Variability in Daphnia behavior following fish community manipulations

Vertical migration and feeding periodicity of daphnids weremonitored for three summers in three lakes, two of which hadundergone reciprocal fish transfers during May of the secondyear. In Peter and Paul Lakes, the diel migratory behavior ofthe Daphnia assemblages was fairly constant. In Tuesday Lake,where fish manipulations greatly reduced planktivory. Daphniapulex became abundant and exhibited high variability in thedirection of diel migration. Daphnids of Tuesday Lake were smallerand more recently established than those of the other lakes.Weather and lunar phase did not correlate well with migratoryor feeding behavior, though there was a negative correlationwith the chlorophyll a content of the mixed layer. In this studyhigh planktivory by fish was associated with pronounced andconsistent nocturnal migration. Low planktivory by fish wasassociated with variable, inconsistent migration. Evidently,planktivorous fish are essential for maintenance of verticalmigration in these daphnid populations.     

Vertical migration and feeding of Euphausia lucens (Euphausiacea) in the Southern Benguela

Diel migration and feeding were examined for populations ofEuphausia lucens living in the near- and offshore waters ofthe southern Benguela. Euphausiids at both stations displayednocturnal diel vertical migration patterns. Animals inshoreascended in a slow-fast-slow manner which seems to be relatedto differential food concentrations throughout the water columnPronounced quantitative and qualitative changes in the dietof E.lucens were observed offshore but not inshore. These differencesare discussed in relation to different ambient food environments     

In situ diel variation in gut pigment contents of Ceriodaphnia sp. in stratification and destratification periods

The short-term, in situ diel grazing of Ceriodaphnia sp. duringperiods of stratification and mixing was investigated usingthe technique of fluorimetric analysis of the gut pigments.There were considerable seasonal differences in feeding behaviourIn mixing, when the concentration of chlorophyll a in the watercolumn was high and Ceriodaphnia abundance was low, gut pigmentcontents showed clear diel variation patterns, probably dueto diel variations of the high values of feeding activity observedin the 24 hour cycle The maximum values were found at dawn.On the other hand, no diel variations in gut pigment were observedduring periods of stratification and while the amounts of pigmentsin the water and in the gut were very low, species abundancewas high. Taking into account the ambient conditions, the authorsdiscuss the possibility that the change of feeding of the Ceriodaphniasp. observed when the environment changed from a mixing periodto one of stratification represents a change from herbivorousto detritivorous behavior.     

A change in the pattern of vertical migration of Chaoborus flavicans after the introduction of trout

The pattern of diel vertical migration exhibited by Chaoborusflavicans in Lake Lenore, Washington has changed over the pastdecade. Formerly Chaoborus larvae and pupae were present inthe water column during the day. Since 1982, a strong patternof diel vertical migration has been evident in this population.Third and fourth instar larvae and pupae reside in the sedimentsduring the day and then ascend to the surface waters at night.The change in migratory activity of C. flavicans was coincidentwith the introduction of cutthroat trout (Salmo clarki henshawi)into Lake Lenore in 1979. Predation by trout on fourth instarlarvae and pupae of C. flavicans was probably responsible forthe observed changes in migratory behavior.     

Diel vertical migration of Cryptomonas and Chromatium in the deep chlorophyll maximum of a eutrophic lake

Large populations of the phytoflagellates Cryptomonas rostratiformis,Cryptomonas phaseolus, Cryplomonas undulata and the phototrophicbacterium Chromatium cf. okenii have been observed in the oxic/anoxicboundary layer of the slightly eutrophic, dimictic lake Schlachtenseeduring the summer stratification. Vertical distribution of thesepopulations was studied with the help of a new close-intervalsampler and by in Situ fluorescence measurements with fine spatial(cm to dm) and temporal (h) resolution on 4 days in 1991, 1994and 1995. All populations lived close to the chemo dine butshowed a regular diel vertical migration with daytime ascentand night-time descent and a low migration amplitude. At leastfor one species— C. rostratiformis—the pattern ofmigration suggests that this behaviour has an endogenous origin.As a result of diel vertical migration, the population of C.rostratiformiswas spatially separated from the other dominating populationsduring daytime. Ultimate cause of diel vertical migration wasa better light supply of all populations. Another probable advantagewas the reduction in grazing pressure, as large percentagesof all populations moved into the anoxic, hydrogen sulphidecontaining water layers during night-time. ¹Present address Department of Lowland Rivers and Shallow Lakes,Institute of Freshwater Ecology and Inland Fisheries, Müggelseedamm260, D-12587 Berlin, FRG     

Diel vertical migration in zooplankton: rapid individual response to predators

While diel vertical migration in zooplankton has been shownrecently to be a predator avoidance behavior, the mechanismby which predators induce and maintain such behavior has beendebated. We report results of an in situ predator manipulationexperiment during which enclosed populations of the marine planktomccopepod Acaraa hudsonica rapidly changed their vertical distributionand diel migration behavior depending on presence or absenceof the planktivorous fish Casterosteus aculeatus These resultspoint unambiguously to phenotypic behavioral plasticity of individualplanktonic prey, not, as previously hypothesized, population-geneticlevel behavioral changes caused by selective fish predation,as the mechanism underlying changes in diel vertical migrationin this copepod.     

