Photophobic responses of calanoid copepods: possible adaptive value

A genual pattern of photophobic responses has been observedwhich differs for calanoid copepods from freshwater, estuarineand oceanic environments. Using a video-computer system formotion analysis, the photophobic responses of light and darkadapted calanoid copepods were compared. Dark-adapted copepodswere exposed to 600 ms flashes of dim blue light at 5 s intervalswhich simulated the flashes of biolumines-cent marine zooplankton.Light-adapted copepods were exposed to 600 ms intervals of darknessat 5 s intervals to simulate the shadows of organisms passingoverhead. Four species of coastal marine copepods (Acartia hudsonica,Centropages hamatus, Pseudocalanus minutus and Temora longicornis)all showed photophobic responses to both flashes and shadows.These responses may have adaptive value to the copepods sincethey live in an environment with predators that are bioluminescentat night and cast shadows on their prey during the day (e.g.ctenophores and cnidarian medusae). Two species of oceanic copepods(Euchaeta marina, Pleuromamma abdominalis) showed strong photophobicresponses to flashes but no response to shadows. This may correspondto the abundance of bioluminescent predators on copepods inthe oceanic environment (fish, ctenophores, siphonophores, etc.)and their lack of exposure to the shadows of predators, sinceboth these species are rarely found in the euphoric zone duringthe day. Two species of freshwater copepods (Diaptomus sanguineus,Epishwa massachusettsensis) showed no similar photophobic responseto flashes of light. This lack of startle response may relateto the lack of bioluminescence in the freshwater environment.Freshwater copepods showed a weak photophobic response to shadows.The adaptive value of this behavior is unclear, however, sincethe responses seem to be too weak to function for escape, andthe dominant predators large enough to cast shadows (fish) tendto approach their prey laterally. ¹Present address: Marine Science Institute, University of Texasat Austin, Port Aransas, TX 78373–1267, USA     

Environmental factors influencing diurnal distribution of zooplankton and ichthyoplankton

The diurnal vertical distribution of a large number of speciesof zooplankton, icbthyoplankton and micronekton were determinedin the top 150 m in three locations in the Shelf Water, on theNova Scotia Shelf, and Slope and on Georges Bank during springand fall periods. Species were categorized as to their trophiclevel and their type of diurnal migration behaviour. The influenceof temperature, salinity, and water density on the diurnal verticaldistribution of the species was examined. Temperature was foundto have the greatest influence on the distribution of the largestnumber of species. Diurnal migration behavior of the same speciesin Shelf and Slope water and at different times of the yearwas examined. Results showed that species changed their behaviorin the two water masses, while some species changed their migrationbehavior at different times of the year. During the night inApril the most abundant copepod species, Calanus finmarchicus,making up about 80% of the biomass, was found concentrated abovethe thermocline and the main chlorophyll layer. The majorityof the less abundant species of copepods were found below thethermocline and the chlorophyll layer. At night in August thetwo most abundant copepod species, Centropoger typicus and Paracalanusparvus, making up at least 80% of the zooplankton biomass, werealso concentrated above the thermocline and the main chlorophyllLayer. Three species of copepods were concentrated at the depthof the main chlorophyll layer and two species were concentratedbelow the chlorophyll layer and thermocline. The vertical distributionof other zooplankton and ichthyoplankton species was examinedin relation to the thermocline and chlorophyll layer. Relationshipsbetween concentrations of six species of fish larvae and allspecies of copepods in the same samples showed a general increasein the numbers of larvae m^–3 as the numbers of copepodsm^–3 increased in a range of 500–4000 m^–3.However, the concentration of Merluccius bilinearis decreasedas the concentration of copepods exceeded 4000 m^–3 suggestingthat high concentrations of copepods may not be a favourableenvironment for the larvae.     

