Predation and priority effects in experimental zooplankton communities

During colonization of new habitats, the sequence of arrival among species is in many cases determined by chance. Priority effects imply that differences in arrival time may lead to long-lasting differences in species dominance. To evaluate the importance of priority effects, we performed a community assembly experiment, manipulating the inoculation order of three large cladoceran zooplankton species. The inoculation treatments were crossed with a predation treatment to test whether the presence of a predator (larvae of the phantom midge Chaoborus ) results in changes in the strength of species sorting and priority effects. Our results clearly demonstrate priority effects, but also that the presence of a predator impacts both community assembly and the strength of priority effects. In the predation-free treatments, communities were dominated by either Daphnia magna or Daphnia obtusa , depending on the species arrival sequence. Whenever D. obtusa was inoculated after D. magna , the species displayed negative growth. In the presence of Chaoborus predation, the communities were generally dominated by the third species, Simocephalus vetulus . Here, the growth of D. magna was negative when the species was inoculated as second or third. Overall, our results underscore the importance of both priority effects and species sorting during community assembly.     

Patchiness and composition of coral reef demersal zooplankton

Zooplankton samples were collected weekly for a full year withdemersal traps on a coral reef off the west coast of Barbados.There was a marked temporal variability in weekly catches bothin terms of abundance and biomass. Patchiness occurred at allsampling frequencies from 2 to 26 weeks, but spectral analysisindicated a variance shift at a frequency of 8–10 weeksAggregations of the two most abundant taxa, the copepoditesand the microzooplankton, occurred at 8–12 week intervalsand significant differences in abundance and biomass were foundbetween mean bimonthly zooplankton catches Lagged cross-correlationsat 7 and 11 weeks between chlorophyll and microzooplankton andcopepodites suggest that aggregations are connected to cyclesof primary production. There was a negative correlation betweenzooplankton abundance and surface water salinity in 8 of 16taxa Copepods were the most abundant taxon overall. Microzooplanktonand copepodites comprised 96% of the abundance and 66% of thebiomass Decreases in taxonomic richness and in diversity wereassociated with patchiness of small-sized copepodites and microzooplankton,suggesting that composition was altered and stability temporarilylessened during peaks of abundance     

Influence of overwinteringDaphnia on spring zooplankton communities: An experimental study

To evaluate the influence of overwintering individuals of zooplankton on spring zooplankton communities, the dynamics of zooplankton communities with or without overwintering individuals were observed in experimental ponds from fall to spring. An insecticide, carbaryl, was used to regulate the overwintering individuals. In ponds which received insecticide applications in November or January, all cladoceran and rotiferan species were eliminated by the treatments and did not reappear until late March or early April, even when the chemical disappeared rapidly. The low water temperature may delayed the establishment of the populations from resting eggs. In these ponds, populations of various cladoceran and rotiferan species, which seemed to be originated from resting eggs, were built up in the spring. In control ponds,Daphnia ambigua orD. longispina overwintered as juveniles and adults and established a large spring population earlier than other cladocerans and rotifers overwintering as resting eggs. The latter zooplankters did not increase in the spring probably because their growth was suppressed by the precedingDaphnia species through competition. In nature, even if the number of overwintering individuals is small, they may have a potential to build up a large population earlier than the individuals hatching from resting eggs. As a result, the species which have overwintered as individuals seem to predominate in the spring and have a large influence on the spring zooplankton community.     

Complex interactions in the control of coral zonation on a Caribbean reef flat

Summary This study uses short-term assays and long-term transplant experiments to document the potential importance of fish predation and herbivory to the distribution and abundance of reef-building corals in a Caribbean back-reef system. Experimental manipulations of fish access reveal that the zonal patterns of the two reef-building corals Porites astreoides and P. porites f. furcata, dominant on shallow back-reef habitats, are strongly associated with the feeding intensity of parrotfishes. Differential palatability of the two corals to parrotfishes, the proximity of protective cover for large grazers and the availability of small refugia to harbor a cryptic grazer fauna are suggested as major features contributing to the observed patterns. A model predicting the interactions of various algivore/corallivore guilds on the relative dominance of Porites and algal populations is presented.     

