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The Swarming Behavior of the Copepod Dioithona Oculata: in Situ and Laboratory Studies Author(S): Edward J

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The Swarming Behavior of the Copepod Dioithona Oculata: in Situ and Laboratory Studies Author(S): Edward J The Swarming Behavior of the Copepod Dioithona oculata: In Situ and Laboratory Studies Author(s): Edward J. Buskey, Jay O. Peterson, Julie W. Ambler Source: Limnology and Oceanography, Vol. 41, No. 3 (May, 1996), pp. 513-521 Published by: American Society of Limnology and Oceanography Stable URL: http://www.jstor.org/stable/2838586 Accessed: 25/10/2010 20:11 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=limnoc. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Society of Limnology and Oceanography is collaborating with JSTOR to digitize, preserve and extend access to Limnology and Oceanography. http://www.jstor.org Limnol. Oceanogr.,41(3), 1996, 513-521 ? 1996, by the American Society of Limnologyand Oceanography,Inc. The swarmingbehavior of the copepod Dioithona oculata: In situand laboratorystudies EdwardJ. Buskeyand Jay0. Peterson Marine Science Institute,750 Channelview Dr., Universityof Texas at Austin,Port Aransas 78373 JulieW. Ambler Department of Biology, MillersvilleUniversity, Millersville, Pennsylvania 17551 Abstract The behaviorof the swarm-formingcopepod Dioithonaoculata was studiedboth in situ and in the laboratoryusing a video-computersystem for motion analysis. In nature,swarms form in lightshafts between theprop roots of redmangroves. Swarms maintain their position within these light shafts despite currents ofup to 2 cm s- 1.In thelaboratory, swimming speeds and turningrates of swarming copepods in stillwater werelower than those observed in thefield. Copepods studied in a flowthroughchamber in thelaboratory had swimmingbehaviors similar to thoseobserved in nature;the stimulation from water movement caused increasesin bothswimming speed and rateof change of direction. Increased current speeds also causedthe swarmsto becomemore tightly packed within the center of a verticallight shaft. Nonswarming copepods wereunable to maintaintheir position in a currentin darkness.In laboratoryexperiments, the presence of activelyfeeding planktivorous fish caused swarmsto temporarilydisperse due to escape responsesof the copepods.However, planktivorous fish were rarely observed feeding on swarmsin nature,perhaps due to thepresence of predatory fish hiding among the prop roots. Copepods have been frequentlyreported to formdense swarmin theabsence of active swarming behavior (Oku- aggregations,called swarms,in a wide range of habitats, bo 1980). includinglakes (Byron et al. 1983), temperateand sub- Numerousdetailed studies have focusedon theswim- tropical marine bays (Ueda et al. 1983), coral-reefenvi- mingbehavior of captivemarine and freshwatercope- ronments(Emery 1968), and near mangrove cays (Am- pods underlaboratory conditions (e.g. Strickler1975; bler et al. 1991). The cyclopoid copepod Dioithona ocu- Wonget al. 1986; Tiselius 1992), but fewstudies have lata is a swarm-formingcopepod commonlyfound near beendone on theswimming behavior of copepods in their coral reefsand mangrovecays (Hamner and Carlton 1979; naturalenvironment (Schulze et al. 1992).The lackof in Ambleret al. 1991). Swarms ofD. oculata near mangrove situstudies of copepodbehavior is partlybecause cope- cays typicallyform between prop roots in shaftsof sun- podsusually are found at lowdensities in nature,so view- lightthat penetrate the mangrovecanopy. These swarms ingthem on a spatialscale appropriate for detailed studies form at dawn and disperse at dusk and are composed of theirswimming behavior (field of view of a fewmil- primarilyof adults and late-stagecopepodites (Ambleret limeters)is difficult.In addition,copepods usually exist al. 1991). The suggestedadaptive advantages of swarm in mixedspecies assemblages in nature,so preciseiden- formationin planktoniccopepods includeprotection from tificationof speciesfrom videotape records may be dif- predators, enhanced opportunitiesfor mating, and re- ficultwhen images are small enoughto allow extended strictionof dispersal by currents(Hamner and Carlton swimmingbouts to be observed.The copepod swarms 1979; Folt 1987). The formationand maintenanceof these thatform between the mangrove prop roots in Belizeare swarms must include a behavioral component because