Escape behavior of Acartia hudsonica copepods during interactions with scyphomedusae

Calanoid copepods possess remarkable abilities to detect andescape from hydrodynamic disturbances, such as those createdby approaching predators. At the same time, a number of studiesin coastal ecosystems have suggested that gelatinous predators,including medusae in the Class Scyphozoa, exert top-down controlon copepod populations. Although prey escape behavior playsa critical role in predation models, we have relatively littleempirical data on how copepods respond to encounters with scyphomedusae.In this study, I used video to quantify encounter rates andescape behaviors of the copepod Acartia hudsonica during interactionswith two scyphomedusae, Aurelia aurita and Cyanea sp., in twoflow regimes. Escapes were complex, variable and effective.Fewer than 1% of encounters resulted in ingestion. Typically,A.hudsonica avoided contact by responding when predators remainedseveral body lengths (4 to 10 mm) distant and stringing togethermany escape jumps at submaximum velocities (33 to 59 mm s^–1).In addition, copepodite stages behaved passively—or failedto respond—following encounters with medusae more oftenthan did adults. Because escape behavior exhibited by A.hudsonicawas so variable, it is unlikely that medusae capture copepodsusing a single, quantifiable mechanism. A range of responseswithin populations and individuals may be the best strategyfor zooplankton faced with strong predation pressure from avariety of predators.     

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.     

Behavioral interactions between a cyclopoid copepod predator and its prey

Behavioral observations on the predatory interactions betweenMesocyclops edax and several different types and sizes of preyrevealed that prey size alone was less important than otherspecific morphological and behavioral characteristics of theprey in deterring successful predation by the copepod. The behavioralresponses of Bosmina and Asplanchna to an attacking copepodwere passive and consisted of a simple retraction of vulnerableswimming appendages which made the prey more difficult to grasp.Daphnia and Diaphanosoma on the other hand exhibited very activeswimming escape responses. Tropocyclops usually avoided M. edaxby fleeing before the larger predator could detect them. Thehard carapaces of Daphnia, Bosmina and Keratella were effectiveat reducing ingestion following capture by M. edax. The resultsof these behavioral observations were supported by enclosureexperiments in which the predator was offered a choice betweentwo prey simultaneously. Cyclopoid copepods are capable of successfullyattacking, capturing and ingesting prey organisms several timestheir own body length. Although size alone may influence thepreference of cyclopoid copepods on large and small individualsof the same or similar prey species, it is not a dependabledeterminant of the preference of cyclopoids on multispecificprey assemblages. ¹Present address: Department of Biology, Williams Hall No. 31,Lehigh University, Bethlehem, PA 18015, USA     

Feeding and metabolism of the siphonophore Sphaeronectes gracilis

The in situ predation rate of the siphonophore Sphaeronectesgracilis was estimated from gut content analysis of hand-collectedsiphonophores and from laboratory data on digestion rates ofprey organisms. At daytime prey densities of 0.25 copepods 1^–1,S. gracilis was estimated to consume 8.1 – 15.4 prey day^–1siphonophore^–1. From data on abundances of siphonophoresand copepods, S. gracilis was estimated to consume 2–4%of the copepods daily. In laboratory experiments, ingestionrates averaged 13.8 prey day^–1 siphonophore^–1 atprey densities of 5 copepods 1^–1 and 36.9 at 20 copeods1^–1. This was equivalent to a specific ingestion rate(for both carbon and nitrogen) of –17% day^–1 and45% day^–1, respectively, while specific ingestion in situwas only 2% day^–1. Ammonium excretion averaged 0.095 µg-atsiphonophore^–1 day^–1 at 5 prey 1^–1, and 0.162at 20 prey 1^–1. The specific respiration (carbon) andspecific excretion (nitrogen as ammonium) were calculated tobe 3% day^–1 at the lower experimental food level, and5% day^–1 at the higher food level. ¹Contribution from the Catalina Marine Science Center No. 66. ²Present address: Dept. of Biology, University of Victoria,Victoria, B.C., Canada V8W 2Y2.     

