Predator-limited population growth of the copepod Pseudocalanus sp.

The impact of predators on population growth of Pseudocalanussp. was investigated in Dabob Bay, Washington. Mortality ofPseudocalanus sp. was determined from stage-specific survivorship,from seasonal changes in mortality rates of adult males andfemales and from incidence of injuries to adult copepods. Theprincipal predators of adult Pseudocalanus were identified asthe predatory copepod Euchaeta elongata, the omnivorous euphausiidEuphausia pacifica and the chaetognath Sagirta elegans. Predatorattack rates - and prey mortality rates - are highly density-dependentand thus sensitive to prey dispersion in the water column, particularlyto layering in the vertical plane. Predation rates by the threeprincipal predators exceeded 100% of the recruitment rate toadult Pseudocalanus sp. beginning in early summer, thus restrictingpopulation growth. Planktivorous fish predation (by adult three-spinestickleback, Gasterosteus aculeatus, and juvenile chum salmon,Oncorhynchus keta) on Pseudocalanus sp. adults was estimatedto be two orders of magnitude lower than consumption rates bypredatory zooplankton, at a deep water station in July. Analysisof seasonal changes in prey ingested by Sagitta elegans revealedthat Pseudocalanus sp. was the major prey item of S. elegansin April (61.0% of prey) and in June (67.0% of prey), thereafterdeclining seasonally in importance. Predation by S. elegansvaried seasonally with changes in chaetognath stage structure,vertical distribution and diapause, not size structure alone.Although chaetognath recruitment and population growth appearto be directly coupled to the abundance of Pseudocalanus sp.,predation by S. elegans has little reciprocal impact on Pseudocalanussp. population growth; hence asymmetries may occur in the interaction of planktonic prey and predators.     

Effects of chaetognath predation and nutrient enrichment on enclosed estuarine copepod communities

Summary The effects of predation by the chaetognath Sagitta hispida and nutrient enrichment on estuarine copepod community structure were studied by experimental manipulations in large enclosures. Chaetognath abundance and nutrient addition rates were manipulated in a factorial design. Predation by chaetognaths resulted in a significant decline in the relative abundance of Acartia tonsa and an increase in the relative abundance of Oithona colcarva. However, these effects were evident only at chaetognath densities far higher than observed in natural populations. Nutrient enrichment resulted in a decline in relative abundance of Paracalanus crassirostris, and, in the absence of chaetognath predation, an apparent increase in the relative abundance of A. tonsa. The effects of chaetognath predation were independent of enrichment level, apparently because of the absence of effects of enrichment on total copepod and chaetognath densities.     

Composition, spatial and temporal variations of Chaetognatha in Guanabara Bay, Brazil

This paper presents the specific composition and distributionof adult chaetognaths in Guanabara Bay, Brazil. A comparisonhas been made between chaetognath and copepod distribution patterns.Three areas were distinguished within Guanabara Bay: an externalarea, influenced by oceanic waters; an internal area, influencedby fluvial waters; and a transitional area between the two.Monthly samples were collected by horizontal surface hauls usinga conical net with 200µm mesh. Three species were identified:Sagitta friderici, S.enflata and S.hispida. Sagitta fridericiwas the dominant species and occurred throughout the year atthe three areas. Sagitta enflata and S.hispida were limitedto the more saline waters. Generally, greater densities of chaetognathswere observed following copepod peakden sities. Temperatureand salinity associated with copepod abundance were the mostimportant factors affecting the distribution pattern and theabundance of chaetognath species in Guanabara Bay.     

Trophic importance of the chaetognaths Eukrohnia hamata and Sagitta gazellae in the pelagic system of the Prince Edward Islands (Southern Ocean)