Vertical zonation of coral reef fishes in the Sudanese Red Sea

The vertical zonation of 38 species of fish inhabiting coral reefs in the Sudanese Red Sea is described. A marked degree of vertical zonation of the fishes is shown to exist. The usefulness of a depth oriented survey method for studying reef fish is discussed in relation to these results.General relationships between vertical zonation and species diversity are noted.     

An analysis of some effects of light and soil temperature on clubroot disease

In a glasshouse study, 54 batches of cabbage seedlings were each grown for 6 wk in clubroot inoculated soil. The batches were sown at weekly intervals during the experiment which lasted 60 wk. Soil temperature and light were monitored throughout the study while other potential variables were maintained as constant as possible. Records of clubroot incidence, severity and plant weights were kept and causal relationships for disease severity and percentage infected plants sought among the light and temperature data by regressions. It was found that clubroot severity was influenced most by the light level in the second and third weeks, while the percentage of diseased plants was most influenced by temperature, that in the second week probably being the most important. The optimum light value in the second and third weeks for expression of maximum clubroot severity was c. 600 Wh m^-2 day^-1 and a mean daily temperature of not less than c. 19-5 ^oC was required to give close to 100% infection. Possible reasons for the findings are discussed.     

An enclosure for experimental field studies with fish and zooplankton communities

A wooden framed enclosure screened by plastic sheeting is described. This was sited along the bank to alleviate circulation problems and to allow for easy construction, surveillance and maintenance and sampling. An aeration system was incorporated. The suitability of the enclosure for use in field experiments on fish and zooplankton was tested by using 10 enclosures to investigate the effect of fish on the density of zooplankton. Fish survival varied from 15.5 to 47% and was similar to that for the pond. Statistically significant inferences were obtained using a one way ANOVA which showed that fish predation did have an effect on the density of some species of zooplankton. Cluster analysis was used to illustrate the closeness of relationship of the zooplankton communities in the enclosures with and without fish. Problems associated with the use of enclosures in experimental studies are discussed.     

The effects of clove oil on coral: An experimental evaluation using Pocillopora damicornis (Linnaeus)

Clove oil solution (10% clove oil, 90% ethanol) is an anaesthetic that is widely used to catch demersal fish on coral reefs. This study assessed the effects of clove oil solution on colonies of Pocillopora damicornis, a cosmopolitan reef coral. In the laboratory, low concentrations (0.5 ppt) of clove oil solution had no effect on coral colour or photosynthetic efficiency, irrespective of exposure time (1-60 min). Corals treated with high concentrations (50 ppt) of clove oil solution died immediately, including those that were exposed briefly (1 min). Intermediate concentrations (5 ppt) of clove oil solution produced variable results: a 1 min exposure had no effect, a 10 min exposure caused bleaching and reduced photosynthetic efficiency, and a 60 min exposure caused total mortality. To validate these observations, clove oil solution was applied to corals in situ. Sixty-three days after application, corals treated with 10 ml of clove oil solution appeared to be unaffected. It was concluded that (1) limited amounts of clove oil solution are unlikely to harm this coral, and (2) clove oil solution may represent an ‘eco-friendly’ alternative to cyanide for use in the live reef-fish trade.     