Effects of prey motility and concentration on feeding in Acartia tonsa and Temora longicornis: the importance of feeding modes

Feeding experiments were conducted with the ambush-feeding copepodAcartia tonsa and the feeding-current-generating copepod Temoralongicornis. The copepods were offered a mixed diet of the dinoflagellateHeterocapsa triquetra and the ciliate Balanion comatum of similarcell size. The dinoflagellate was offered at a constant concentrationof 10–15 cells mL^–1, whereas the ciliate was offeredat a variety of concentrations, ranging from 7 to 57 cells mL^–1.Copepods with different feeding modes possess different mechanismsfor prey detection, suggesting that the two copepods would responddifferently to the two prey types. Both copepods had significantlyhigher clearance rates on the highly motile ciliate than onthe less motile dinoflagellate. In encounters between A. tonsaand its prey, we argue that this is due to the higher hydromechanicalsignal generated by the ciliate. The advection feeding copepodT. longicornis fed on the two prey according to their relativeconcentrations; in this case, we suggest that although B. comatumis capable of detecting feeding-current-generating predators,the feeding current velocity generated by T. longicornis isgreater than the escape velocity of this ciliate.     

The swimming behavior of the freshwater calanoid copepods Limnocalanus macrurus Sars, Senecella calanoides Juday and Epischura lacustris Forbes

The swimming behavior of the freshwater calanoid copepods Epischuralacustris, Senecella calanoides, and Limnocalanus macrurus wasexamined using a videotape analysis technique. E. lacustrisand S. calanoides alternate between periods of slow glidingand passive sinking. L. macrurus does not exhibit sinking behavior;its swimming activity consists of slow and steady gliding interruptedoccasionally by jumps. All three copepods exhibit behavioralresponses when exposed to prey organisms, as expressed by anincrease in the duration of sinking for E. lacustris and S.calanoides, a decrease in sinking speed for S. calanoides, anda less circuitous swimming path for L. macrurus. These behavioralchanges may allow the copepods to allocate more time for preydetection, to be less detectable by potential prey and to reducethe frequency of path re-crossing. The results demonstrate therelationship between the swimming and feeding behavior of calanoidcopepods and are consistent with previous findings that copepodshave the ability to modify their foraging behavior in responseto changes in food conditions.     

Remote prey detection in Oithona similis: hydromechanical versus chemical cues

We quantified prey encounter rates and prey reaction distancesin the ambush-feeding cyclopoid copepod Oithona similis by videorecording freely swimming copepods at different concentrationsof prey, the dinoflagellate Gymnodinium dominans. Prey encounterrate increased with prey concentration, and a maximal clearancerate of 0.42 ± 0.10 ml h^–1 was estimated. The averagedistance (from the antennules) at which O.similis reacts toprey is 0.014 ± 0.007 cm. A simple prey encounter modelwas used to combine observed predator and prey velocities andprey reaction distance, and yielded a clearance rate similarto that estimated directly from prey encounter rates. The observedprey reaction distance was consistent with that estimated froma published model of hydromechanical prey perception. The possibilityof remote chemodetection was examined by modeling the distributionof solutes leaking out of a swimming cell. The cell leaves along slender chemical trail in its wake. However, since theambush-feeding O.similis is essentially stationary when perceivingprey, it is the width rather than the length of the trail thatmatters. Owing to advection, the chemical signal vanishes almostinstantaneously off the sides of the swimming flagellate, andsolute concentrations are below any likely detection thresholdwithin 40–50 µm from the flagellate. Our observationsare thus inconsistent with remote chemodetection in O.similis.The considerations are generalized, and it is concluded thatambush-feeding copepods, unlike cruisers and suspension feeders,cannot utilize chemical signals for the detection of individualprey, but rely on either hydromechanical detection or directinterception of prey.     

Effect of prey size on vulnerability of copepods to predation by the scyphomedusae Aurelia aurita and Cyanea sp.

