Impact of epibenthic predation on estuarine intertidal harpacticoid copepod populations

The extent to which energy is transferred directly from benthic meiofauna to epibenthic predators was investigated on an intertidal sand-flat in the Exe estuary, southwest England, during 1981–1982 and compared with data obtained from an intertidal mud-flat in the Lyhner estuary, also in south-west England, between 1978 and 1981. Two species of flatfish (Pleuronectes platessa L. and Platichthys flesus L.), two species of goby [Pomatoschistus microps (Krøyer) and P. minutus (Pallas)], brown shrimp (Crangon crangon L.) and shore crabs (Carcinus maenas L.) are the most common epibenthic predators feeding on the benthic invertebrates in these locatites. Harpacticoid copepods are the only component of the meiofauna to form a significant part of the diet of early juvenile stages of these predators, particularly the invertebrates. Harpacticoids are a more important source of food for predators feeding over the sand-flat than for those feeding on the mud-flat because in the sand-flat alternative prey of suitable size, such as small annelids, are absent. Moreover, the impact of predation on the mud-flat is spread over the whole harpacticoid species spectrum whereas on the sandflat it is confined almost entirely to a single species, Asellopsis intermedia (T. Scott). Flatfish, gobies and shrimp consume daily an estimated 0.01 to 0.1% of the standing stock of A. intermedia and account for between 12 and 22% of the observed reduction in the population of this species between July and October. Therefore, only a very small proportion of total meiofauna biomass is transferred directly to higher trophic levels.     

Meiofauna on the seagrass Thalassia testudinum: population characteristics of harpacticoid copepods and associations with algal epiphytes

The composition and abundance of bladedwelling meiofauna was determined over a 15 mo period (1983–1984) from a Thalassia testudinum Banks ex König meadow near Egmont Key, Florida, USA. Harpacticoid copepods, copepod nauplii, and nematodes were the most abundant meiofaunal taxa on T. testudinum blades. Temporal patterns in species composition and population life-history stages were determined for harpacticoid copepods, the numerically predominant taxon. Sixteen species or species complexes of harpacticoid copepods were identified. Harpacticus sp., the most abundant harpacticoid, comprised 47.8% of the total copepods collected, and was present throughout the study. Copepodites dominated the population structures of the blade-dwelling harpacticoid species on most collection dates. Ovigerous females and/or copepodites were always present, indicating continuous reproductive activity. Results suggest that epiphytic algae influence meiofaunal abundance on seagrass blades, as densities of most meiofaunal taxa at Egmont Key were positively associated with percent cover of epiphytic algae throughout the study. The majority of significant correlations between meiofaunal density and cover of epiphytic algae involved filamentous algae, although encrusting algae dominated the epiphytic community. It appears that resources provided by epiphytic algae to seagrass meiofauna (additional food, habitat, and/or shelter from predation) may be associated with algal morphology.     

Patterns of selective predation by juvenile,benthivorous fish on estuarine macrofauna

Benthic feeding on macrofauna was studied in juveniles of the sparids Lithognathus lithognathus and Rhabdosargus holubi in the upper reaches of the Gamtoos Estuary, South Africa. Fish and benthic macrofauna were sampled simultaneously, and the selection of invertebrate prey assessed. Both fish species strongly selected for corophioid amphipods and consumed other benthic taxa in low numbers. R. holubi also exploited aquatic autotrophs, while L. lithognathus had a narrow prey-spectrum, feeding almost exclusively on the tube-dwelling amphipod Grandidierella lignorum. G. lignorum was the most abundant prey species, both in the benthos and the fish's diet. This species also showed prominent behavioural differences between the sexes; males were markedly more active on the sediment surface than females, who rarely left their tubes during the day. Males switched from an infaunal to epifaunal microhabitat in search of receptive females, concurrently increasing their exposure to fish predators. Consequently, L. lithognathus selected significantly more males than female amphipods, causing a marked bias towards females in the sex ratio and age-structure of the amphipod population. Juvenile amphipods were less preyed upon, presumably as a result of lower prey-detection or capture efficiency by the predators. Accepting current notions about predation as an important structuring element for benthic communities, our data also stress the prominence of size-and sex-selective predation in structuring individual prey populations.     