Trophodynamics and predation impact of the 2 dominant chaetognaths Eukrohnia hamata and Sagitta gazellae were investigated at 19 stations in the vicinity of the Prince Edward Islands and at a 24-h station occupied at the sub-Antarctic Front in late summer (April/May) 1996. During the entire investigation, the zooplankton assemblages were numerically dominated by copepods with densities ranging from 21 to 170 ind. m⁻³. Amongst the copepods, Clausocalanus brevipes, Metridia gerlachei and M. lucens dominated accounting for >90% of the total. Generally, chaetognaths were identified as the second most important group composing at times up to 30% (mean = 14.7%) of total zooplankton abundance. Of the two chaetognath species, E.␣hamata was generally numerically dominant. Gut content analysis showed that both chaetognath species are opportunistic predators generally feeding on the most abundant prey, copepods. No feeding patterns were evident during the 24-h station, suggesting that both species feed continuously. The feeding rates of E. hamata ranged from 0 to 0.50 prey ind. day⁻¹ and between 0 and 0.90 prey ind. day⁻¹ for S. gazellae. The maximum total predation impact of E. hamata was equivalent to 5.2% of the copepod standing stock or up to 103% of copepod production per day. For S. gazellae the predation impact was lower, reaching a level of 3.2% of the copepod standing stock or 63% of the daily copepod production. Chaetognaths can, therefore, be regarded as an important pelagic predator of the Prince Edward Islands subsystem. Received: 27 March 1997 / Accepted: 11 September 1997     

Vertical distribution, life cycle and production of the chaetognath Sagitta crassa in Tokyo Bay, Japan

The daytime vertical distribution of Sagiita crassa in TokyoBay was examined from February 13, 1988 to February 20, 1989.High densities of larger-size chaetognaths were found near thesea bottom, whereas the smaller animals tended to inhabit theupper layers. This feature of distribution is discussed in relationto the distribution of their main food organisms, e g. Pseudodiaptomusmarinus, Acartia omoru, Centropages abdomialis and Oithona davisae.The two periods of replacement of two morphs were confirmedby the variation only in mean body length of this chaetognath,unlike the previous authors who made additional morphologicalobservations. It was hypothesized that S.crassa has at leastfive generations Two generations, including mostly the largerforms, had higher growth rates than the generation consistingmainly of the small form. Yearly respiration of S.crassa was8.2 g C m^–Abstract. Yearly production of this animal wasestimated to be 3.8 g C m^–. A feeding estimate revealedthat chaetognaths require a prey production of 13.1 g C m^–year^–1. The impact of this chaetognath on the prey populationin Tokyo Bay and the propriety of an estimated value of annualproduction of S crassa is discussed.     

Feeding of Sagitta enflata and vertical distribution of chaetognaths in relation to low oxygen concentrations

Zooplankton samples were collected in Mejillones Bay, northernChile (23°00'15'S, 70°26'43'W ). Sampling was conductedat 4 h intervals, for 24 h during three seasons, austral spring(October 2000), summer ( January 2001) and winter (August 2001)at three different strata (0–25, 25–50 and 50–100m). Five species of chaetognaths were collected. Sagitta enflatawas the most abundant species, representing up to 65% of allchaetognaths in total numbers, followed by Sagitta bierii, makingup 34% of the total abundance of chaetognaths. S. enflata wasdistributed mainly above the Oxygen Minimum Zone, while S. bieriiremained below this zone. Feeding rates were relatively constantwithin the upper layer (0–25 m depth), for each samplingdate, averaging 1.2 prey S. enflata day^–1, and decreasingwith depth. Gut content analyses demonstrated that predationwas principally focused on small copepods (<1500 µm),with greatest feeding activity occurring at night. The dailypredation impact on the total standing stock of small copepodsvaried seasonally between 6% in spring and 0.4% in winter. Thispercentage may represent a negligible impact on the entire copepodcommunity, but it is relevant at the species or genus level,since S. enflata removed more than 20% of the standing stockof Centropages brachiatus and Corycaeus sp. Thus, during someperiods of the year, chaetognaths may strongly influence theabundance and size distribution of copepods in coastal upwellingecosystems.     

Predation impact of Cyclops vicinus on the rotifer community in Lake Constance in spring

The predation impact of Cyclops vicinus on rotifers was studiedunder near-natural conditions in small enclosures to evaluatewhether copepod predation is responsible for the decline ofrotifers in Lake Constance in spring. Cyclops vicinus fed selectivelyon Synchaeta spp.; Keratella and Polyarthra spp. were not selectedfor. Predation rates increased with prey density up to a maximumof 37 Synchueta day^–1 at a density of 1.6 x 10⁶ Synchaetam^–2, i.e. at -1200 Synchaeta l^–1. Calculation ofcropping rates suggests that Cyclops alone can control the abundanceof Synchaeta in spring, i.e. that mainly Cyclops is responsiblefor the decline of Synchaeta species in Lake Constance in May.     

