Vertical migration and diel activity patterns of hydromedusae: studies in a large tank

Vertical distributions and swimming behaviors of 7 species ofhydromedusae (Aequorea victoria, Bougainvillia principis, Gonionemusvertens, Mitrocoma cellularia, Phialidium gregarium, Polyorchispenicillatus, and Stomotoca atra) were observed over 24-h periodsin a 2 m high, 1500 1 transparent tank. In this tank, most speciesperformed well-marked diel vertical migrations that were mediatedby swimming. Manipulation of the light regime showed the didswimming behaviors to be light-dependent rather than intrinsic,even in species that do not possess recognized photoreceptors.Correlations between vertical migration and spawning times forseveral species of medusae suggest that in cases when verticalmigration reduces the distances between individuals (e.g., inmass movements to the surface) just prior to spawning, enhancedfertilization success may result from such movements. ¹Present address: Friday Harbor Laboratories, Friday Harbor,WA 98250, USA.     

Synchronized release of medusae from three species of hydrozoan fire corals

 The release of medusae from three hydrozoan fire corals, Millepora dichotoma, M. murrayi and M. platyphylla, was investigated at three sites in southern Taiwan from February 1994 to July 1995. All three species were gonochoristic, and developed and released several batches of medusae between April and May. The duration of open ampulla appearing on the surface of coralla was short, about three months, and could be used to infer the reproductive season of the fire corals between April and May. No obvious lunar cycles of medusa release were found in these species. Medusa release started before dark at approximately 17:00 h and continued for several hours. Males began releasing medusae earlier than females. Synchronization of medusa release between colonies, i.e., the probability of occurring on the same nights, was significantly higher within a species than between different species. Hybridization in nature among the three species is, therefore, unlikely due to segregation in the spawning dates. Moreover, the synchronization within each species was often significantly higher within versus between sites. The free-swimming medusae released gametes within approximately one hour, and the spent medusae lived for a few more hours. Medusae may facilitate fertilization rates as a result of an apparently negatively geotactic swimming response that results in medusa aggregation at the surface. No differences in the sizes of medusae, eggs and sperm were detected among the three species; however, some characteristic differences of medusa nematocysts were found. Accepted: 25 September 1997     

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.     

Carbon content of the neritic scyphomedusa Chrysaora fuscescens

An analysis of the role of scyphomedusae in a planktonic ecosystemrequires that biomass or numerical abundance estimates be convertibleinto standard units for comparison with other components ofthe planktonic community. One species under investigation isthe brown sea nettle, Chrysaora fuscescens, which is very abundantin coastal waters of the west coast of North America (Shenker,1984). For this species, the carbon content of whole immatureanimals and the body components of sexually mature medusae weredetermined. Immature medusae contained a mean carbon content0.202% of the wet weight. The bell, oral arms and gonadal tissueof mature medusae had mean carbon levels 0.156, 0.554 and 0.576%of the wet weight, respectively. When the relative proportionsof these body tissues were calculated, the mean carbon contentof whole mature medusae was determined to be 0.280% of the wetweight.     

The Scyphomedusae of the Rockall Trough, northeastern Atlantic Ocean

An extensive series of samples between the surface and 2700m depth have been taken in the Rockall Trough between 1973 and1978. The coronate medusae, in decreasing order of abundanceoccurring in the Trough are: Periphylla periphylla, Atolla wyvillei,A. parva, A. vanhoeffeni and Nausitho? globifera. The last specieswas rare and no information on its biology was obtained. Theother species breed within the Trough, the three Atolla speciesthroughout the year. Few large mature Periphylla periphyllawere caught and its breeding season remains undefined. Theirmorphology and vertical and geographical distributions are brieflydiscussed. The semaeostomid medusae, Pelagia noctiluca, Aureliaaurita and Cyanea lamarckii also occurred, the latter two speciesconfined to the Rockall Bank.     

Bathymetric distribution of medusae in the open waters of the middle and south Adriatic Sea during spring 2002

Twenty-eight species of hydromedusae and one scyphomedusa werecollected from the middle and south Adriatic in Spring 2002.Haliscera bigelowi was recorded for the first time in the AdriaticSea and Dicodonium adriaticum for the southern Adriatic. Thecosmopolitan species Rhopalonema velatum, Aglaura hemistomaand Persa incolorata averaged 88% of total medusan abundance.The Mediterranean species Rhabdoon singulare and Solmissus albescenswere numerous at the deeper stations. These species were foundeither in the southern or in middle Adriatic, and their abundanceis greater than in other regions of the Mediterranean Sea. Speciesdiversity and abundance generally were highest in the upper200 m and decreased with depth. Diel vertical migration patternsincluded some species that nocturnally sought shallower or deeperlayers, and some species migrated in both directions. A fewspecies did not show any vertical migration. Although lightlikely triggers vertical movements, data suggest that foragingstrategies also influence the vertical migration of medusae.     