The diel migrations and vertical distributions of zooplankton and micronekton in the Northwestern Mediterranean Sea. 2. Siphonophores, hydromedusae and pyrosomids

Vertical distributions and diel migrations in the 0–1000m water column at a position in the Northwestern MediterraneanSea are given in detail for the main species of gelatinous macrozoo-plankton:three siphonophores (Abylopsis tetragona, Chelophyes appendiculataand Lensia conoidea), one hydromedusa (Solmissus albescens)and one pyrosomid (Pyrosoma atlanticum). Extensive diel verticalmigration occurred in these five species, particularly in A.tetragona,S.albescens and P.atlanticum for which the bulk of the populationmigrated as a compact unit. Pyrosoma atlanticum underwent thelargest migration, with a mean amplitude of 515 m. The migrationpattern of L.conoidea was more complex. Its day-distributionwas bimodal and at night part of the population ascended towardsthe surface, while the other part appeared to undergo a weakreverse migration of     

Diel periodicity and selectivity in the feeding rate of the predatory copepod Mesocyclops edax

The diel periodicity and selectivity in the feeding behaviorof the predatory cyclopoid copepod Mesocyclops edax was investigatedat 3 h intervals over two 24 h sampling periods in nature. Gutcontent analyses revealed an increase in gut fullness at 11.30and 20.30 in July, and at 20.30 in August. The increase in gutfullness at these times could not be explained by an increasein prey density, changes in predator-prey overlap or differencesin prey vulnerability. We suggest that M.edax exhibits a truediel periodicity in the intensity of its predatory activities,although the alternative hypothesis of a diel periodicity ingut passage time cannot be ruled out. Vanderploeg and Scavia'sE^* selectivity index indicated a preference for rotifer andcladoceran prey over copepods.     

Diel reproduction and vertical distributions of the marine cladocerans, Evadne tergestina and Penilia avirostris, in contrasting coastal environments

Evadne tergestina and Penilia avirostris were sampled duringseveral diel cycles in the well-mixed, turbid Inland Sea ofJapan and the more stratified, less turbid Gulf of Mexico shelf.Parthenogenetic female Evadne contained embryos with pigmentedeyes only at night, and apparently released neonates near dawnor in the morning. There was no significant diel vertical migrationin the Gulf of Mexico, and a ‘reverse’ migrationin the Inland Sea. Female Penilia could contain mature embryosat any time, but were most likely to do so at night. Peniliadid not migrate dielly, and in the Gulf of Mexico was concentratedin a nepheloid layer near the bottom.     

Short-term feeding rates of A carrier grani in natural conditions: diurnal variation

Short-term in situ diel grazing activity of adult male and femalesofAcania graniSars. were investigated at a coastal station inthe Alboran sea (SE of Spain). Gut pigment contents were measuredfluorometrically, and the ingestion rates, filtration ratesand daily ratios were calculated from gut fullness, evacuationrates and pigment concentrations in sea water. The results indicatethat there were size-related differences in the amount of foodingested by males and females. Gut contents of females wereon average >1.5 times that for males. Body-volume-specificfeedingestimates were, however, the same for both sexes. Gutpassagetimes, estimated from animals collected at the time of day whengut pigments were maximal, were shorter in females. Both sexesshowed diel changes in feeding activity and faecal pellet productionof about one order of magnitude;the maximum rates were observedat dusk, the minimum rates at midday. There was a directrelationshipbetween copepod grazing and pigment concentration in sea waterwhen a time lag was considered. Daytime and night-time in situincubations using natural sea water enriched withlaboratorycultured algae provided evidence of non-continuous feeding andof light intensity control of the diel patterns. The possibleadaptive advantages of differences in sexual size on feedingbehaviour and diurnal variability are discussed.     