The production and ingestion of faecal pellets by nauplii of marine calanoid copepods

The downward transport of organic matter as zooplankton faecalmaterial is influenced by copepods which fragment, ingest andrecycle some of the pellet contents. Most of this activity hasbeen attributed to the later copepodite stages and the adults,but little is known about the role of nauplii. Stage-relateddefaecation rates during the naupliar development of two speciesof copepod, Calanus helgolandicus and Pseudocalanus elongatus,were quantified in a series of laboratory experiments. The productionof faecal material commenced soon after the appearance of theNIII in both species and increased throughout naupliar development.The causes of the increase were the formation of larger pelletsby later stages in Calanus and an increased rate of productionby Pseudocalanus. Calanus nauplii, when supplied with algalfood at concentrations that would support full naupliar development,ingested or broke up the pellets of the smaller Pseudocalanusspecies at rates of 1.15 pellets nauplius^–1 h^–1This consumption increased to 2.96 pellets nauplius^–1h^–1 when the concentration of algal food was reduced toa limiting level. Pseudocalanus was not able to consume thepellets of Calanus. Ingestion of Pseudocalanus faecal pelletsby Calanus could supply a nutritional benefit to a food-limitednauplius.     

Response of ciliates and Cryptomonas to the spring cohort of a cyclopoid copepod in a shallow hypereutrophic lake

The impact of a cyclopoid copepod population on the protozoacommunity (two ciliate categories and Cryptomonas) was assessedweekly during the spring cohort of Cyclops vicinus (one monthduration) in hypereutrophic Lake Søbygård by insitu gradient experiments with manipulation of ambient zooplanktonabundance. As C.vicinus always made up >92% of the zooplanktonbiomass, the response of protozoa is assumed to be a resultof predation by the copepod. Significant effects of copepodbiomass on protozoa net population growth rates were obtainedin the four experiments. Copepod clearance rates were significantlyhigher on oligotrichs than on prostomatids and Cryptomonas butdeclined for all three protozoa categories during the firstthree weeks of the copepod cohort, probably because of the changein developmental instar composition of the copepod population.Grazing impact on protozoa at ambient copepod abundance wasconsiderable (range, 0.05–0.87 day^–1) and could,together with the estimated reproductive potential of protozoans(range, –0.20–0.87 day^–1), account for thedecline in abundance and biomass of protozoa during the cohortdevelopment. Carbon flow from the protozoa to C.vicinus (range,2.8–23.5 µg C l^–1 day^–1) documents thepresence of a trophic link between protozoa and the spring cohortof C.vicinus in Lake Søbygård.     

Distribution and species-specific egg production of Pseudocalanus in the Gulf of Alaska

Pseudocalanus species are important contributors to the secondaryproduction of the northern hemisphere mid- to high-latitudeoceans. In the coastal Gulf of Alaska, Pseudocalanus are presentyear round and are represented by three species. In 2001, Pseudocalanusmimus was the dominant Pseudocalanus species on the shelf duringspring and summer, comprising 30–100% of the total, whilePseudocalanus newmani dominated in Prince William Sound (10–90%).Pseudocalanus minutus were only abundant in Prince William Soundduring early spring. Egg production (by number and volume) wasa function of female prosome length and decreased from springto summer; however, significant variability was attributableto regional influences that were independent of size. For thesame sized female, P. newmani produced more eggs per clutchthan P. mimus. Pseudocalanus mimus, however, tended to havea larger mean egg size than P. newmani. Consequently, clutchvolumes of the two species were indistinguishable. Pseudocalanusegg production rates (EPRs) (eggs female^–1 day^–1)were lower in July and August (ca. 2–4) than April andMay (ca. 1–9), but total egg production by the population(eggs day^–1) was nearly equivalent for the two time periodsdue to higher female concentrations in summer.     