A field colonization experiment with meiofauna and seagrass mimics: effect of time,distance and leaf surface area

From a conservation point of view, it is essential to know how fast an ecosystem can recover after physical disturbance. Meiofauna and especially harpacticoid copepods are abundant in seagrass beds and are therefore useful to study ecosystem recovery after disturbance. In the western Caribbean coast, a fragmented Thalassia testudinum seagrass bed was selected to conduct a colonization field experiment by means of plastic seagrass mimics. Meiofauna colonization, with special emphasis on harpacticoid copepods, was followed in relation to: (1) colonization time (2, 4, 6, 10, 14 and 21 days); (2) distance to source of colonizers (close and far series) and (3) leaf surface area to colonize (small, medium, large). Colonization was recorded after 2 days with average meiofauna densities of 480 ind/100 cm² (close) and 1350 ind/100 cm² (far) of leaf surface area, while on average 400 ind/100 cm² were collected from the natural seagrass plants. In this early phase, the meiofauna diversity was high, with on average 8 taxa. A longer period of colonization (21 days) showed an increased meiofaunal density and diversity (average density: 3220 ind/100 cm², 13 taxa). Increasing meiofauna colonization with time is probably related to the development of a biofilm making the leaf more attractive for meiofauna. The effect of distance was not so pronounced as that of time. Total absolute densities were highest in the far series (5 m away from natural seagrass patch), mainly because of nematode densities. Meiofauna diversity was lower in the far series than in the close series (at the border of the natural seagrass patch). A larger individual leaf surface area did not affect the overall meiofauna densities but had a significant positive effect on copepod densities. Larger surface areas promoted the presence of epiphytic copepod families such as Tegastidae and Dactylopusiidae. Overall, we found a rapid recovery of meiofauna in fragmented seagrass beds with primary colonizers (both nematodes and benthic opportunistic copepods) originating from the sediment and later colonizers as epiphytic copepods and their nauplii from the local seagrass regeneration pool.     

Environmental stability and seasonality of a harpacticoid copepod community

The southwest monsoon on the west coast of India brings about dynamic changes in estuaries and coastal waters. The response of the meiofauna to monsoonal rain is obvious, but the impact of such environmental changes on the community structure of harpacticoid copepod species and their seasonality in the estuaries influenced by the tropical monsoon is poorly understood. In this study the spatial and temporal variability in abundance and community structure of meiobenthic copepods was investigated over an annual cycle (June 1983 to June 1984), in an estuary influenced by the tropical monsoon. Total meiofaunal abundance showed wide variations in space and time. Minimum and maximum densities were observed in the monsoon and pre-monsoon seasons, respectively. Quick recovery of harpacticoid populations in the early post-monsoon season indicated their recuperative power under adverse conditions. Of the 25 species recorded from lower, middle and upper reaches, eight comprised over 70% of the total copepod population. The peak and low occurrence of dominant species displayed striking correlations with the summer and rainy seasons, respectively. While other species were restricted in distribution, Stenhelia longifurca was recorded from all salinity regimes. The harpacticoid community was greatly influenced by the onset of the monsoon period, and their spatial and temporal variabilities were related with physico-chemical parameters and the variability of these parameters in the estuary.     

On predation of pelagic larvae and early juveniles of marine bottom invertebrates by adult benthic invertebrates and their passing alive through their predators