A growing number of studies correlate changes in zooplanktonpopulations with abundance of medusae, but we cannot yet explainor predict the specific factors driving these interactions.This study demonstrates that the size of copepods has a significantinfluence on their vulnerability to predation by scyphomedusae.This finding is important because prey size, independent ofbehavior, has been neglected in theoretical models of predationby medusae. In experiments in a planktonkreisel, we used liveand heat-killed prey (Acartia hudsonica adults and copepodites)to separate the effects of copepod size and behavior on feedingrates by two medusae (Aurelia aurita and Cyanea sp.). Resultsrevealed that: differences in copepod size had a significantimpact on feeding rates, and thus small size can provide a refugefrom predation; behavior of adults diminished the liabilityassociated with larger size; and medusae with different morphologiesingested A.hudsonica at similar rates. Other experiments demonstratedthat medusae digested copepods at different rates based on preysize and predator species, findings that have implications forall future laboratory and field studies that assess feedingby scyphomedusae. Finally, this study illustrates how laboratorystudies serve as critical supplements to field observations.The effect of prey size on feeding rates can be confounded bydifferences in prey behavior, yet explains why small copepodswere typically ingested at relatively low rates by medusae.Size was clearly a dominant factor influencing copepod vulnerabilityacross scyphomedusan species, even those with very differentmorphologies. Future work should focus on the mechanisms ofsize selection, or the factors influencing contact and retentionrates.     

The effect of container size on the feeding rate of Heterocope septentrionalis, a freshwater predaceous copepod

The feeding rates of freshwater predaceous copepods have beenmeasured many times using enclosure experiments. Typically theseexperiments involve enclosing a known number of zooplanklonprey with a known number of predaceous copepods and determiningthe number of prey remaining after a set length of time. Variousfactors such as prey species and size have been shown to influencethe feeding rate of copepods. However, little attention hasbeen paid to the influence that container size may have on feedingrates. Using previously published data and some new data itwas found that container size has a major impact on the feedingrate of Heterocope septentrionalis, a predaceous freshwatercopepod common in North American arctic waters. In experimentswith Daphnia puiex, where container size varied from 0.3 to54 l (180 times), the measured feeding rate coefficient variedfrom –0.04 to –1.7 (42.5 times). Other prey speciesshowed similar changes, but the vulnerability of particularprey species to Heterocope predation remained consistent acrossthis range of expenmental containers. The reason for the containereffect is not known, but it is postulated that edge avoidanceby Heterocope may account for the observed change in feedingrate.     

Mesozooplankton grazing under conditions of extreme eutrophication in Guanabara Bay, Brazil

The objective of the present study was to quantify mesozooplanktongrazing in the eutrophic waters of Guanabara Bay. Mesozooplankton(>200 µm) was dominated by the copepods Acartia lilljeborgi,Acartia tonsa, Parvocalanus crassirostris and Paracalanus furcatus.Dinoflagellates, specifically the species Prorocentrum triestinum,were an important group for mesozooplankton nutrition, beingingested in significant amounts during all experiments. On average,12.3 ± 2.9 P. triestinum cells were ingested copepod^–1min^–1 (other dinoflagellates: 11 ± 8 cells copepod^–1min^–1). Filamentous cyanophyceae and nanoplankton wereingested in one experiment each, but the mesozooplankton communitygenerally preferred dinoflagellates to these groups, which werealways abundant in the water column. Euglenophyceae were notingested, although they dominated in one experiment. Mesozooplanktoningested, on average, only 0.2% of the nano- and microplanktonbiomass per day. The results suggest that grazing was not acontrolling process for the nano- and microplankton communityin the study area. Addition of zoeae larvae of Chasmagnatusgranulata (Decapoda: Brachyura: Grapsidae) in one experimenthad a significant effect on the mortality of adult copepods,probably due to a predator–prey relationship.     

The role of feeding behavior in sustaining copepod populations in the tropical ocean

