The notal organ of the scorpionfly (Panorpa vulgaris): an adaptation to coerce mating duration

The notal organ in P. vulgaris (Insecta: Mecoptera) is a clamplikestructure on the dorsum of the middle of the male’s abdomenthat holds one of the female’s forewings throughout mating.Males often provide salivary masses as food to their mates duringmating, and a male’s ability to produce saliva dependsupon whether he has fed adequately before mating. To test thehypothesis that the evolutionary function of the notal organis to coerce longer matings than are in the interests of females,the notal organ and the amount of food males and females receivebefore mating were experimentally manipulated. In support ofthe hypothesis, copulation was reduced when the notal organwas made inoperative under the following four conditions: (1)female fed and male starved before mating, no saliva duringmating; (2) same as in condition 1 but with one salivary massduring mating; (3) both sexes starved before mating, no salivarymass during mating; and (4) both sexes fed before mating. However,when females were starved before mating and males were fed,resulting in multiple masses being provided during mating, thenotal organ had no effect on length of copulation, and thusthere was no sexual conflict over mating duration. In addition,the larger the female relative to her mate, the briefer thecopulation when the notal organ was operative, which suggeststhat a physical struggle between the male and female may occurduring sexual conflict about mating duration. This is one ofonly a few studies that provide evidence of adaptation to thedomain of sexual coercion. [Behav Ecol 1991;2:156–164]     

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.     

FIELD OBSERVATIONS ON FEEDING OF THE LAND SNAIL HELIX ASPERSA MULLER

Feeding of the land snail Helix aspersa (Müller) was observedat monthly intervals. Three natural populations in Galicia (NW-Spain)were studied. At two sites only a few plants constituted thebulk of the diet and in spring the snails' diet had the highestdiversity (H'). In the third population feeding and distributionof Helix aspersa (Müller) were observed in a small plotwith permanent patches of Urtica dioica. Nearly one half offeeding snails fed upon Urtica dioica. Most of the other observations wereon Mentha suaveolens, Ranunculus repens and Gramineae. The diversity ofthe snails' diet showed seasonal variation with the maximumin the autumn months. Comparison between the availability ofthe different plant species and their contribution to the snails'diet showed that the snails did not eat at random; Urtica dioicawas eaten much more than expected from its occurrence and grasseswere strongly under-represented in the snails' diet. Temporalchanges of availability were significantly correlated with the amountseaten in the case of Urtica, but not for the other food plants.The distribution of the snails in the plot was significantlycorrelated with that of Urtica. Chemical analyses of the foodplants revealed Urtica as the species with the higher protein,ash and calcium contents. The strong preference of Helix aspersafor Urtica dioica could be explained by the value of Urticaas food or by its suitability as habitat for the snails. The largestproportions of green material in the snails' diet occurred inthe spring and juveniles ate more green material than adultsin the three populations. (Received 16 March 1998; accepted 30 November 1998)     

POPULATION DENSITY EFFECTS ON GROWTH IN CULTURE OF THE EDIBLE SNAIL HELIX ASPERSA VAR. MAXIMA

The effects of population density on the growth of H. aspersaMÜller var. maxima under controlled environmental conditionswere examined. Inhibitory effects on snail growth and maturityresulting from increased population density, between 100–800snails m^–2 of floor area, were observed for a range ofcontainer cleaning frequencies. At all population densities,enhanced snail growth was observed when the frequency of containercleaning was increased to a two-day interval. No significantdifferences were recorded, following 19 weeks growth, betweenfinal mean weights of snails from containers cleaned less frequently.The lowest snail mortality was consistently recorded at thelowest population density in the most frequently cleaned containers.At all snail population densities three phases of growth wereobserved: (a) lag (0–5 weeks), (b) rapid (6–15 weeks)and (c) stable (16 weeks and over). During the first three weeksof growth, high population density had a positive effect ondiet consumption, food conversion efficiency and snail growth.Adverse population density effects increased progressively duringphase (b), typically following 9 weeks growth. Juvenile snailstransferred from high to low population densities during phase(b) continued to exhibit slower growth rates associated withhigh population densities. Food conversion efficiency of snailsin all treatments decreased throughout the experimental periodbut with no overall effect of container cleaning frequency apparent.Inherent growth variability of sibling snails was unaffectedby population density or container cleaning frequency. The importanceof the results for intensive snail culture is discussed. (Received 23 June 1994; accepted 1 December 1994)     