Feeding, excretion and egg production by individuals and populations of the marine, planktonic copepods, Acartia spp. and Centropages furcatus

Diel variations in vertical distribution, gut pigment content,ammonium excretion and egg production were investigated foradult females of Acartia erythraea and A.pacifica in the verticallymixed Inland Sea of Japan and Centropages furcatus in the stratified,neritic Gulf of Mexico. Gut pigment content and egg productionrate were maximal at night and ammonium excretion was maximalduring the daytime. Neither A.erythraea nor A.pacifica adultfemales showed an apparent diel migration, but the former werehighly concentrated in the surface layer during the afternoon.In contrast, C.furcatus adult females showed a clear diel migration,residing immediately above the bottom during the daytime andbeing concentrated between 10 and 25 m depth during the nighttime.Individual-based data on gut content and excretion and egg productionrates were combined with vertical-distribution data to calculatepopulation values. In the Inland Sea of Japan, the resultantpattern for Acartia spp. reflected the diel variation in physiologicalrates and even distribution of adult females, except for theafternoon, surface aggregation of A.erythraea. In the Gulf ofMexico, the pattern for C.furcatus reflected largely the dielvariation in each rate process and the heterogeneous distributionof adult females in the water column. Elevated nocturnal feedingactivity of these copepods may be due to an endogenous rhythm.The daytime maximum in ammonium excretion and night-time maximumin egg production rate indicated approximate half-day and daytime lags, respectively, after the intake of food until itsconversion into dissolved excreta and released eggs.     

Relationships between geotaxis/phototaxis and diel vertical migration in autotrophic dinoflagellates

Marine dinoflagellate diel vertical migrations are often conceptuallyexplained by a species' geotactic and phototactic preferences,but actual simultaneous measurements are rare. Newly collectedsimultaneous measurements on Heterocapsa (Cachonina) illdefina(Herman and Sweeney) and Gymnodinium breve (Davis) are combinedwith similar literature information on Amphidinium carterae(Hulbert), Peridinium faeroense (Paulsen) and Prorocentrum micans(Ehrenberg) to explore several examples of the actual relationshipsbetween diel vertical migration and geotaxis/phototaxis. Amphidiniumcarterae does not migrate, but it exhibits a negative geotaxisthat may counter a small sinking velocity. The four other speciesall exhibit diel vertical migrations that yield surface aggregationsduring daylight, but the associated combinations of geotaxisand phototaxis precision (which is strongest when every cellin a population exhibits the same response to a stimulus andweakest when the response is random) and sign [which is positive(negative) when motion is toward (away from) the stimulus] aredifferent in each case. These different taxis combinations maybe related to species-specific sensor structure and/or placement.Furthermore, variations in the different biochemical pools overa species' cell cycle may contribute to structural/mechanicalchanges that influence how a given sensory array functions ata given time. If so, this coupling may be an important linkin the growth optimization mechanisms and occasional bloom successesof different autotrophic dinoflagellate species under varyingenvironmental conditions.     

Diel patterns of zooplankton grazing in a shallow lake

A diel survey of in situ species-specific zooplankton clearancerates (with radioactively labelled Chlorella cells) was conductedin the shallow lake of Cr?teil characterized by small-sizedplanktonic forms (algae<10µm and zooplankton <1.3mm). Experiments were performed every 4h at two depths (1 and4m). Power functions relating individual filtering rates tobody length were established for the three most abundant cladoceransand for calanoids (nauplii being included in this feeding group),for each depth and time. No filtering penodicity was observedin Ceriodaphnia spp., adults and copepodites of Eudiaptomusgracilis and Eurytemora velox, and nauplii. On the contrary,clear nocturnal filtering peaks were obtained for Daphnia spp.and for Diaphanosoma brachyurum, these being more pronouncedfor the larger individuals at 1 m deep. The observed diel periodicitycannot be explained by variations in physico-chemical parametersor food concentration alone. The ecological significance ofthis phenomenon in the polymictic lake of Cr?teil is discussedin the light of previously published data and the hypothesesaccounting for it.     

Diet changes in the carbon and nitrogen content of the copepod Metridia lucens

Diel changes in the near-surface (0–50 m) abundance, prosomelength, and carbon and nitrogen content of the copepod Metridialucens were measured in Deep Cove, Doubtful Sound, New Zealand(45°27'S, 167°9'E) between 3 and 6 September 1996. Metridialucens showed maximal abundance a night, suggesting a patternof normal diel vertical migration (DVM). The change in abundancesuggested that the descent of the population occurred –1h prior to dawn and the ascent –1 h after dusk. However,a proportion of the population remained near the surface duringthe day. Although there was no diel pattern in the prosome lengthof M.lucens collected near the surface, there was a marked dielcycle in the measured carbon and nitrogen contents, with maximalvalues being measured towards the end of the night prior tothe downward migration. We suggest that this diet cycle in themeasured elemental content was caused by DVM occurring morestrongly in those animals which had a better body condition,i.e. a higher elemental content per unit length.     