A fundamental question regarding marine copepods is how themany species coexist and persist in the oligotrophic environment(i.e. Hutchinson’s paradox). This question is addressedwith a stochastic, object-oriented Lagrangian model that explicitlysimulates the distinct foraging behaviors of three prominenttropical species: Clausocalanus furcatus, Paracalanus aculeatusand Oithona plumifera. The model also individually tracks allprey cells. Each particle’s motion combines sinking, turbulentdiffusion and active swimming when applicable. The model successfullysimulates observed size-partitioned carbon uptake rates. Basedon the model results, the wide-ranging translational ambit employedby C. furcatus is best suited for the acquisition of passiveprey while the relatively stationary behavior of O. plumiferapromotes the capture of larger, quickly sinking cells. The modelresults further suggest that the slow velocities and feedingcurrent employed by P. aculeatus are best suited for acquiringthe smallest cells though it also has a slight advantage overC. furcatus in acquiring the largest prey. A resource threshold,at a prey concentration of 530 cells mL^–1, is consistentlyexhibited by all three modeled species. Overall, these resultsimply that the size-partition preferences due to their differentforaging behavior contribute to the coexistence of these threespecies. This paper is one of six on the subject of the role of zooplanktonpredator–prey interactions in structuring plankton communities.     

The feeding strategies of two large marine copepods

We compared the feeding behaviour of the two copepods Paraeuchaetanorvegica and Chiridius armatus, allowing them to prey on othercalanoids in small-scale laboratory experiments. Several differenceswere found. When fed either live, free-swimming or dead, non-movingprey, P.norvegica seemed unable to locate the dead prey itemswhile C.armatus foraged heavily on them. When starved, P.norvegicaincreased its feeding rate while C.armatus reduced its feedingafter an initial increase. None of the predators changed theirfeeding rates when exposed to light. Nighttime versus daytimefeeding was tested only with P.norvegcia, which seemed to possessan endogenous feeding rhythm with increased rates at night.Chiridius armatus infected with epizooic ciliates appeared tohave increased feeding rates.     

From polar night to midnight sun: photoperiod, seal predation, and the diel vertical migrations of polar cod (Boreogadus saida) under landfast ice in the Arctic Ocean

The winter/spring vertical distributions of polar cod, copepods, and ringed seal were monitored at a 230-m station in ice-covered Franklin Bay. In daytime, polar cod of all sizes (7–95 g) formed a dense aggregation in the deep inverse thermocline (160–230 m, −1.0 to 0°C). From December (polar night) to April (18-h daylight), small polar cod <25 g migrated into the isothermal cold intermediate layer (90–150 m, −1.4°C) at night to avoid visual predation by shallow-diving immature seals. By contrast, large polar cod (25–95 g), with large livers, remained below 180 m at all times, presumably to minimize predation by deep-diving mature seals. The diel vertical migration (DVM) of small polar cod was precisely synchronized with the light/dark cycle and its duration tracked the seasonal lengthening of the photoperiod. The DVM stopped in May coincident with the midnight sun and increased schooling and feeding. We propose that foraging interference and a limited prey supply in the deep aggregation drove the upward re-distribution of small polar cod at night. The bioluminescent copepod Metridia longa could have provided the light needed by polar cod to feed on copepods in the deep aphotic layers.     

Capelin larvae (Mallotus villosus) and community structure of zooplankton off the coast of northern Norway

Capelin larvae (Mallotus villosus) and mesozooplankton werecollected in May 2001 in the southern part of the Barents Sea.During the period of hatching and first feeding, capelin larvaeexperience high mortality rates. In order to understand theunderlying mechanisms influencing larval survival, we monitoreda selected site for 10 days using a variety of different samplingapproaches. Hydrological parameters conductivity–temperature–depth–fluorescence(CTDF) were recorded and zooplankton samples obtained simultaneouslyusing MOCNESS at three time periods. The mesozooplankton consistedessentially of copepods (85), dominated by Calanus finmarchicus.The zooplankton community consisted of different assemblagesderived from different areas (i.e. different water masses; Atlanticand coastal). Capelin larvae were found in surficial coastalwaters. Copepod nauplii and C. finmarchicus were less abundantin coastal waters where small copepodites (such as Acartia spp.and Temora longicornis) were found together with early stagesof coastal invertebrates that are possible prey for capelinlarvae. We suggest that capelin larvae base their foraging onsmall prey organisms frequently found among the coastal zooplanktonand that these larvae are less dependent on the propagationof the recruiting generation of C. finmarchicus present fromApril to June.     