The dynamic quantitative balance between prey and predator invertebrate species inhabiting the same shallow-shelf (sublittoral level bottom) benthic communities was first discussed by Thorson (1953). Thorson considered the exact timing of larval settlement of prey and predator species possessing pelagic development and temporal supression of the adult predators' feeding activities during reproduction at the time of the preys' settlement to constitute the major factors which facilitate survival of the prey species in such communities. However, information obtained demonstrates that Thorson's “mechanism of balance between predator and prey species of benthic communities” is not always effective in securing survival from predation not only of the prey's spat but even sometimes of the predator's spat also. Because of this, the “mechanism” can not be rated as universally effective in all situations. Analysis of the data so far published demonstrates that, in marine benthic communities, especially in shallow-shelf waters, it is not uncommon for gametes, larvae, or early juveniles of different prey species to pass alive through suspension (filter)-feeding and deposit-feeding adult invertebrates preying on them. Sometimes development can even continue after excretion by predators. The hypothesis of Voskresensky (1948) and Goycher (1949) of the importance of this phenomenon for the maintenance and recruitment of the mussel Mytilus edulis and other filter-feeding lamellibranchs of nearshore waters preying on their own and other lamellibranch pelagic larvae must be rejected on the basis of accumulated data on their feeding and general biology and on the adverse influence of the mucous of their faecal pellets and pseudofaeces on the larvae excreted by them alive. The data considered here demonstrate that, although the passing alive of larvae and spat of benthic invertebrates through benthic predators is not uncommon in shallow-shelf bottom-communities, it plays no important role in the processes of maintenance and recruitment of the species and communities involved nor of the marine benthos as a whole. The actual ecological significance of predation on pelagic larvae and bottom spat of benthic invertebrate prey species by all three main trophic groups of marine benthos (suspension or filter-feeders, deposit-feeders, carnivores) and its importance to predator-prey dynamics in marine benthic communities remains open to debate until more reliable quantitative data become available.     

Effect of temperature on life cycles of nematodes associated with the mangrove (Rhizophora mangle) detrital system

The effect of temperature on the life history of various representatives of the meiofauna associated with decaying mangrove (Rhizophora mangle) leaves was investigated. Life cycles, recorded in days, for 6 species of marine nematodes cultured at a temperature of 24°C are as follows: Rhabditis marina 2 1/4; Diplolaimelloides sp., 7; Diplolaimella ocellata, 11 1/2; Enoplus paralittoralis, 22; Oncholaimus sp., 29; Haliplectus dorsalis, 34. In general, life cycles become shorter with increased temperatures; however, as temperatures approach the upper limits which support reproduction, life cycles become slightly lengthened. For most species, the ability to complete a life cycle was inconsistent within the temperature range of 33° to 35°C. Studies with two harpacticoid copepod and two foraminifera species tend to support the 33° to 35°C range as being the thermal stress zone.Contribution No. 1697 from the University of Miami, Rosentiel School of Marine and Atmospheric Science, 10 Rickenbacker Causeway, Miami, Florida 33149, USA.     

Predation risk assessment by green frog (<Emphasis Type="Italic">Rana clamitans</Emphasis>) tadpoles through chemical cues produced by multiple prey

Many prey assess predation risk through predator chemical cues. Numerous studies have shown that (1) prey sometimes respond to chemical cues produced by heterospecifics and (2) that many species are capable of associative learning. This study extends this research by focusing on predation risk assessment and antipredator behavior in environments containing chemical cues produced by multiple prey species. The results show that green frog (Rana clamitans) tadpoles (1) assess risk from the chemical cue produced during predation by a heterospecific (gray tree frog, Hyla versicolor, tadpoles) and (2) can exhibit similarly strong behavioral responses to a mix of conspecific and heterospecific cues compared to conspecific cue alone, depending on their conditioning environment. I then discuss how the prey choice of the predators and the relative abundances of the prey species should influence the informational value of heterospecific cues.     

The effect of predation on artificial reef juvenile demersal fish species

There is a concern that artificial reefs (AR) may act purely as fishing aggregation devices. Predators attracted to ARs can influence the distribution and abundance of prey fish species. Determining the role of predators in AR is important in advancing the understanding of community interactions. This paper documents the effects of predation on fish assemblages of AR located near a coastal lagoon fish nursery. The Dicentrarchus labrax is a very opportunistic species preying on juveniles (0⁺ and 1⁺ age classes) of several demersal fish species on the ARs. Reef prey and sea bass abundance were negatively correlated. The mean numbers of prey per sea bass stomach increased with the increase of reef fish prey abundance, suggesting that predation has a significant influence, resulting in a decrease in prey abundance. Prey mortality (4–48%) of demersal reef fish associated species depends on bass density. Prey selection was related both with prey abundance and vulnerability. Results showed that D. labrax predation on AR-fish associated species can increase prey natural mortality. However, the role of bass predation on the ecological functioning of exploited ARs is not clear. There may be increases in local fishing yields due either to an increase in predator biomass through aggregation of sea bass attracted to ARs or to greater production. In contrast, predation on juveniles of economically important reef fish preys, especially the most frequent and abundant (Boops boops), can contribute to a decrease in recruitment to the fishery. Our results indicate that inter-specific interactions (predator–prey) are important in terms of conservation and management, as well as for the evaluation of the long-term effects of reef deployment. Thus, it is necessary to consider ecological interactions, such as predation, prior to the development and deployment of artificial habitats as a tool for rehabilitation.     