Trophodynamics of the plankton community at Dogger Bank: predatory impact by larval fish

The trophodynamics of a coastal plankton community were studied,focusing on fish larvae and their copepod prey. The major objectiveswere to describe distributional overlap and evaluate the predatoryimpact by larval fish. The study was carried out across DoggerBank in the North Sea, August-September 1991. Sampling transectscrossed tidal fronts off the Bank and plankton at all trophiclevels showed peak abundance within frontal zones. Also Verticallythere was a significant overlap in distributional patterns ofthe plankton. Seven species of fish larvae were abundant, ofthese sprat (Sprattus sprattus) dominated. The abundance ofone group of fish larvae peaked in the shallow water close tothe Bank, whereas other species, including sprat, were foundin deeper water. Prey preference and predation pressure of fishlarvae were assessed using information on prey sizes and growthrates of larvae and the copepod prey. We estimated larval removalof preferred prey sizes to 3–4% day^–1, counterbalancedby a 3–7% day^–1' replenishment from copepod productionand growth. Additional predation pressure on copepods by aninvertebrate predator was estimated to 1–3%day^–1.In conclusion, the dynamics of fish larvae and other zooplankterswere closely linked. At peak abundances of fish larvae (>35mg dry weight m^–2), the accumulated predation on specificsize ranges of copepods, exerted by larvae and other predators,could exceed the ability of copepod replenishment and intra-/interspecificcompetition among predators might take place.     

The life history and vertical distribution of the chaetognath, Sagitta elegans, in Dabob Bay, Washington

Seasonal changes in the population structure and patterns ofvertical distribution and migration of the dominant chaetognath,Sagitta elegans, in a fjord off Puget Sound were determinedwith samples collected at approximately semi-monthly intervalsfrom November 1972 to November 1973. S. elegans bred continuouslyfrom March through August, with major recruitment in June andJuly. Seasonal changes in the vertical distribution of the populationand in the proportion of the population that vertically migratedreflected the stage of maturity and size of the individuals.Small chaetognaths were non-migratory and distributed in thetop 100 m. Stage II individuals were dispersed throughout thewater column; some did migration was evident. Breeding stageswere restricted to a layer between SO and 100 m during the dayand migrated to the surface layer at night, enhancing the probabilityof successful reproduction.     

Distribution and trophic ecology of chaetognaths in the western Mediterranean in relation to an inshore-offshore gradient

This study examines the distribution patterns and feeding ecologyof chaetognaths in the Catalan Sea in relation to mesoscalefeatures along an inshore–offshore gradient. The studywas conducted during two different periods of the year: latespring of 1995 and late summer of 1996. The two periods differedin hydrographic conditions and mesoscale processes, which affectedthe distribution patterns of the different species of chaetognathsfound. The diet of the chaetognaths was mainly composed of copepodsand differed between species. Prey size was not always stronglyrelated to chaetognath size and for certain species, there wasan overlap in prey size spectrum. Trophic niche breadth (ona ratio scale) appeared to be constant with growth. Ingestionrates and predation pressure by chaetognaths did not followa clear trend related to the mesoscale features in the area,such as the presence of a density front. The impact of chaetognathson copepod standing stock appeared to be extremely low (<1%),but it became more relevant when the species and prey size specificityof the chaetognaths was taken into account.     

Vertical distribution of pelagic chaetognaths and feeding of Sagitta enflata in the Central Equatorial Pacific

The vertical distribution and feeding of pelagic chaetognathsat 5°S, 160°W in the Central Equatorial Pacific wereinvestigated using a series of 0–500 m vertical haulswith a VMPS net over a 24 h period between 6 and 7 October 1990.The total number of individuals per haul was between 370 and688. Fourteen species in four genera were found at this station.The most abundant species was Sagitta enflata which comprised32.4–61.1% of the individuals collected from the 0–500m layer. Mesopelagic species made up 9.3–15.1% of thetotal number of individuals. Sagitta enflata and Pterosagittadraco were found in the upper part of the thermocline both byday and at night. The fraction of the population containingfood items (FCF) of S.enflata in the 0–50 m layer variedbetween 4.8 and 12.5% (mean 10.8%) and feeding activity washighest between sunrise and noon. The percentages of Copepoda,Foraminifera, crustacean larvae, Chaetognatha, Pteropoda, Ostracoda,fish and unidentified material in the gut of S.enflata were51.9,6.7,3.8,2.9,1.9,1.9 and 30.9%, respectively. Sagitta enflataconsumed food organisms which were mainly between 0.5 and 1.0mm in length. The daily feeding rate of S.enflata was 1.81 preyper individual, which was equivalent to 8.06 mg C m^–2day^–1. This corresponded to     

Predation and respiration by the small cyclopoid copepod Oithona similisr: How important is feeding on ciliates and heterotrophic flagellates?