THE INFLUENCE OF INGESTIVE CONDITIONING ON FOOD CHOICES IN THE LAND SNAIL HELIX ASPERSAMULLER (GASTROPODA: PULMONATA: STYLOMMATOPHORA)

An experiment of food choice in one-day-old naïve landsnail Helix aspersa Müller was carried out, involving threephases. First, a training period on a monophagous diet of leafdiscs of Taraxacum officinale or Urtica dioica was given for1, 5, 10, 15 or 30 days. The area of leaf discs consumed byeach snail during this stage was assessed. In the intermediatephase, the animals were not fed for 24hours. Finally, food choicewas tested over 24 hours, when animals could eat leaf discsof each species of plant usedfor training. A coefficient offood preference for Urtica dioica was calculated (UPC = areaof Vrtica leaf disc consumed/area of Urtica and Taraxacum discsof leaves consumed for 24 hours). The length of the training phase did not reduce the percentageof individuals preferring the plant previously consumed. Onthe contrary, after 15 days of monophagous diet, animals formerlyfed on Urtica showed a coefficient of food preference for Urticasignificantly higher than Taraxacum fed animals, and after 30days of training this phenomenon was more pronounced. Thus,during the choice test, snails ate more of the previously eatenplant leaf. Our experiment highlighted the feeding behaviourplasticity of H. aspersa. The significant correlation betweenthe area of Urtica eaten during the training phase and the coefficientof food preference for Urtica is discussed. (Received 10 April 1994; accepted 10 January 1995)     

GROWTH OF JUVENILE BITHYNIA TENTACULATA (PROSOBRANCHIA, BITHYNIIDAE) UNDER DIFFERENT FOOD REGIMES: A LONG-TERM LABORATORY STUDY

The growth of juvenile Bithynia tentaculata (Proso-branchia,Bithyniidae) was measured under controlled laboratory conditionsover 18 months. The animals were fed with different concentrationsof suspended food (Chlamydomonas reinhardii at 2, 000 cells.ml^–1and at 10, 000 cells.ml^–1), solid food (lettuce) and combinationsof both (lettuce with 2, 000 cells.ml^–1 and with 10, 000cells.ml^–1 Chlamydomonas). The growth of all animals wasmeasured weekly, and after 1 years final shell sizes, shelldry weights and ash-free dry weights, as well as soft body dryweights and ash-free dry weights were determined. Survival rateof the animals increased from 20% when fed with 2, 000 algalcells.ml^–1 to 75% with lettuce and 10, 000 cells.ml^–1as food. Final shell sizes and shell weights were not influencedby food, but soft body dry weights were significantly reducedwhen animals were fed on suspended food only. Reproduction (i.e.egg laying) was observed at the end of the second summer whenlettuce (with or without algal addition) was given as food.The ecological consequences of these results are discussed. (Received 7 April 1993; accepted 9 August 1994)     

DISTRIBUTION AND PRODUCTION OF THREE HYDROBIA SPECIES (GASTROPODA: HYDROBIIDAE) IN A SHALLOW COASTAL LAGOON IN THE BAY OF CADIZ, SPAIN

The abundance, life span, growth and production of the mud snailsHydrobia minoricensis, H. ulvae and H. ventrosa in a semi-naturallagoon system were studied by taking monthly samples at threesites during 1991 and 1992. The most abundant species, H. minoricensisoccurred at mean densities of 12834 to 26264 snails m^–2(10.7 to 25.8g dry weigh m^–2), depending on the site.The least abundant species, H. ulvae, occurred at mean densitiesof 185 to 353 snails m^–2 (3.2 to 2.2g dry weight m^–2).The numerical abundance and biomass of the three Hydrobia specieswere positively related to the biomass of benthic macroalgae(P<0.01). Although H. ulvae egg capsules were recorded throughoutthe year, newly hatched snailsof this species were not observed,in contrast to the other two species. The early spring and summercohorts of H. minoricensis and H. ventrosa seemed to be themost numerous. The average life spans of these two species wereestimated to be about 18 and 13 months respectively. Annualproduction estimates for the whole lagoon system were 29.0 (6.3),5.5 (0.8) and 5.2 (1.0)g dry weight (ash-free dry weight) m^–2yr^–1 for H. minoricensis, H. ulvae and H. ventrosa respectively.The annual P/B ratio was about 2 for H. minoricensis and H.ventrosa. (Received 5 July 1994; accepted 5 October 1994)     