Submarine experiments on benthic colonization of sediments in the western Baltic Sea. II. Meiofauna

From July, 1972 to February, 1973, meiofauna samples were taken from submerged platforms. Substrates were gravel, fine sand, and clayey mud. The individual samples consisted of 20 cm² 100 cm³. A diver collected the samples by hand. Some days after submersion of the platforms, colonization by meiofauna began. The submerged substrates were colonized almost exclusively by nematodes and harpacticoids; gastrotrichs were never present. In general, the harpacticoid fauna can be classified into two groups: the first, confined to the mesopsammon, did not colonize the platforms; the second, not limited to the interstitial habitat, did colonize. Two harpacticoid species, Ameira parvula and Danielssenia typica, were markedly dominant. The main migration route to the submerged platform was up the securing lines.Contribution no. 35 from the Sonderforschungsbereich 95, Kiel.     

Patterns of shell repair in articulate brachiopods indicate size constitutes a refuge from predation

The cost of overcoming prey defenses relative to the value of internal tissues is a key criterion in predator/prey interactions. Optimal foraging theory predicts: (1) specific sizes of prey will result in the best returns to predators, and (2) there will often be a size at which the cost/benefit balance is low enough to effectively exclude predation. Data presented here on styles of repaired shell damage and size at which injury had been sustained was collected from samples of terebratulide brachiopods from the Antarctic Peninisula (Liothyrella uva), Falkland Islands (Magellania venosa and Terebratella dorsata) and Chile (M. venosa). The predominant form of damage on shells was indicative of predators attacking the valve margins. The modal size for repaired damage was more than 10 mm smaller than the modal size for the overall size distribution in each species and there were no repaired attacks in the largest size classes of any species. These data suggest that size forms a refuge from predation, as would be predicted by optimal foraging theory. The optimal sizes that predators appeared to attack vary between species, as do the sizes that provided a refuge from predation. High levels of multiple repairs (19% of the M. venosa population from the Falkland Islands sampled had 2 or more repairs) suggest that the mortality following attack is low, suggesting that many predators abandon their attacks.     

Increased production of faecal pellets by the benthic harpacticoid Paramphiascella fulvofasciata: importance of the food source

The re-use of faecal pellets in the water column before sinking to the seafloor is known as an important pathway in marine food webs. Especially planktonic copepods seems to be actively use their faecal pellets. Since benthic copepods (order Harpacticoida) live in the vicinity of their pellets, it remains unclear how important these pellets are for their feeding ecology. In the present study a laboratory experiment was conducted to analyse the importance of faecal pellets for the feeding ecology of the harpacticoid Paramphiascella fulvofasciata and its grazing pressure on two diatom species (Seminavis robusta, Navicula phyllepta). By quantifying the amount and volume of the produced faecel pellets in different treatments, it was tested to what extent the food source and the lack of faecal pellets would influence the production of faecal pellets. We found that the grazing pressure of P. fulvofasciata depended on the diatom density since only a top-down effect could be found on the smaller Navicula cells during its initial exponential growth phase. The grazer had a negative effect on the diatom growth and controlled the cell density to about 4,000 cells/cm². In spite of the fact that the addition of faecal pellets did not show a significant positive effect on the assimilation of diatoms, the removal of faecal pellets strongly promoted the pellet production. Especially when grazing on Navicula the harpacticoid P. fulvofasciata produced significantly more and smaller faecal pellets when the pellets were removed. This outcome illustrates the need for faecal pellets of this harpacticoid copepod when grazing on the diatom Navicula. Apart from its selection for smaller diatom cells, it was suggested that the colonisation of heterotrophic bacteria enriched these pellets. This study is the first to indicate that trophic upgrading occurs on faecal pellets and not only on the initial autotrophic food sources per se.     