Response of two zooptankton grazers to an ichthyotoxic estuarine dinoflagellate

The dinoflagellate Pfiesteria piscicida (gen. et sp. nov.).a toxic ‘ambush predator’, has been implicated asa causative agent of major fish kills in estuanne ecosystemsof the southeastern USA. Here we report the first experimentaltests of interactions between P.piscicida and estuarine zooplanktonpredators. specifically the rotifer Brachionus plicatilis andthe calanoid copepod Acartia tonsa. Short-term (10 day) exposureof adult B.plicatilis to P.piscicida as a food resource, aloneor in combination with the non-toxic green algae Nannochlorisand Tetraselmis. did not increase rotifer mortality relativeto animals that were given only non-toxic greens Similarly,short-term (3 day) feeding trials using adult A.tonsa indicatedthat the copepods survived equally well on either P.piscicidaor the non-toxic diatom Thalassiosira pseudonana. Copepods giventoxic dinoflagellates exhibited erratic behavior, however, relativeto animals given diatom prey. The fecundity of B.plicatiliswhen fed the toxic dinoflagellate was comparable to or higherthan that of rotifers fed only non-toxic greens We concludethat, on a short-term basis, toxic stages of P.piscicida canbe readily utilized as a nutritional resource by these commonestuarine zooplankters. More long-term effects of P.piscicidaon zooplankton, the potential for toxin bioaccumulation acrosstrophic levels, and the utility of zooplankton as biologicalcontrol agents for this toxic dinoflagellate. remain importantunanswered questions.     

Diel variations in gut pigment content, diel vertical migration and estimates of grazing impact for copepods in the southern Benguela upwelling region in October 1987

The gut fluorescence technique was used to estimate ingestionand filtration rates of the adult female copepods Paracalanusparvus, Cenlropages brachiatus and Calanus austrails, and copepoditestages 3, 4 and 5 of C.australis in the southern Benguela upwellingregion. During the study period chlorophyll concentrations withinthe upper 20 m of the water column were high, 5 µg I^–1in mid-shelf waters and 15–30 µg I^–1 in innershelf waters. Copepod gut pigment content was low and constantduring the day then increased sharply during the first 2 h aftersunset. Gut pigment content was 2–6 times higher duringthe night compared with daytime values. Small non-migratingcopepods (Paracalanus parvus) showed the smallest diel differencein gut pigment content and large migrating copepods (Centropagesbrachiatus and Calanus australis) the largest difference. Eggproduction rates were 20 and 50% of maximum at the mid-shelfand inner shelf stations respectively, suggesting food-limitation.Comparison of ingestion rates calculated from egg productiondata with ingestion rates calculated from gut pigment data suggestedthat the copepods were feeding omnivorously at the inner shelfstations but herbivorously at the mid-shelf stations. Assumingthat all of the phytoplankton was available as food, the nearshorecopepod assemblage grazed {small tilde}1% of the standing cropeach day, and the mid-shelf assemblage grazed 5% day^–1.Because of errors and uncertainties associated with the gutfluorescence technique, the feeding impact could be underestimatedby 2–4-fold. We discuss several approaches which couldlead to more precise estimates of feeding rates. ³Present address: Marine Sciences, SUNY, Stony Brook, NY, 11794-5000,USA     

Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions

Prey species must adapt their behavior to avoid predation. Asa key prey item for lions (Panthera leo), plains zebras (Equusburchelli) were expected to respond to immediate threats posedby lions in their area. In addition, zebras were predicted toexhibit behavior tuned to reduce the potential for encounterswith lions, by modifying their movement patterns in the timesof day and habitats of greatest lion danger. We studied a populationof approximately 600 plains zebra living in Ol Pejeta Conservancy,Kenya. We found that zebra abundance on or near a grasslandpatch was lower if lions had also been observed on that patchduring the same day. Predation danger was highest in grasslandhabitat during the night, when lions were more active. Zebrasightings and global positioning system radio collar data indicatedthat zebras also reduced their use of grassland at night, insteadusing more woodland habitat. Zebras moved faster and took sharperturns in grassland at night. It is hypothesized that these moreerratic movements assist zebras in avoiding detection or captureby lions.     

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.