The fate of the carotenoid pigment fucoxanthin during passage through the copepod gut: pigment recovery as a function of copepod species, season and food concentration

The amount of fucoxanthin, a taxonomically diagnostic carotenoid,recovered after passage through the guts of the copepods Acartiacalifomiensis and Calanus pacificus, was determined after thecopepods had fed on low (50 µg Cl^–1) and high (350µg C1^–1 for Acartia; 500 ug C H for Calanus) concentrationsof the diatom Thalassiosira weissftogii, during spring (May)and winter (December). Changes in pigment concentrations andcell abundances were assessed in experimental (with copepods)and control (without copepods) samples by standard incubationexperiments. Pigment recovery was assessed by (i) comparingthe amount of ingested pigment recovered in the experimentalgroups with that predicted to have been ingested from cell countdata and (ii) comparing fuco-xanthin/cell ratios in controland experimental samples. Both techniques suggested that pigmentloss is substantial (usually 60–100%), regardless of species,food availability or season. Patterns of pigment conservationdiffered between species, although pigment recovery was alwayshigher at high, than at low, food concentrations. Pigment recoveryin Acartia was higher (9.4–28.0%) in the spring than duringthe winter (0 recovery), regardless of food concentration. InCalanus, however, pigment recovery was always higher at high(34.9–67.8%) than at low (0 recovery) food concentrations,regardless of season.     

WATER EXCHANGE, TEMPERATURE TOLERANCE, OXYGEN CONSUMPTION AND ACTIVITY OF THE NAMIB DESERT SNAIL, TRIGONEPHRUS SP.

Water exchange, temperature tolerance and oxygen consumptionof the snail, Trigonephrus sp., from the southern Namib desertof Namibia were examined and related to activity. At 25°Cand 15% R.H. mean water loss and food and water uptake were5.95 mg. day^–1 and 630 mg.day^–1, respectively. Bodytemperature tracked sand temperature. Snails tolerated sandtemperatures as high as 45°C. Mean ± S.D. oxygenconsumption rates were 32.0 ± 2.94 µlO₂.g totalbody mass^–1.h^–1 at 15°C, when the snails wereactive, and 11.27 µlO₂.g total body mass^–1.h^–1at 25°C, when the snails were inactive. These values are2-6 times lower than those recorded for the similarly sizedmesic snail, Helix aspersa. Activity experiments indicated thatlow ambient temperatures and high humidities were favoured bythe snails. This, together with the burying behaviour of thesesnails during high temperatures, suggests that they limit stressby restricting activity to physiologically-favourable periods,even though more-extreme conditions may be tolerated. (Received 7 June 1990; accepted 20 November 1990)     

Feeding selectivities and food niche separation of Acartia clausi, Penilia avirostris (Crustacea) and Doliolum denticulatum (Thaliacea) in Blanes Bay (Catalan Sea, NW Mediterranean)

Selectivity-size spectra, clearance and ingestion rates andassimilation efficiencies of Acartia clausi (Copepoda), Peniliaavirostris (Cladocera) and Doliolum denticulatum (Doliolida)from Blanes Bay (Catalan Sea, NW Mediterranean) were evaluatedin grazing experiments over a wide range of food concentrations(0.02–8.8 mm³ L^–1 plankton assemblages from BlanesBay, grown in mesocosms at different nutrient levels). Acartiaclausi reached the highest grazing coefficients for large algae>70 µm (longest linear extension), P. avirostris forintermediate food sizes between 15 and 70 µm, and D. denticulatumfor small sizes from 2.5 to 15 µm. Penilia avirostrisand D. denticulatum acted as passive filter-feeders. Acartiaclausi gave some evidence for a supplementary raptorial feedingmode. Effective food concentration (EFC) decreased linearlywith increasing nutrient enrichment for D. denticulatum andfollowed domed curves for A. clausi and for P. avirostris withmaximum values at intermediate and high enrichment levels, respectively.Clearance rates of crustacean species showed curvilinear responseswith narrow modal ranges to increasing food concentration. Clearancerates of D. denticulatum increased abruptly and levelled intoa plateau at low food concentrations. Mean clearance rates were13.9, 25.5 and 64.1 mL ind.^–1 day^–1, respectively.No clearance could be detected for A. clausi at food concentrations<0.1 mm³ L^–1 and for P. avirostris at food concentrations     