Feeding on natural plankton populations and respiration of thesmall cyclopoid copepod Oithona similis were measured duringthe warm season in Buzzards Bay, Massachusetts, USA. AlthoughO.similis did not significantly ingest small autotrophic andheterotrophic flagellates (2–8 µn), this copepodactively fed on >10 µm particles, including autotrophic/heterotrophic(dino)flagel-lates and ciliates, with clearance rates of 0.03–0.38ml animal^–1 h^–1. The clearance rates increased withthe prey size. O.similis also fed on copepod nauplii (mainlycomposed of the N1 stage of Acartia tonsa with a clearance rateof 0.16 ml animal^–1 h^–1. Daily carbon ration fromthe combination of these food items averaged 148 ng C animal^–1day^–1 (41% of body C day^–1), with ciliates and heterotrophicdino-flagellates being the main food source ({small tilde}69%of total carbon ration). Respiration rates were 020–0.23µl O₂ animal^–1 day^–1. Assuming a respiratoryquotient of 0.8 and digestion efficiency of 0.7, the carbonrequirement for respiration was calculated to be 125–143ng C animal^–1 day^–1, close to the daily carbon rationestimated above. We conclude that predation on ciliates andheterotrophic dinoflagellates was important for O.similis tosustain its population in our study area during the warm season.     

Species composition and depth distribution of chaetognaths in a Kuroshio warm-core ring and Oyashio water

Chaetognath species composition, diversity and depth distributionwere investigated inside and outside a warm-core ring off thenorth-east coast of Japan. Time series samples were collectedat a station inside a Kuroshio warm-core ring (KWCR) and ata station in the surrounding Oyashio water. Greater mean abundanceof 2.51 ± 0.116 (mean ± SE) chaetognaths m^-3 wasfound outside the ring compared with 1.75 ± 0.321 chaetognathsm^-3 collected within the ring. However, species diversity valueswere higher within the KWCR (Shannon–Weaver index). Eukrohniahamata was the dominant species outside the ring, comprising79.8–87.9% of the total chaetognaths per haul. Inside,Sagitta minima was dominant, comprising 35.1–44.3%. MostE. hamata were collected in deeper layers within the KWCR andtheir abundance was on average only 9% of that found in theOyashio region. Only E. hamata and Sagitta scrippsae had differentdepth distributions in the KWCR. The vertical distribution ofE. hamata by body size appeared altered by the KWCR. Althoughthe mean length of E. hamata was not significantly differentbetween regions, sexual development appeared inhibited in thering. Sagitta elegans collected in the KWCR were mostly smallin size (<10 mm), significantly smaller than in the Oyashiowater.     

Biogeographic study of the planktonic communities of the Prince Edward Islands (Southern Ocean)

Microphytoplankton and zooplankton composition and distributionin the vicinity of the Prince Edward Islands and at the Sub-antarcticFront (SAF) were investigated in late austral summer (April/May)1996. Samples were collected for analysis of chlorophyll a concentration(Chi a), microphytoplankton and zooplankton abundance. Generally,the highest Chl a concentrations (up to 2.0 µg l^–1)and zooplankton densities (up to 192 ind. m^–3) were recordedat stations within the inter-island area while the lowest values(<0.4 µg l^–1) were observed at stations upstreamof the islands. High Chl a and zooplankton biomass values werealso associated with the SAF. Microphytoplankton were dominatedby chain-forming species of the genera Chaetoceros (mainly C.neglectus),Fragilariopsis spp. and the large diatom Dactyliosolen antarcticus.The zooplankton assemblages were always dominated by mesozooplanktonwhich at times contributed up to 98% of total zooplankton abundanceand up to 95% of total biomass. Among mesozooplankton, copepods,mainly Clausocalanus brevipes and Metridia lucens numericallydominated. Among the macrozooplankton euphausiids, mainly Euphausiavallentini, E.longirostis and Stylocheiron maximum, and chaetognaths(Sagitta gazellae) accounted for the bulk of abundance and biomass.Cluster and ordination analysis did not identify any distinctbiogeographic regions among either the microphytoplankton orzooplankton.     