Controlled activation of species typical behaviour to low salinity,seawater movement and seawater depth in <Emphasis Type="Italic">Aurelia labiata</Emphasis> (Scyphozoa) Jellyfish in Roscoe Bay,Canada

Controlled observations in Roscoe Bay examined whether Aurelia labiata medusae would respond to three environmentally significant stimuli: low salinity, seawater movement and seawater depth. Significantly more upward-swimming medusae turned and swam down or sideways when they encountered an experimentally created low-salinity plume than did upward-swimming medusae that encountered a seawater plume or a seawater plume containing milk powder. Downward-swimming medusae that encountered a plume containing freshwater continued swimming down. Significantly more upward-swimming medusae exposed to an experimentally created stream of seawater that gently pushed them horizontally turned and swam down than did upward-swimming medusae not exposed to the seawater stream. Downward-swimming medusae continued to swim down when pushed horizontally. A stronger seawater stream that tumbled as well as pushed medusae horizontally was less effective. In a shallow area near the time of a low tide, most medusae were in the top metre of a 2 m water column whilst in an adjacent area where the water was 4–6 m deep, almost all medusae were below 1.5 m. These observations add to a growing body of evidence that the ecological distribution of A. labiata is influenced by their ability to respond adaptively to significant environmental stimuli.     

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.     

Chemical composition and biomass parameters of a population of Craspedacusta sowerbii Lank1880 (Cnidaria: Limnomedusa)

Until now, very little information about the ecology of thefreshwater jellyfish, Craspedacusta sowerbii, has been available.Although many publications deal with Craspedacusta, most ofthem contain only observations. Detailed analyses are rare.In this study, investigations on size-dependent fresh and dryweights, and carbon, nitrogen and phosphorus, are presented.The water content of the medusae ranged between 96.74 and 99.87%.Fresh and dry weights ranged from 0.06 to 331.86mg FW ind.^0–1and 0.01 to 2.50 mg DW ind.^–1, respectively. The molarC:N:P relationship was calculated as 39:9:1. The C:P ratio waslow in contrast to other freshwater zooplankton species, andimplied a high demand for P. A comparison with the C:P stoichiometryof their prey indicates the possibility of P limitation of Craspedacustaat a gross growth efficiency above 25%. The results are discussedin relation to medusae abundance, bioaccumulation and theirimpact on food web structure.     

Field observations of four Aurelia labiata jellyfish behaviours: swimming down in response to low salinity pre-empted swimming up in response to touch,but animal and plant materials were captured equally

Jellyfish live in complex environments and must continually make behavioural choices. In field observations, adult Aurelia labiata were confronted with a conflict between swimming up elicited by touch of the manubrium and swimming down elicited by low salinity. Following a touch, downward-swimming medusae (1.5–2.0 m deep) turned and swam to within 0.5 m of the surface when the salinity in the top 1.5 m of the water column was greater than 20 ppt but medusae uniformly refused to swim up into the top 1.25 m when the salinity was less than 20 ppt even after being touched three times. The central nervous system of A. labiata appears to have neural circuitry that specifies their response when medusae encounter stimuli that elicit incompatible behaviours. Upward-swimming adult medusae had animal, vegetable or cellulose (paper) material dispersed ahead of them. Medusae captured each material on the bell margin and transported it to a gastric pouch. Medusae displayed only minor behavioural differences in the process. Having sensory, neural and muscular systems organized to capture and pass to the stomach, a huge variety of materials allows medusae to survive in different seasons and environments.     

Propulsion in Cubomedusae: Mechanisms and Utility

Evolutionary constraints which limit the forces produced during bell contractions of medusae affect the overall medusan morphospace such that jet propulsion is limited to only small medusae. Cubomedusae, which often possess large prolate bells and are thought to swim via jet propulsion, appear to violate the theoretical constraints which determine the medusan morphospace. To examine propulsion by cubomedusae, we quantified size related changes in wake dynamics, bell shape, swimming and turning kinematics of two species of cubomedusae, Chironex fleckeri and Chiropsella bronzie. During growth, these cubomedusae transitioned from using jet propulsion at smaller sizes to a rowing-jetting hybrid mode of propulsion at larger sizes. Simple modifications in the flexibility and kinematics of their velarium appeared to be sufficient to alter their propulsive mode. Turning occurs during both bell contraction and expansion and is achieved by generating asymmetric vortex structures during both stages of the swimming cycle. Swimming characteristics were considered in conjunction with the unique foraging strategy used by cubomedusae.     