Seasonal patterns of food limitation in Daphnia galeata: separating food quantity and food quality effects

Food limitation for the cladoceran Daphnia galeta was measuredgrowing them with natural lake seston for one growing seasonunder standardized conditions in the laboratory. Growth rateswere related to several measures of food quantity. Particulateorganic carbon (POC) <30 µm was the best predictorof total food availability measured. It was better than chlorophyll-a(Chl-a) of the same size fraction, algal volume calculated fromcell counts or total particle volume from Coulter Counter measurements.Daphnids were also grown with Scenedesmus acutus under the sameconditions and their growth rates were compared. Sestonic carbonconcentrations were related to Scenedesmus carbon concentrations,that provided equal growth rates. The ratio of the carbon concentrationsof both food types (seston, Scenedesmus) represented the foodquality of seston. Thus, the nutritional quality of naturallake seston relative to a standard alga was determined overone growing season. Growth rates of seston fed animals revealedthat they were food limited during long periods of the year.This was also ascertained by standard clutch sizes of fieldanimals. However, the causes of food limitation varied withinthe duration of the studied period. Whereas during the clear-waterphase the food quantity was limiting (while the quality washigh: 100% Scenedesmus), shortly after on 10 June and 17 Junefood conditions were close to threshold concentrations due tolow food quality. For the remainder of the summer, a POC of0.3–0.6 mg C I^–1 with a quality of     

Prey size, prey nutrition, and food handling by shrews of different body sizes

We tested some predictions relating metabolic constraints offoraging behavior and prey selection by comparing food handlingand utilization in four sympatric shrew species: Sorex minutus(mean body mass = 3.0 g), S. araneus (8.0 g), Neomys anomalus(10.0 g), and N. fodiens (14.4 g). Live fly larvae, mealwormlarvae, and aquatic arthropods were offered to shrews as smallprey (body mass <0.1 g). Live earthworms, snails, and smallfish were offered as large prey (>0.3 g). The larvae werethe high-nutrition food (>8 kJ/g), and the other prey werethe low-nutrition food (<4 kJ/g). The smallest shrew, S.minutus, utilized (ate + hoarded) <30% of offered food,and the other species utilized >48% of food. The largerthe shrew, the more prey it ate per capita. However, highlyenergetic insect larvae composed 75% of food utilized by S.minutus and only >40% of the food utilized by the other species. Thus, inverse relationships appeared between shrewbody mass and mass-specific food mass utilization and betweenshrew body mass and mass-specific food energy utilization:the largest shrew, N. fodiens, utilized the least food massand the least energy quantity per 1 g of its body mass. Also,the proportion of food hoarded by shrews decreased with increase in size of shrew. With the exception of S. araneus, the sizeof prey hoarded by the shrews was significantly larger thanthe size of prey eaten. Tiny S. minutus hoarded and ate smallerprey items than the other shrews, and large N. fodiens hoardedlarger prey than the other shrews.     

GROWTH, MORTALITY AND FECUNDITY IN SUCCESSIVE GENERATIONS OF HEIX ASERSA MULLER CULTURED INDOORS AND CROWDING EFFECTS ON FAST-, MEDIUM- AND SLOW-GROWING SNAILS OF THE SAME CLUTCH

This paper examines the optimum conditions for edible snailsHelix aspersa to be cultured indoors successfully in successivegenerations (originating from the crossing of snails comingfrom different clutches of a previous generation), and the effectof crowding on growth and reproduction in fast-, medium-, andslow-growing snails coming from the same clutches. The timeneeded for the snails to reach marketable size (25–32mm)varied from 2.5 to 5 months up to the 7th generation. The timeneeded for the snails to mature and reproduce from 4 to 7 monthsuntil the fifth generation. After the F5 x F5 generation, thefinal size of the snails decreased. The number of eggs did notdiffer statistically among the different generations but thereproductive success (how many snails reproduced/cage) increasedfrom Fl = Fl generation onwards to F5 x F5. In F6 x F6 onlythree (out of 26) snails reproduced and in F7 x F7 none, althoughthe snails remained under controlled conditions for 15 moremonths. Mortality in the different generations varied from 0–10%up to F5 x F5 but from F6 x F6 onwards increased and reached25%. Concerning the origin of snails, it was found that largersnails (originating from Kyparissia, Peloponnesos) lay statisticallymore eggs (138.40 ± 29.60, N =5) than smaller ones (77.38± 40.42, N=4) (originating from Hania, island of Crete).Hatching success was greater, too. (Received 10 September 1996; accepted 24 March 1997)     