Near-bottom depletion of zooplankton over a coral reef II: relationships with zooplankton swimming ability

Intense predation by corals and associated fauna can generate vertical gradients of their zooplankton prey. The goal of this study was to characterize the small-scale distribution of zooplankton above the coral reef of Eilat, Israel. Four vertical arrays, each consisting five underwater pumps attached 0.5–3 m apart on a taut mooring, were deployed for 10 days at two sites over the reef slope. A distinct layer of depleted zooplankton was repeatedly found in the 1.5 m high benthic boundary layer. The gradient was sharpest for strong swimmers (copepods and polychaeta), intermediate for weak swimmers (nauplii, mollusks and appendicularia), and lacking for passive taxa (eggs and foraminifera). Spatio-temporal changes in the abundance of copepods and polychaetes were highly correlated, more so in the water aloft (Pearson r>0.9) than near the bottom (r>0.8). The spatio-temporal correlations between weak swimmers and between passive taxa were much weaker (r<0.8 and r<0.4, respectively). Flow and shear stress did not affect the distribution patterns. The correspondence between the zooplankton distribution and their swimming ability indicates that bottom avoidance contribute to the formation of depleted layer over the reef.     

Preliminary results of an experimental study of the effects of mysid predation on estuarine zooplankton community structure

In a shallow estuarine system near Beaufort, North Carolina, a period of high winter abundance of the mysidsMysidopsis bigelowi andNeomysis americana was associated with a change in zooplankton species composition, from dominance byAcartia tonsa to dominance byCentropages spp. andSaphirella sp. Both mysids feed onA. tonsa at higher rates than the other copepods. Experiments were carried out in 600–1 000 liter enclosures, in which the initial mysid density was manipulated and the effects on the enclosed copepod community were monitored. Mysid predation had a significant effect on copepod densities. The effects of mysid predation on species composition appeared to depend on the relationship between their prey preferences and the dominant copepod species present in the communities. Under conditions favoring dominance byA. tonsa, the preferred prey species, the results suggested that mysid predation may reduce dominance and increase diversity. But when the less preferredCentropages was dominant, mysid predation had no effect on species composition.     

Shedding new light on scleractinian coral recruitment

Studies of the distribution of scleractinian coral recruits within stacks of settlement plates at Orpheus Island and Lizard Island revealed that corals are recruited almost exclusively on the lower surfaces of the settlement plates. The absence of recruitment on the upper surfaces of the plates is attributed to the observed high levels of sediment on these surfaces. All hermatypic coral spat were found near the edge of the settlement plates, whereas ahermatypic corals were located towards the center. Growth and survivorship of hermatypic corals were dependent on the position of attachment, decreasing with distance from the edges of the plates. Survivorship of ahermatypic corals was uniformly high. Light levels on the lower surfaces of the plates within the stacks were shown to decay asymptotically as a function of distance from the plate edge. There was a highly significant non-linear relationship between relative light intensity on the settlement plates and the positions of coral spat. In the absence of evidence for selective hydrodynamic deposition of spat, these results suggest that light intensity is the most important factor in determining the position of attachment of coral spat on the underside of these plates.     

An experimental analysis of species sorting and mass effects in freshwater bacterioplankton

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Effects of light dynamics on coral spawning synchrony

Synchrony of spawning in many hermatypic corals, typically a few nights after the full moon, is putatively dependent on solar and lunar light cycles in conjunction with other possible cues such as tides and temperature. We analyze here the contributions of separate components of light dynamics, because the effects of twilight and lunar skylight on coral spawning synchrony have previously been conflated and the alternative hypothesis that these components have differential contributions as proximate cues has not been tested. Moonlight-dependent changes in spectra during twilight, rates of decreasing twilight intensities, and changes in lunar photoperiod were experimentally decoupled using programmed light-emitting diodes and compared for their separate effects on spawning synchrony in Acropora humilis. Effects on synchrony under the control of synthetic lunar cues were greatest in response to changes in lunar photoperiod; changes in light intensities and spectra had lesser influence. No significant differences among treatment responses were found at the circa-diel time scale. We conclude that spawning synchrony on a particular lunar night and specific time of night is a threshold response to differential periods of darkness after twilight that is primarily influenced by lunar photoperiod and secondarily by discrete optical components of early nocturnal illumination.     