Effects of water motion and prey behavior on zooplankton capture by two coral reef fishes

Water motion is an important factor affecting planktivory on coral reefs. The feeding behavior of two species of tube-dwelling coral reef fish (Chaenopsidae) was studied in still and turbulent water. One species of blenny, Acanthemblemaria spinosa , lives in holes higher above the reef surface and feeds mainly on calanoid copepods, while a second, A. aspera , lives closer to the reef surface, feeds mainly on harpacticoid copepods, and is exposed to less water motion than the first. In the laboratory, these two blenny species were video recorded attacking a calanoid copepod ( Acartia tonsa, evasive prey) and an anostracan branchiopod (nauplii of Artemia sp., passive prey). Whereas A. spinosa attacked with the same vigor in still and turbulent water, A. aspera modulated its attack with a more deliberate strike under still conditions than turbulent conditions. For both fish species combined, mean capture success when feeding on Artemia sp. was 100% in still water and dropped to 78% in turbulent water. In contrast, when feeding on Acartia tonsa, mean capture success was 21% in still water and rose to 56% in turbulent water. We hypothesize that, although turbulence reduces capture success by adding erratic movement to Artemia sp. (passive prey), it increases capture success of Acartia tonsa (evasive prey) by interfering with the hydrodynamic sensing of the approaching predator. These opposite effects of water motion increase the complexity of the predator-prey relationship as water motion varies spatially and temporally on structurally complex coral reefs. Some observations were consistent with A. aspera living in a lower energy benthic boundary layer as compared with A. spinosa: slower initial approach to prey, attack speeds modulated according to water velocity, and lower proportion of approaches that result in strikes in turbulent water.Communicated by P.W. Sammarco, Chauvin     

Selective feeding of littoral harpacticoids on diatom algae: hungry gourmands?

We studied benthic harpacticoid grazing on diatom algae from two sites on the White Sea intertidal sandflat. Diatoms from sediments and from harpacticoid gut contents were sampled in situ, identified and counted, and grazing rates were calculated by two ways: (1) using potential daily ration estimations and (2) from the gut content and gut-residence time data. Paraleptastacus kliei did not contain any diatoms in the guts and presumably fed on other objects (bacteria or flagellates). Two other dominating species studied, Heterolaophonte minuta and Huntemannia jadensis, contained an average of 604 and 222 diatom cells per specimen. Diet composition differed significantly from the natural algal community. Two diatoms of intermediate cell size (Nitzschia palea var. debilis and Navicula sp.) contributed 92–97% of gut content for H. minuta and 58–81% for Hn. jadensis, whereas these diatoms amounted to only about 10% of biomass in native community. Mean consumption rates were estimated as 50–200 μg of wet biomass/day/cm², so the harpacticoids grazed only between 3 and 11% of the total microalgae biomass per day. The grazing impact on the two preferred diatom populations, however, was much more intensive, 10–30% per day for Navicula sp. and 55–228% for N. palea. Therefore, native harpacticoid populations demonstrate highly selective feeding and could be strongly limited by their food in spite of seemingly plentiful total abundance of microphytobenthos. This disproportionally high grazing pressure upon some species apparently could affect the structure of microalgae communities resulting in low relative abundance of mid-sized forms. We hypothesize that a very dynamic spatio-temporal distribution of epibenthic harpacticoids (short-living micropatches) may be the possible adaptation to such local food limitation.     

Cercopagis pengoi and Mysis spp. alter their feeding rate and prey selection under predation risk of herring (Clupea harengus membras)

The anti-predator behaviour of Baltic crustacean planktivores was studied in feeding experiments under predation pressure of herring. The experiments were conducted with pelagic mysids: Mysis mixta and Mysis relicta, and with Cercopagis pengoi, a non-indigenous cladoceran, which invaded the Baltic Sea in 1992. Zooplankton was offered as prey. Two kinds of experiments were performed in the absence and presence of chemical predator cues: (1) two-prey experiments with prey, which have poor or good escape responses and all three planktivores and (2) natural prey experiments with mysids in natural zooplankton assemblages. The results showed that all three species reacted to the chemical cue of herring by decreasing their feeding rate and altering prey selection. C. pengoi selected easily captured prey (rotifers) in two-prey experiments under predation risk while selection for any prey was evident in mysids in natural prey experiments only in the absence of predator cues. This indicates that planktivores have different anti-predator strategies, which are modified by their own prey capture abilities. C. pengoi was a very efficient predator on small prey with size-specific prey consumption rate 5 to 18 times the rate of mysids. Results show that the studied planktivores are capable of adjusting their feeding behaviour to decrease their conspicuousness in order to increase survival under predation risk. Further, results support the view that C. pengoi has adapted well to the Baltic ecosystem, sharing food niche with pelagic mysids and most probably having a strong influence on the whole pelagic food web.     