The effects of food deprivation, prey density and volume on clearance rates and ingestion rates of Diacyclops thomasi

The ingestion rates of the copepod, Diacyclops thomasi, on thesoft-bodied rotifer, Synchaeta pectinata, increased 10-fold(0.07–0.77 Synchaeta h^–1) over the range 50–250prey l^–1. The saturating functional response curve appearedsigmoid but was statistically indistinguishable from a parabola.The response curve was more linear and 10 times lower over thesame range of density when Diacyclops was offered Kerarellacochleans, a species having a stiffened lorica. Diacyclops maximizedits ingestion rate on Synchaeta as a function of the availablegut space. Predation effort, measured as clearance rates, waslinked tentatively to changes in swimming speed of Diacyclopsand was a function of hunger level. Diacyclops, which were starvedfor varying periods of time, increased their ingestion rateson Synchaeta up to a maximum (-3.0 h^–1) after 7–10h of food deprivation. The gut passage time of Diacyclops wasestimated to be 7–8 h. Therefore, ingestion rates (andclearance rates) appeared to be strongly correlated to the volumeof food in the gut.     

Trophodynamics of the scyphomedusae Aurelia aurita. Predation rate in relation to abundance, size and type of prey organism

Ephyra larvae and small medusae (1.7–95 mm diameter, 0.01–350mg ash-free dry wt, AFDW) of the scyphozoan jellyfish Aureliaaurita were used in predation experiments with phytoplankton(the flagellate Isochrysis galbana, 4 µm diameter, {smalltilde}6 x 10^–6 µg AFDW cell^–1), ciliates (theoligotrich Strombidium sulcatum, 28 µm diameter, {smalltilde}2 x 10^–3 µg AFDW), rotifers (Synchaeta sp.,0.5 µg AFDW individual^–1) and mixed zooplankton(mainly copepods and cladocerans, 2.1–3.1 µg AFDWindividual^–1). Phytoplankton in natural concentrations(50–200 µg C I^–1) were not utilized by largemedusae (44–95 mm diameter). Ciliates in concentrationsfrom 0.5 to 50 individuals ml^"1 were consumed by ephyra larvaeand small medusae (3–14 mm diameter) at a maximum predationrate of 171 prey day^–1, corresponding to a daily rationof 0.42%. The rotifer Synchaeta sp., offered in concentrationsof 100–600 prey I^–1, resulted in daily rations ofephyra larvae (2–5 mm diameter) between 1 and 13%. Mixedzooplankton allowed the highest daily rations, usually in therange 5–40%. Large medusae (>45 mm diameter) consumedbetween 2000 and 3500 prey organisms day^"1 in prey concentrationsexceeding 100 I^–1. Predation rate and daily ration werepositively correlated with prey abundance. Seen over a broadsize spectrum, the daily ration decreased with increased medusasize. The daily rations observed in high abundance of mixedzooplankton suggest a potential ‘scope for growth’that exceeds the growth rate observed in field populations,and this, in turn, suggests that the natural populations areusually food limited. The predicted predation rate at averageprey concentrations that are characteristic of neritic environmentscannot explain the maximum growth rates observed in field populations.It is therefore suggested that exploitation of patches of preyin high abundance is an important component in the trophodynamicsof this species. ¹Present address: University of Bergen, Department of MarineBiology, N-5065 Blomsterdalen, Norway     

UMBONIUM VESTIARIUM (L.) (GASTROPODA, TROCHACEA) AS THE FOOD SOURCE FOR NATICID GASTROPODS AND A STARFISH ON A MALAYSIAN SANDY SHORE

Umbonium vestiarium (L.) forms virtually the entire diet of3 (possibly 4) species of naticid snails and the starfish Astropectenvappa Mueller annd Troschel on some north Penang sandy shores.Umbonium comprises about 99% of numbers and tissue of macrofauna.Predation totalled some 1.75 Umbonium (ca. 33 mg dry tissue).m^-2.day^-1 across much of the downshore sand flats rising to 2.3Umbonium (ca. 45 mg). m^-2. day^-1 near MLWS. Natica maculosaLamarck comprised > 80% of the predators and took 77–94%of the Umbonium eaten. Natica antonii Phillippi alone addedto this toll on the upper reaches of the zone while Polinicesspp and Astropecten appear to have taken 12–14% of thetotal toll of Umbonium near MLWS. Total predation is indicatedat 237–327 kJ.m^-2. year^-1 across the shore and this representsalmost the total flow of energy from primary consumers to intertidalbenthic predators on such shores and accounts for some 15.6%(lower shore)—20.5% (upper shore) of total Umbonium production. (Received 10 September 1982;