FORAGING BEHAVIOUR OF TERRESTRIAL GASTROPODS: INTEGRATING FIELD AND LABORATORY STUDIES

The foraging range of adult snails, Helix aspersa, has beenstudied using radio-transmitters. Snails did not return to anexact roost site after foraging, but often remained in the foodpatch, or returned only to the general roost area. Time lapse video films of the slug Deroceras reticulatum madeunder infra-red lighting in arenas were analysed for tracklengthsand degree of turning, in order to simulate slug movements inan unbounded situation. The results suggest that many food itemsare found by random encounter. Slugs usually ate the first food item found, but often ignoredfood items encountered later. If food was scarce, the slugsfed almost every time. Electronic recordings of bites on a wheatflour pellet over 24 hours show that feeding is most intensein the first two and a half hours from starting to feed, andlater meals are both shorter and less regular. Starved slugsdiffered from fed slugs principally by taking a second mealshortly after the first. When given a choice of a more preferredfood (maize pellets) and a less preferred food (pea pellets)in different ratios, the slugs appeared to encounter pelletsat random, but they fed more from the preferred pellets unlessthe ratio was 1 maize: 7 pea. Starved slugs ate twice as muchas fed ones.     

ASPECTS OF PREDATION BY TONNA ZONATUM (PROSOBRANCHIA: TONNOIDEA) FEEDING ON HOLOTHURIANS IN HONG KONG

A laboratory-held specimen of Tonna zonatum was fed holothurianprey over a period of 21 weeks. At first, the Tonna consumedHolothuria cinerascens, H. insignis and H. fuscocinerea, butsubsequently consumed only H. leucospilota. Prey size chosenwas in the range 54–200 g total weight with approximately1 holothurian being eaten each week by extension of the proboscis.Consumption time was 2–5 minutes and records of rejectionof living prey showed that food is not cut up in the proboscisby the jaws or the radula but is engulfed whole. Holothurians reacted violently to the hunting Tonna, typicallyby taking in large quantities of water to increase in size,particularly in diameter. Holothuria fuscocinerea ‘fled’the predator while H. scabra and H. leucospilota became buoyantand, writhing, effected simple ‘swimming’. Calculations of consumption show that this adult specimen consumed2% (wet weight) or 1% (dry weight) of its body weight. day^–1.This is much lower than figures for other predatory gastropods,i.e. 5%. day^–1. Analysis of faeces production, correlatedwith consumption, suggests a residence time in the gut of 4days and an assimilation efficiency of 80.02%. This study andliterature data suggest that the Tonnidac are highly specialized,exclusive, predators of holothurians. (Received 16 November 1989; accepted 22 January 1990)     

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     

Diet-shifts and food-dependent survival in Engraulicypris sardella (Cyprinidae) larvae from Lake Malawi, Africa

The diet of Engraulicypris sardella (Cyprinidae) larvae wasdeterinmed from the open waters of Lake Malawi, Africa. Theguts of first-feeding larvae of 2–3 mm total length (TL)usually contained many cells of 5–9 µm diametertentatively identified as a non-colonial green alga (Chlorophyta).The number of these cells in the guts of larvae declined aslarvae increased in size, and were not found in larvae greaterthan 9 mm TL. Other types of phytoplankton were rarely seenin the guts of larvae. Copepod nauplii were eaten by larvaegreater than 4 mm TL, and copepodite copepods and cladoceansby larvae greater than 5 mm TL. The biomass of open water crustaceanzooplankton and E.sardella larvae were determined over a 2-yearsampling programme. The mortality rate of E. sardella was negativelycorrelated with zooplankton biomass, but was not significantlycorrelated with the amount of zooplankton food in the guts oflarvae.     