Predation on zooplankton by the benthic anthozoans Alcyonium siderium (Alcyonacea) and Metridium senile (Actiniaria) in the New England subtidal

The Alcyonacean octocoral Alcyonium siderium Verrill and the sea anemone Metridium senile (L.), the only common perennial zooplanktivores on shallow (16-m depth) subtidal rock walls in much of northern New England, USA, are of similar heights and overlap in their habitat and microhabit distributions. The coelenteron contents of both species were sampled at four-hour intervals over a diel cycle and were compared to zooplankton available in the water at 1 to 5 cm from the rock wall, the height at which the cnidarians held their feeding tentacles. Prey in coelenterons of A. siderium were significantly smaller (means of 256 to 345 m), and those in coelenterons of M. senile were equal to or slightly greater in length (means of 415 to 1006 m) than the available zooplankton. The diets of A. siderium and M. senile differed significantly from each other and from the available zooplankton. A. siderium showed strong positive electivites for ascidian larvae and for foraminiferans, and strongly negative electivities for most crustaceans. M. senile had strong positive electivities for barnacle cyprids, ascidian larvae, and gammarid amphipods, and strong negative electivities for invertebrate eggs, foraminiferans, calanoid and harpacticoid copepods, and ostracods. Electivities may reflect tentacle avoidance or escape by motile prey as well as predator preference. Substratum-associated organisms (e.g. demersal crustaceans, larvae of benthic invertebrates) were the most common items in the diets of both species, suggesting a tight benthic food web, similar to the situation for coral reef anthrozoans which rely on reef-generated zooplankton. A. siderium ate large numbers of ascidian larvae which, as benthic adults, compete for space with A. siderium and can overgrow small colonies. Predation on the larvae of a competing species may alleviate competition by decreasing the competitor's recruitment.     

Population dynamics of three estuarine meiobenthic harpacticoids (Copepoda) in South Carolina

To elucidate the population dynamics of abundant meiofauna, a 15-month field study of 3 species of harpacticoid copepods was conducted in a South Carolina (USA) estuary. Per capita birth and death rates and average brood size (number of eggs per adult female) were lowest for Enhydrosoma propinguum (Brady), Microarthridion littorale (Poppe), and Stenhelia (Delavalia) bifidia (Coull) from November–February, indicating that food or density-independent factors such as temperature regulated winter densities. Reproduction for all species began in February, with the first appearance of copepodites approximately 1 month later. From March–October, M. littorale displayed high per capita birth and death rates (>0.6 individual^-1 day^-1), and high average brood size (8 to 9 eggs per adult female). As this species is epibenthic, and thus susceptible to grazing by detritivores, these data suggest that M. littorale was limited by predation. S. (Delavalia) bifidia displayed a sharp increase in density, birth rate and average brood size in July, but birth and death rates (0.3 individual^-1 day^-1) were always lower than those of M. littorale and average brood size gradually decreased from July–December. As this species burrows and is less susceptible to grazing predation, these data are consistent with the hypothesis that S. (Delavalia) bifidia became food-limited as population growth approached its carrying capacity. Although E. propinguum was the most abundant species, quantification of birth potential was not accomplished due to an underestimate of the number of ovigerous females. Therefore, little can be concluded about the mechanism of the summer density regulation of E. propinguum.Contribution No. 274 from the Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina.     