Marine cladocerans in Toyama Bay, southern Japan Sea: seasonal occurrence and day-night vertical distributions

Marine cladocerans in the zooplankton samples collected vertically(0–500 m) with Norpac nets at 2–4 week intervalsfrom February 1990 through January 1991 at a station in ToyamaBay, southern Japan Sea, were examined. From mid-March throughearly December, cladoceran populations exhibited five distinctpeaks. in which seven species occurred and showed a species-specific pattern of seasonal distribution. In mid-July. Peniliaavirostris exhibited a prominent peak in population density.Three species of Evadne, E.nordmanni, E.spinifera and E.tergestina,showed a clear ecological separation in time. The first twospecies had a peak density in April and July. respectively,but the last species demonstrated three distinct peaks betweenJune and September. Two congeneric species of the genus Podon,P.leuckarti and P.schmackeri, also did not co-exist in time.Simultaneous horizontal hauls with MTD nets made in June andSeptember 1986 revealed a strikingly marked surface distributionof all species. Evadne nordmanni occurred in June and demonstrateda clear ‘reverse’ did vertical migration, residingin the 30 m depth during the night time and ascending up tothe uppermost water column of 0–10 m during the daytime.Among the four species which occurred in September, E.spinifera,E.tergestina and P.schmackeri also showed a reverse migrationbetween the 30 m depth during the night and the very surfaceduring the day. A different pattern was observed in P.avirostrzs,which was distributed at a depth of 30 m during both the dayand the night with no sign of diel vertical migration. Amongthe more prevalent parthenogenetic individuals, some gamogeneticforms were found to exist in most species occurring in ToyamaBay.     

Predator-dependent diel migration by <Emphasis Type="Italic">Halocaridina rubra</Emphasis> shrimp (Malacostraca: Atyidae) in Hawaiian anchialine pools

Diel migration is a common predator avoidance mechanism commonly found in temperate water bodies and increasingly in tropical systems. Previous research with only single day and night samples suggested that the endemic shrimp, Halocaridina rubra, may exhibit diel migration in Hawaiian anchialine pools to avoid predation by introduced mosquito fish, Gambusia affinis, and perhaps reverse migration to avoid the predatory invasive Tahitian prawn, Macrobrachium lar. To examine this phenomenon in greater detail, we conducted a diel study of H. rubra relative abundance and size at 2-h intervals in three anchialine pools that varied in predation regime on the Kona-Kohala Coast of Hawai‘i Island. We found two distinct patterns of diel migration. In two pools dominated by visually feeding G. affinis, the abundance of H. rubra present on the pool bottom or swimming in the water column was very low during the day, increased markedly at sunset and remained high until dawn. In contrast, in a pool dominated by the nocturnal predator M. lar, H. rubra density was significantly lower during the night than during the day (i.e., a pattern opposite to that of shrimp in pools containing fish). In addition, we observed that the mean body size of the shrimp populations varied among pools depending upon predator type and abundance, but did not vary between day and night in any pools. Our results are consistent with the hypothesis that H. rubra diel migratory behavior and size distributions are influenced by predation regime and suggest that diel migration may be a flexible strategy for predator avoidance in tropical pools where it may be a significant adaptive response of endemic species to introduced predators.     

Diel vertical migration and gut pigment rhythm of Paracalanus parvus,P. crassirostris,Acartia erythraea and Eucalanus subcrassus (Copepoda,Calanoida) in Tolo Harbour,Hong Kong

Diel vertical migration and gut pigment rhythm of calanoid copepods were studied in the highly eutrophic waters of Tolo Harbour, Hong Kong in summer between July 30 and August l, 1991 and in winter between January 24 and 25, 1992. Paracalanus parvus and P. crassirostris showed neither diel vertical migration nor diel gut pigment rhythm during the summer study. Acartia erythraea and Eucalanus subcrassus exhibited prominent diel vertical migration and diel gut pigment rhythm. Diel changes in gut pigment content of surface dwelling individuals and increase of gut pigment content before the onset of nocturnal upward migration suggest that gut pigment rhythm in E. subcrassus was not the result of animals moving in and out of a food-rich surface layer. During the winter study, the harbour was affected by a bloom of Noctiluca scintillans. P. parvus and P. crassirostris resided in deeper waters presumably to avoid the dense populations of N. scintillans at the surface. P. crassirostris remained non-migratory during the winter study. There was a nocturnal increase in the number of P. parvus in the surface waters, although no diel change in the mean depth of the population was observed. Both Paracalanus species showed diel gut pigment rhythm. The presence of diel gut pigment rhythm in the non-migratory P. crassirostris during the winter study suggested the presence of an independent feeding rhythm.     

