declined as a function of food concentration. The shape of these curves are most consistent with Holling's (1966) Type 3 functional response. We found little support for the Type 3 functional response in published laboratory studies of Daphnia; most investigators report either a Type 1 or Type 2 response. The one study in which the Type 3 response was observed involved experiments where animals were acclimated at low food concentrations for 24 h, whereas those studies associated with response Types 1 or 2 had acclimation periods of only 1 to 3 h. We therefore assembled relevant data from the literature to examine the effect of acclimation period on the feeding rates of Daphnia at low food concentrations. In the absence of any acclimation, animals filtered at extremely low rates. After 2 h of acclimation, however, filtering rates increased 4 to 5-fold but declined again with longer durations; after > 70 h of pre-conditioning, filtering rates were almost as low as they had been with no acclimation. We also found little support for the Type 3 functional response in published studies of copepods. The only study associated with a Type 3 response involved a marine copepod that had been subjected to a starvation period of 48 h; however, an analysis of the effects of acclimation period did not yield conclusive evidence that filtering rates of freshwater copepods (Diaptomus and Eudiaptomus) decrease significantly with acclimation duration. The low filtering rates associated with long acclimation periods in laboratory experiments appears to be a direct result of animals becoming emaciated from prolonged exposure to low food concentrations, a situation which renders them incapable of high filtering rates. This may explain the Type 3 functional response for field cladocerans, since zooplankton in food-limiting situations are constantly exposed to low food concentrations, and would therefore have low body carbon and consequently less energy to filter-feed. We cannot, however, use this to explain the Type 3 response for field diaptomids, since copepods in the laboratory did not appear to lose body carbon even after 72 h of feeding at very low food levels, and there was inconclusive evidence that either Diaptomus or Eudiaptomus decrease their filtering rates with acclimation period. Although Incipient Limiting Concentrations (ILC) for Daphnia ranged from 1 to 8.5 × 10³ μg 1⁻¹, more than half of these fell between 1 and 3 × 10³ μg l⁻¹, bracketing the value of 2.7 × 10² μg l⁻¹ for field cladocerans. There was, however, a great deal of variation in reported maximum ingestion rates (MIR), maximum filtering rates (MFR) and ILC values for Daphnia magna. ILC values from the few laboratory studies of freshwater copepods ranged between 0.5 to 2.8 × 10³ μg 1⁻¹, and was higher than the ILC value of approximately 0.2 × 10³ μg l⁻¹ calculated for field populations of D. minutus. Generally, there was considerable agreement among laboratory studies regarding the shape of grazing-rate and ingestion-rate curves when data were converted to similar units and presented on standardized scales.     

Foraging behavior by <Emphasis Type="Italic">Daphnia</Emphasis> in stoichiometric gradients of food quality

Mismatches in the elemental composition of herbivores and their resources can impact herbivore growth and reproduction. In aquatic systems, the ratio of elements, such as C, P, and N, is used to characterize the food quality of algal prey. For example, large increases in the C:P ratio of edible algae can decrease rates of growth and reproduction in Daphnia. Current theory emphasizes that Daphnia utilize only assimilation and respiration processes to maintain an optimal elemental composition, yet studies of terrestrial herbivores implicate behavioral processes in coping with local variation in food quality. We tested the ability of juvenile and adult Daphnia to locate regions of high-quality food within a spatial gradient of algal prey differing in C:P ratio, while holding food density constant over space. Both juveniles and adults demonstrated similar behavior by quickly locating (i.e., <10 min) the region of high food quality. Foraging paths were centred on regions of high food quality and these differed significantly from paths of individuals exposed to a homogeneous environment of both food density and food quality. Ingestion rate experiments on algal prey of differing stoichiometric ratio show that individuals can adjust their intake rate over fast behavioral time-scales, and we use these data to examine how individuals choose foraging locations when presented with a spatial gradient that trades off food quality and food quantity. Daphnia reared under low food quality conditions chose to forage in regions of high food quality even though they could attain the same C ingestion rate elsewhere along a spatial gradient. We argue that these aspects of foraging behavior by Daphnia have important implications for how these herbivores manage their elemental composition and our understanding of the dynamics of these herbivore–plant systems in lakes and ponds where spatial variation in food quality is present. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Selective grazing of Temora longicornis in different stages of a Phaeocystis globosa bloom—a mesocosm study

Selective grazing of a calanoid copepod Temora longicornis was measured during different stages of a Phaeocystis globosa bloom, in order to reveal (1) if T. longicornis feeds on single cells and/or colonies of P. globosa in the presence of alternative food sources, (2) if copepod food selection changes during the initiation, maintenance, collapse and decay of a P. globosa bloom and (3) if P. globosa dominated food assemblage provides a good diet for copepod egg production. Our results show low but constant feeding on small colonies of P. globosa, irrespective of the type or concentration of alternative food sources. In contrast, feeding on single cells was never significant, and the total contribution of P. globosa to carbon ingestion of T. longicornis was minor. T. longicornis fed most actively on the decaying colonies, whereas during the peak of the bloom copepods selected against P. globosa. Mostly, T. longicornis fed unselectively on different food particles: before the bloom, the major part of the diet consisted of diatoms, whereas during and after the bloom copepod diet was dominated by dinoflagellates and ciliates. Egg production was highest during the decay of the bloom, coinciding with highest proportional ingestion of heterotrophic organisms, but was not seriously reduced even during the peak of the bloom. We conclude that P. globosa blooms should not threaten survival of copepod populations, but the population recruitment may depend on the type (and concentration) of the dominant heterotrophs present during the blooms. Due to relatively unselective grazing, the impact of T. longicornis to the initiation of a Phaeocystis bloom is considered small, although grazing on decaying colonies may contribute to the faster termination of a bloom.     

Learning in an invasive and a native predatory crab

The invasion of the green crab Carcinus maenas in the northeastern U.S. and its competition with the native blue crab Callinectes sapidus and other native crustaceans has been well-documented and researched. Various reasons for the invader’s success against native crabs have been examined (juvenile predation, food source flexibility, etc.), but another possibility is a difference in the learning ability of invasive versus native crab species. In this study, the learning ability of C. maenas and C. sapidus was tested by their increased speed in locating hidden food over successive days. The data suggest that C. maenas possesses a learning ability significantly greater than that of C. sapidus, which may partially contribute to its success.     

Plasticity in filtering screens of Daphnia cucullata×galeata hybrids and parental species at two food concentrations

The coexistence of Daphnia cucullata × galeata hybrids with the parental species D. galeata and D. cucullata was investigated by measuring areas and mesh sizes of filtering structures of these herbivorous zooplankton taxa cultivated at low and high food concentrations. The clearance rates and somatic growth rates were also determined. When reared at low food concentration, all taxa had larger filtering areas. Larger filtering areas also resulted in higher clearance rates. Differences between taxa in both filtering area and clearance rate were caused mainly by interspecific size differences. Hybrids had the largest absolute mesh sizes, and the parental species had smaller mesh sizes. Hybrids also showed heterosis in somatic growth rate at high food concentration. The observed taxon-specific differences in mesh size and somatic growth rate contribute to resource partitioning between the taxa and thus to their successful coexistence in lakes. Received: 1 March 1999 / Accepted: 25 May 1999