Swimming behavior of Daphnia: its role in determining predation risk

Individual swimming behavior of zooplankton can play an importantrole in determining how planktivorous fishselect their prey.Although several studies have documented the effect of preysize, contrast or degree of pigmentation, escape ability, encounterrate and abundance in determining predation risk, the importanceof individual behavior has received relatively little attentionby aquatic ecologists. Recent advances in the technology ofvideo recording and computer analysis of motion have allowedus to collect digitized three-dimensional videorecords of free-swimmingzooplankton such as Daphnia. We found that Daphnia clones, includingthose within a single species, exhibit a wide range of swimmingbehaviors as measured by swimming speed. The individual behaviorof a species cannot be adequately described by looking at oneclone. We alsoshow that different behavior observed in liveDaphnia can play an important role in determining attractivenessto visual predators. Given a choice between two clones of equalsize and visibility contrast, fish selected indi viduals fromthe faster swimming clone. Our results suggest that currentmodels of prey selection would be improved by the incorporationof individual swimming behavior because it is an important factordetermining overall prey visibility.     

The ontogeny of swimming behavior in the scyphozoan, Aurelia aurita. II. The effects of ions and drugs.

1. The responses of Aurelia medusae to pharmacological agents and ionic variation were classified into four response types: Type I, no response; Type II, inhibition of pacemaker activity; Type III, inhibition of both pacemakers and swimming muscles; and Type IV, increase in pacemaker output. 2. The swimming pacemakers of Aurelia medusae become hyperactive in Mg+2-free solutions (Type IV). This response appears to be general in swimming scyphozoa. 3. The response pattern to pharmacologically-active compounds indicates that the coelenterate neuromuscular system is quite different than those in other phyla. In fact, the response spectrum is not consistent within the Cnidaria. 4. Similarly, the responses of adult medusae to ionic variation show no consistent pattern within various scyphomedusae. 5. Test solutions from each response type established with medusae were selected and tested on the scyphistoma and strobila stages. The comparison of the responses to the test solutions between the medusa, scyphistoma, and strobila showed that the neuromuscular systems are physiologically different. The strobila, specificially the ephyra, is a mixture of both polypoid and medusoid response types. The strobila, therefore, is physiologically an intermediate stage in the development of the adult medusa.     

Comparative behaviour of two species of Lepadogaster (Pisces: Gobiesocidae) living at different depths

The two clingfish species studied occupied similar habitats but occurred at different depths. When compared with the subtidal species Lepadogaster candollei , the intertidal species Lepadogaster l. purpurea was less active, spent more time in shelters, visited fewer shelters, showed more site fidelity, and spent less time swimming. Feeding, swimming, and agonistic behaviours were performed mainly in close contact with the substrate in this species. It is hypothesized that these contrasts in behaviour may have evolved under different levels of turbulence.     

Host activity and the risk of nest parasitism by brown-headed cowbirds

Proportions of nests parasitized by brown-headed cowbirds (Molothrus ater) vary greatly among host species, but factors underlyingthis variation remain poorly understood. Cowbirds are believedto find nests by watching host behavior. We tested the hypothesisthat the activity of hosts during nest building correlateswith the probability of parasitism among and within four sympatrichosts: dusky flycatchers (Empidonax oberholseri), warblingvireos (Vireo gilvus), yellow warblers (Dendroica petechia),and American redstarts (Setophaga ruticilla). Daily probabilityof parasitism varied substantially among these species, from3% for dusky flycatchers to more than four times that for warblingvireos. The four species did not differ in the proportion ofcowbirds fledged from their nests. Differences in nest placementdid not explain differences in probability of parasitism amongor within species. Parasitism frequencies increased among speciesthat made longer nest-building visits, had a greater propensityto perch during nest approach, spent more time near their nests,and had males that vocalized more often near nests. Within species, females that visited their nests less often, spent more timeon the nest per visit, and males that sang more and were activein a smaller area around their nests were more likely to beparasitized by cowbirds. These data support the hypothesisthat cowbirds use the activity of hosts during nest buildingto locate nests.     