Zooplankton-phytoplankton interactions in a eutrophic lake

Enclosure experiments were made in a cyanobacteria dominatedlake (Lake Rotongaio) to assess the impact of zooplankton (>150µm) grazing on algal growth rates and determine the effectof diel and vertical changes in zooplankton grazing intensityand nutrient (NH₄-N) regeneration upon abundance of phytoplankton.The filamentous cyanobacterium Anabaena minutissima var. attenuataand diatom Cyclotella meneghiniana showed a negative linearchange in abundance with a gradient in zooplankton grazing intensity.Phytoflagellates were not grazed and showed a positive linearchange in abundance with increasing zooplankton biomass. Theseeffects, as well as shortening of filament length of Anabaena,were caused by raptorial feeding by the alanoid copepod Boeckellapropinqua which dominated the zooplankton. Phytoplankton growthwas not stimulated by addition of nutrients, suggesting nutrientregeneration was not important. Diel and vertical changes infeeding and NH₄-N regeneration rates were measured in Marchand June 1988. Diel differences were more pronounced in Marchwhen the water column was stratified. Specific feeding rateswere more important than vertical changes in zooplankton biomassin determining community grazing rates in March, but in Junewhen the water column was mixed, vertical distribution of zooplanktonbiomass was important. Zooplankton grazing was an importantloss process for phytoplankton in the lower part of the epilimnionin Lake Rotongaio.     

The diel migrations and vertical distributions of zooplankton and micronekton in the Northwestern Mediterranean Sea. 1. Euphausiids, mysids, decapods and fishes

Vertical distributions and diel migrations of the main speciesof micronekton, four euphausiids, one mysid, one decapod andthree fishes, were described in detail in the 0–1000 mwater column on a fixed station in the Northwestern MediterraneanSea. The euphausiids Euphausia krohni and Thysanopoda aequalis,the decapod Gennadas elegans and, to a lesser extent, the fishArgyropelecus hemigymnus were shown to perform clear diel verticalmigrations. Results of horizontal hauls at a given depth aroundsunrise and sunset showed a marked diurnal symmetry of the migratorycycles, particularly for E.krohni, T.aequalis and G.elegans.The behaviour of the euphausiid Nematoscelis megalops was morecomplex: it presented a repetitive bimodal day distributionand only part of its population migrated. As very weak or non-migratorswe found the euphausiid Stylocheiron longicorne and the bathypelagicmysid Eucopia unguiculata, for which migration concerned onlysome of the older individuals. The fishes Cyclothone braueriand Cyclothone pygmaea appeared to be non-migrants. As depthincreased, C.braueri was replaced by C.pygmaea, with maximumconcentrations at 350–550 and 550–700 m depth, respectively.     

The importance of macrophyte bed size for cladoceran composition and horizontal migration in a shallow lake

Cladoceran composition and diel horizontal migration were studiedin 2, 10 and 25 m diameter macrophyte exclosures establishedin the littoral zone of shallow Lake Stigsholm, Denmark. Theexclosures were protected from waterfowl grazing, but open tofish. The macrophyte community cornprized Potwnogeton pectinatus,Potamogeton pusillus and Callitriche hemaphroditica. Cladoceranswere sampled randomly every third hour inside and outside themacrophyte exclosures during a 24 h period. In the 2m exclosure,the pelagic species Ceriodaphnia spp. and Bosmina spp. dominatedduring the day, mean density being as high as 3430 indiv. l^–1During the night, density decreased to 10–20% of the daytimedensity thus indicating diel horizontal migration. In the 10and 25 m exciosures, the daytime mean density of Ceriodaphniaspp. was 865 and 202 indiv. l^–1, respectively, and didnot decrease at night. In contrast to the pelagic species, thedensity of macrophyte-associated species tended to be higherin the 10 and 25 m exclosure than in the 2 m exclosure. In thedaytime, Eurycercus lamellatus density in the 2, 10 and 25 rnmacrophyte exclosures was 7, 28 and 16 indiv. l^–1 respectively,while that of Simocephalus vetulus was 11, 171 and 92 mdiv.l^–1, respectively. There was no thy-night difference inthe density of macrophyte-associated species. We conclude thatcladoceran community composition varies with macrophyte bedsize, and that the edge zone between the bed and open wateris an important daytime refuge for potentially migrating pelagiccladocerans.