Effect of light and predator abundance on the habitat choice of plant-attached zooplankton

1. The diurnal variations in the habitat choice of the periodically plant-attached cladoceran Sida crystallina together with light environment and predator abundance were studied.
2. The density of S. crystallina attached to floating leaves of Nuphar lutea increased between 18:00 and 20:00 hours, when light intensity underneath the leaves was temporarily increased, and decreased again when light intensity declined. A proportion of S. crystallina remained in the swimming mode underneath the leaves even during daylight, indicating that the water column sheltered by the leaves is safer than the open water.
3. In the water adjacent to the leaves, the density of S. crystallina increased steeply in the dark. The increase was not accompanied by a decrease in S. crystallina attached to plant leaves, indicating that the nocturnal increment in the open water density of S. crystallina was due to migration from daytime refuges other than floating leaves.
4. Sida crystallina was most intensively consumed by perch ( Perca fluviatilis ). Predation threat by fish had weaker effects on the density of S. crystallina attached to plant leaves than on cladocerans in the adjacent water. Cladocerans underneath floating plant leaves, whether attached or not, are probably less vulnerable to fish predation than those outside the leaf cover.
5. The results suggested that light intensity is the proximate factor regulating the attachment of zooplankton to the lower surfaces of floating macrophyte leaves. Light intensity has a positive effect on the density of S. crystallina attached to the floating leaves and a negative effect on density in the water. Predation threat by fish has a strong effect on the migration of zooplankters into the open water habitat.     

Production and supply of larvae as determinants of zonation in a brooding tropical coral

Production and settlement of planktonic larvae of the coral Favia fragum (Esper) were studied. The species is restricted to shallow back- and fore-reef habitats throughout the Caribbean Sea. Adults are in their greatest abundance on the reef-crest and shallow reef slope (<3 m) at Tague Bay, St. Croix, in the US Virgin Islands. Because F. fragum broods larvae that are capable of immediate settlement, this distribution pattern may be due to variation in fecundity among depths. Corals were collected from shallow (1.0 m) and deep depths (10-13 m) and cultured in individual containers exposed to shaded ambient light. Corals from shallow depths had greater fecundity (polyp⁻¹ lunar cycle⁻¹) and were larger than deep corals. To test the hypothesis that fecundity was related to successful fertilization, corals were kept in different densities in an area with sea-grass, where there were no natural adults. Production of larvae 6 months later was not affected by density of adults, possibly due to self-fertilization. Larval choice of habitat was also examined. In the laboratory, twice as many larvae settled on coral rubble fragments collected from depths where adults were common (1.5 and 3 m) than on those from depths where adults were rare (10 m). Larval supply may establish the vertical distribution of adults on St. Croix.     

Direct and indirect impacts of predation by fish on the zooplankton community: an experimental analysis using tanks

The impact of Pseudorasbora parva, a common zooplanktivorous fish species in Japan, on a zooplankton community was analyzed in experimental tanks, half of which were stocked with the fish. Different zooplankton species showed different responses to the introduction of the fish. In the presence of the fish, the populations of the large cladoceran Ceriodaphnia and the predatory copepod Mesocyclops were reduced, but the population of the herbivorous copepod Eodiaptomus and the small cladocerans Bosmina fatalis and Bosminopsis deitersi increased relative to the controls. The increase of Mesocyclops seen in the control tanks might have suppressed the populations of the small cladocerans, which are vulnerable to invertebrate predation. The results suggest that the population densities of the large prey items preferred by the fish, Ceriodaphnia and Mesocyclops, were controlled directly by fish predation, but the population densities of the smaller and less preferred zooplankton were controlled indirectly through the food-web cascade.     