EFFECTS OF HEAVY METALS ON SURVIVAL AND GROWTH OF BIOMPHALARIA GLABRATA SAY (GASTROPODA: PULMONATA) AND INTERACTION WITH SCHISTOSOME INFECTION

The effects of dissolved Pb, Cd and Hg on growth and survivalof adult Biomphalaria glabrata Say. uninfected and infectedwith Schistosoma mansoni Sambon, were examined. Pb at concentrationsfrom 0.25–100 µM, Cd from 0.075–0.25 µMand Hg from 0.25–1 µM significantly reduced growthand survival of normal snails. With each metal the effects increasedwith increasing concentration. The LC 25 at 2 wks exposure was82 ± 19, 0.22 ± 0.04 and 0.94 ± 0.13 µM(x ± S.E.) for Pb, Cd and Hg, respectively. Snails exposedto heavy metals continued to be reproductively active, but theegg production was highly variable and no significant effectof heavy metal exposure was demonstrated. Infection also decreasedsurvival and growth of snail hosts and infected individualsexposed to heavy metals displayed the greatest mortality. TheLC 25 for infected snails at 2 wks exposure was 8 ± 3,0.9 ± 0.02 and 0.29 ± 0.08 for Pb, Cd and Hg respectively.A significant interaction between heavy metal exposure and infectionwas apparent. Infected snails were not reproductively active.Cercarial shedding by infected snails was significantly reducedin the presence of heavy metals and by 6 wks shedding had ceasedat the highest metal concentrations. (Received 20 May 1996; accepted 31 July 1996)     

Dynamics of herbivorous grazing by the heterotrophic dinoflagellate oxyrrhis marina

In a series of batch experiments in the dark the heterotrophicdinoflagellate Oxyrrhis marina grazed three phytoplankton prey(Phaeodactylun tricornutum, Isochrysis galbana and Dunaliellateriolecta) with equal efficiency. Growth rates of the dinoflagellateranged between 0.8 and 1.3 day–1 Maximum observed ingestionrates on a cell basis varied according to the size of the preyfrom about 50 cells flagellate^–1 day^–1 when D.tertiolectawas the prey to 250–350 cells fiagellate^–1 day^–1when the other species were eaten. However, when compared ona nitrogen basis, ingestion rates were independent of prey type.Both ingestion and growth ceased when prey cell concentrationsfell below a threshold concentration of about 10⁵ cells ml^–1.Maximum specific clearance rates were 0.8x10⁴0ndash;5.7x10⁴it day which is considerably lower than that found for heterotrophicdinoflagellates in oceanic waters and may explain why O.marinagenerally thrives only in productive waters. The timing of NHregeneration was linked to the C:N ratio of the prey at thestart of grazing. Regeneration efficiencies for NH₄. never exceeded7%; during the exponential phase and were 45% well into thestationary phase. These results are comparable to those obtainedwith heterotrophic flagellates and demonstrate that the bioenergeticpatterns of grazing and nutrient cycling by different protozoaare very similar. Moreover, they support the notion that toachieve 90+% nutrient regeneration in the open ocean, as iscurrently believed, the microbial food loop must consist ofmultiple feeding steps. Alternatively, nutrient regenerationefficiencies may be considerably lower than 90%.     

FACTORS CONTRIBUTING TO SPATIAL HETEROGENEITY IN THE ABUNDANCE OF THE COMMON PERIWINKLE LITTORINA LITTOREA (L.)

Although Littorina littorea (L.) exhibits a relatively consistentpattern of vertical distribution throughout the North Atlantic,ranging from the mid-intertidal to the shallow subtidal zone,its horizontal distribution and abundance are highly variable.In this study, we first described the snail's horizontal distributionpatterns on Appledore Island, ME, USA and then asked whetherwave exposure, rugosity, predator density (e.g. Carcinus maenasand Cancer borealis), the percentage of the substrate coveredby Ascophyllum nodosum, Chondrus crispus, Polysiphonia spp.,and ephemeral green algae, or the percentage of uncovered substrate(bare rock) were correlated with L. littorea abundance in theintertidal zone (0.6 to 0.0 m relative to Mean Lower LowWater [MLLW]) and the shallow subtidal zone (–1.5 to –3.0 mMLLW) at nine study sites. Intertidal densities of L. littoreawere highly variable across sites, ranging from 0 to >600 m^–2.In this zone, L. littorea density showed a significant positivecorrelation with rugosity and percentage of bare rock. Densitieswere very low in the subtidal zone (range: 0–19 m^–2)and varied little among sites. Exploratory multiple regressionanalysis suggested that L. littorea density was positively correlatedwith the density of C. maenas in the shallow subtidal zone.Additionally, snails in the subtidal zone had thicker shellsthan snails of the same size in the intertidal zone. Our resultssuggest that structural elements of the habitat, such as rugosityand percentage of uncovered substrate, are among the most importantpredictors of L. littorea abundance on moderately protected,rocky intertidal shores. (Received 9 February 2005; accepted 10 August 2005)