Minimal predation upon meiofauna by endobenthic macrofauna in the Exe Estuary,south west England

Field manipulation experiments were performed in the Exe Estuary, south west England, in October 1988, to investigate the importance of the meiofaunamacroinfauna trophic link in benthic trophodynamics. Four hypothetically meiofauna-predacious endobenthic macrofauna species were selected for manipulation using the criteria of high abundance and different modes of feeding: Cerastoderma edule (filter-feeder), Nereis diversicolor (omnivorous scavenger), Ophelia bicornis (sand-ingester), Scrobicularia plana (deposit-feeder). Enclosures constructed from plastic tubing, 63 m nylon monofilament mesh and galvanized steel were deployed, containing adult members of these taxa at densities raised to approximately four times that of the surrounding sediment. The experiments ran for 12 tidal cycles. Differences in phyletic meiofaunal abundance between treatment and control enclosures at the termination of the experiment were assessed using both uni- and multivariate dataanalysis techniques. Only two univariate significant differences (p<0.05) existed for the N. diversicolor treatment and two for the S. plana treatment. Annelida, Turbellaria and copepod nauplii were the only meiofauna taxa affected. No univariate significant differences were recorded for either the C. edule or O. bicornis treatments. Multidimensional scaling ordination of the data revealed no consistent changes in community composition between treatments and controls. It is concluded that the experiments provide evidence of minimal predation by macroinfauna upon meiofauna.     

Movements of intertidal whelks (Morula marginalba and Thais orbita) in relation to availability of prey and shelter

The movements of two species of predatory whelks (Muricidae: Gastropoda) were examined on several rocky seashores in New South Wales, Australia, from 1980 to 1984. The smaller species, Morula marginalba Blainville, moved less far than the larger species, Thais orbita (Gmelin). Intermittent but rapid movement allowed T. orbita to shelter in exposed places low on the shore, but to feed on their preferred prey, barnacles, much higher up to shore. This resulted in a dynamic and unpredictable pattern of vertical foraging movements for this species. Compared to areas with intact prey or with barnacles that had been killed but not removed, M. marginalba moved quickly away from areas from which barnacles had been experimentally removed. Many M. marginalba left experimental plots on two shores without prey or crevices in which to shelter, whereas some stayed over 30 d in areas with either prey or crevices (but not both), and most remained in plots where both prey and crevices were available. Thus, the presence of suitable prey and crevices seems to influence the movements of both species, which respond in different ways, corresponding to their different mobilities. These results are discussed in relation to the behavioural cues involved and predation by the whelks.     

Niche segregation and habitat specialisation of harpacticoid copepods in a tropical seagrass bed

Several harpacticoid copepod species are adapted to an epiphytic lifestyle. Previous studies on tropical seagrass meiofauna mainly focussed on the epiphytic communities and neglected the benthic component. The present study aims to document the benthic harpacticoid copepod communities sampled from different sediment depth horizons adjacent to five seagrass species in the intertidal and subtidal zone of a tropical seagrass bed (Gazi Bay, Kenya). Two benthic copepod communities could be identified mainly based on the tidal position of the samples: a first community was collected near the intertidal seagrasses Halophila ovalis and Halodule wrightii; a second community occurred near the subtidal seagrasses Thalassia hemprichii, Syringodium isoetifolium and Halophila stipulacea. The first community was mainly determined by sediment characteristics (e.g. skewness), while the second community was split off based on organic matter content (% TOM), nutrient and pigment values. A subtle combination of horizontal and vertical niche segregation was reported for the dominant copepod families. Species of the families Thalestridae, Laophontidae and Diosaccidae were structured by tidal position and showed a strong preference for the subtidal zone. The opposite strategy, i.e. a clear preference for the intertidal zone, was found for copepods belonging to the families Paramesochridae and Canuellidae. In addition, Apodopsyllus africanus (Paramesochridae) was well-adapted to stress and was concentrated in the deeper sediment layers near the subtidal seagrasses. On the other hand, Canuellidae, as filter feeders, were concentrated in the upper centimetres of the sediment. The families Ectinosomatidae and Cletodidae did not show any vertical or horizontal segregation. On the species level, however, clear horizontal niche segregation was detected for the family Cletodidae. In addition to the reported ecological results, the study material was used to evaluate different niche definitions. We found tidal position to be the most important factor forcing harpacticoids to specialise. Sediment depth horizon was less powerful in dividing the families into different guilds (from specialists to generalists) based on standardised niche breadth. The present study documents the subtle habitat partitioning of co-existing species in a limited area and its role in sustaining high biodiversity in the community.