The zonation patterns of Caribbean coral reefs as controlled by wave and light energy input,bathymetric setting and reef morphology: computer simulation experiments

The computer model COREEF was used to simulate variations in the zonation patterns of Caribbean reefs in relation to parameters that affect the magnitude and distribution of wave and light energy. We first developed a simulated standard reef by exposing a simplified profile of the reef at Discovery Bay, Jamaica, to the known wave and light energy conditions to establish a reference coralgal and sedimentological zonation pattern. We then varied 13 parameters related to the wave and light energy input, bathymetric setting, and gross morphology of this reef to determine the effects of each parameter on the zonation pattern. Analysis of the simulation results indicates that submerging the reef or altering the wave or light energy input to the reef produces the greatest modifications of the zonation pattern. Morphological structures that alter a reef's horizontal dimensions only minimally affect the zonation pattern, but those structures that alter a reef's vertical dimensions-particularly steep-sided, wave reflecting structures-can significantly modify the zonation of the structure itself and that of more leeward areas. The more seaward the location of a morphological structure, the more profoundly it can affect the overall reef zonation. If waves break at the reef crest, wave energy conditions in the back reef are greatly reduced and the bottom consists of lower wave energy zones than those found at the same depths in the fore reef. If waves do not break at the crest, the back reef is subjected to almost the same wave conditions that exist in the fore reef, and the zones tend to be similar. The zonation patterns of some existing reefs resemble those of our simulated reefs, but other zonation patterns cannot be reproduced accurately because our simulation experiments do not consider the interactions between multiple parameters found on many existing reefs.     

Effects of lindane and deltamethrin on zooplankton communities of experimental ponds

Two insecticides, lindane (321 µg l^–1) and deltamethrin (13 µg l^–1) were employed in a four mesocosm experiment (two ponds of 10 m³ and two of 16 m³) to asses the impact of water pollution by pesticides. Resistance of the different zooplankton species was variable and depended upon both the group and the insecticide concentration. No effect of lindane was observed on macrozooplancton such as Cladocera and Copepoda. In the deltamethrin-treated pond, all species of zooplankton were found dead a day after the treatment. The microzooplankton (Rotifera and copepod nauplii) were highly susceptible to both insecticides. Although the larvae of Chaoborus were present in the ponds after the treatments, their density decreased (less than 1 individual l^–1). The elimination of filter-feeding zooplankton by deltamethrin was followed by an increase of the concentration of chlorophyll a in the post-treatment period. Two months later the original zooplankton population recovered, with the addition of a new and dominant species: Ceriodaphnia reticulata.     

Interactive effects of productivity and predation on zooplankton diversity

Recent studies suggest the necessity of understanding the interactive effects of predation and productivity on species coexistence and prey diversity. Models predict that coexistence of prey species with different competitive abilities can be achieved if inferior resource competitors are less susceptible to predation and if productivity and/or predation pressure are at intermediate levels. Hence, predator effects on prey diversity are predicted to be highly context dependent: enhancing diversity from low to intermediate levels of productivity or predation and reducing diversity of prey at high levels of productivity or predation. While several studies have examined the interactive effects of herbivory and productivity on primary producer diversity, experimental studies of such effects in predator‐prey systems are rare. We tested these predictions using an aquatic field mesocosm experiment in which initial density of the zooplankton predator Notonecta undulata and productivity were manipulated to test their interactive effects on diversity of seven zooplankton, cladoceran species that were common in surrounding ponds. Two productivity levels were imposed via phosphorus enrichment at levels comparable to low and intermediate levels found within neighboring natural ponds. We used open systems to allow for natural dispersal and behaviorally‐mediated numerical responses by the flight‐capable predator. Effects of predators on zooplankton diversity depended on productivity level. At low and high productivity, prey species richness declined while at high productivity it showed a unimodal relationship with increasing the predator density. Effects of treatments were weaker when using Pielou's evenness index or the inverse Simpson index as measures of prey diversity. Our findings are generally consistent with model predictions in which predators can facilitate prey coexistence and diversity at intermediate levels of productivity and predation intensity. Our work also shows that the functional form of the relationship between prey diversity and predation intensity can be complex and highly dependent on environmental context.