ideal forstudying copepod behavior in situ;high-density swarms are composed of a single species of copepod in copepodswarms of a knownsingle species form in pre- an environmentwhere numerous species are found and dictablelocations and remainin these locationslong because turbulentdiffusion would lead to dispersalof the enoughto allow filmingof theirbehavior. In addition, this swarmingbehavior makes it easy to capturelarge numbersof individualsof the same speciesfor detailed laboratorystudy, and theseswarms can be re-createdeas- Acknowledgments ilyin thelaboratory. This studywas fundedby a grantfrom the National Science Mostlaboratory observations of copepod behavior are Foundation(OCE 92-18516). madein stillwater, even though such conditions are rarely KeithSchmidt and ChrisCollumb provided assistance with observedin nature.Little is knownabout the behavioral data analysis.We thankKlaus Ruetzlerfor allowing us to use thefacilities at CarrieBow Cay and FrankFerrari for assisting responsesof zooplankton to watercurrents, although sev- witharrangements. eral recentstudies have investigatedthe effectsof tur- Universityof Texas Marine ScienceInstitute contribution bulence on copepod feedingand swimmingbehavior 943 andCaribbean Coral Reef Ecosystem Program contribution (Costelloet al. 1990; Saiz 1994; Ki0rboeand Saiz 1995). 463. In situobservations of copepod swarms during our study 513 514 Buskey et al. revealedthat zooplankton experience both directional Experimentsrun during the day werecarried out be- currentsand turbulenceeven when surfaceconditions neaththe laboratorybuilding in a coveredarea thatre- appearcompletely still, and these water movements clear- ceived no directsunlight. Nighttime experiments and lyaffect their behavior. Most previous studies of swarm- daytimelight-intensity studies were carried out in a dark- formingcopepods were field studies describing the dis- enedstorage room that excluded stray light. A blackfabric tribution,densities, and age structuresof swarms formed enclosurefor the experimental containers further exclud- in thefield (e.g. Emery 1968; Hamnerand Carlton1979; ed straylight. Swarming behavior was inducedin the Ueda et al. 1983). In thisstudy, we made detailedmea- laboratorywith a lightshaft produced by a fiber-optic surementsof swimmingbehavior of swarmingcopepods illuminator(Cole Parmer,150-W quartz-halogen lamp) in situ,and, in laboratoryexperiments, we examinedthe witha 5-mm-diameterlight pipe equipped with a focusing effectsof lightintensity, swarm density, water currents, lens.The fiber-opticlight pipe produceda verticalcone- and visual predatorson the swarmingbehavior of D. shaped shaftof light3 cm in diameterat the water's oculata. surfaceand 6 cm in diameteron thebottom to simulate shaftsof naturalsunlight that penetrate the mangrove canopyin thefield. Swimming behaviors of copepods in Materials and methods laboratoryexperiments were videotaped in the vertical planewith the same system used for in situstudies. Image Our studieswere carried out at theNational Museum contrastof the copepodsunder low lightwas enhanced of NaturalHistory's field station at CarrieBow Cay off withillumination produced by a ringof diodesemitting thecoast of Belize. Copepod swarms were collected near infraredlight (peak wavelength, 890 nm)arranged to pro- TwinCays, a mangrove-coveredisland 2 kmnorthwest duce dark-fieldillumination. These long wavelengths of CarrieBow Cay. Copepod swarmswere filmed in situ probablywere not perceivedby the copepods (Steams alongthe Lair Channelof Twin Cays in July1993 and and Forward1984; Buskeyet al. 1989). inTwin Bays in January 1994 and May 1994.The swarms The effectof light intensity on swarmingbehavior was werein shallowwater (< 1 m) betweenprop roots in shafts examinedbetween 1400 and 1600hours in a 20 x 20 x 20- of sunlightthat penetrated the canopyof the red man- cm clear acrylicaquarium filled with seawater filtered grove(Rhizophora mangle). througha 20-Am-meshsieve. Light intensities of between The swarmswere video recordedin situwith a Cohu 10 and 1,280 Amolphotons m-2 s-' wereproduced by 3315 monochromeCCD video cameraequipped with a adjustingthe rheostat on thefiber-optic illuminator and macrolens (Micro-Nikkor55 mm f2.8)and placedin a by.using neutral density filters. Light intensity was mea- waterproofhousing (Video Vault). The cameraand hous- suredwith a LiCormodel 158A photometer with
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