Trophic interactions of the freshwater jellyfish Craspedacusta sowerbii

The predatory impact and the trophic role of the freshwaterjellyfish, Craspedacusta sowerbii, was studied using microcosmand enclosure experiments as well as a 3-year pond survey. Theresults showed a significant decrease of small herbivorous crustaceans,i.e. Bosmina longirostris and juvenile cyclopoid copepods, inthe medusa treatments of the microcosms and the enclosure experiments.Chlorophyll concentrations in the enclosure experiment weresignificantly increased in the medusa treatment, suggestingthat C. sowerbii may cause cascading effects in the food chain.A comparison of daily zooplankton losses during the pond surveycaused by medusae and fish (roach, Rutilus rutilus), and theirfood selectivities suggest food separation of these two predatorsand reveal a strong negative impact of medusae on the copepodpond community. In the case of a jellyfish bloom, our resultsshow that both food chains can co-occur in lakes because ofa weak interaction between these top predators, fish and jellyfish,with simultaneous impacts on the zooplankton structure.     

Harvest rates and foraging strategies in Negev Desert gerbils

We examined the foraging strategy and quantified the foragingtraits of two nocturnal rodent species, Allenby's gerbil (Gerbillusallenbyi) and the greater Egyptian sand gerbil (Gerbillus pyramidum).In the laboratory, both species used two distinct foragingstrategies: either they immediately consumed seeds found ina patch (seed tray); or they collected and delivered the seedsto their nest box for later consumption. Moreover, we founda transition in foraging strategy among individual G. allenbyi under laboratory conditions; they all began by consuming theseeds on the tray and, after 7 days on average, switched tothe collecting strategy. By contrast, in the field both speciesused only one foraging strategy; they collected and deliveredthe seeds to their burrow or to surface caches for later consumption.Furthermore, G. allenbyi and G. pyramidum collected seeds atsignificantly higher rates in the field than in the laboratorybecause the seed encounter rates for both species were higherin the field. This suggests that in natural conditions, probablyinvolving predation risk and competitive pressure, gerbilsmust respond in two ways: (1) they must choose a foraging strategythat reduces predation risk by minimizing time spent feedingoutside their burrows; and (2) they must forage more efficiently.In the field, seed handling time of the larger species, G. pyramidum, was shorter than that of the smaller one, G. allenbyi.This difference may give G. pyramidum an advantage when resourcelevels are high and when most of a forager's time is spent handling seeds rather than searching for more seeds. Additionally,our field study showed that the seed encounter rate of G. allenbyiwas higher than that of G. pyramidum. This difference may giveG. allenbyi an advantage when resource levels are low and whensearching occupies most of the forager's time. The differentadvantages that each species has over the other, under differentconditions, may well be factors promoting their coexistenceover a wide range of resource densities.     

Food regulation of growth and maturation in a natural population of Aurelia aurita (L.)

Growth and maturity development of the moon jellyfish. Aureliaaurita, were recorded in Vgsbpollen, a small and semi-enclosedbay on the Norwegian west coast, and compared to those of medusaetransferred to excess food and starving conditions, respectively.Mesozooplankton were extremely scarce in Vgsbpollen. The abundanceand biomass of the medusae in the poll were higher than thosetypicallyfound in open waters, reaching a maximum of 22 ind.m^–3 and 710 mg C m^–3 in June. The average diameterof medusae in the p increased to 8 cm until the last part ofJune, with an instantaneous growth rate between 1.5 and 20%day^–1, thereafter retarding somewhat, giving a negativegrowth rate of up to 2.6% day^–1. Starving medusae showeda negative growth rate ofup to 13.4% day^–1, and all thernedusae were dead after 49 days. Well-fed medusae showed avery stable growth over a 56 day period, diverging from thepollpopulation from early June, and with a growth rate between3.8 and 9.8% day^–1. Medusae from the pollpopulation begancarrying planulae on their oral arms when at least 5 cm in diameter,whereas not even the largest medusa of 15.6 cm diameter amongthose in the well-fed group produced any planulae. For the firsttime, it is thus explicitly shown that thesize and maturityof A.aurita are externally controlled through food availability.Scarcity of food reduces the growth rate, but also changes theenergy allocation towards reproduction, which thus occurs ata smaller size than for well-fed rnedusae. Its plasticity makesit possible for this species to exploit environments with lowadvection of food and develop high abundance in such environments,without losing fecundity.     

Locomotion and neuromuscular system of Aglantha digitale

Aglantha digitale swims in two ways: a slow rhythmical swim typical of hydromedusae in general and a sudden rapid movement that appears to be an escape response. The swimming musculature is an extremely well developed striated circular muscle layer that possesses a sarcoplasmic reticulum. The nervous system of this species can be divided into three units: an inner nerve ring and an outer nerve ring, which are joined by unusually large transmesogleal pathways, a group of giant axons that extends over the surface of the swimming muscle, and the radial canal. Well developed ciliated sensory cells are located on the exumbrellar surface of the margin. Consideration of these properties of the organisation of this species suggests that normal slow swimming is controlled by a mechanism similar to that found in other medusae, while the escape response is the result of the action of the giant axons.     

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.