Diel feeding behavior of neritic copepods during spring and fall blooms in Akkeshi Bay,eastern coast of Hokkaido,Japan

In situ diel feeding behavior of neritic copepods was investigated using the gut fluorescence method, during spring and fall bloom periods in Akkeshi Bay, on the eastern coast of Hokkaido, Japan. Acartia omorii and Paracalanus sp. were the dominant species during the fall, and Pseudocalanus spp. and A. longiremis during the spring. During both bloom periods, diel rhythms were always observed for the gut pigment contents of these dominant copepods, although there were interspecific differences in the pattern. The maximum gut pigment content was always observed during the night and the minimum during the day. For all species, except Paracalanus sp., the average gut pigment content during the night was significantly higher (p<0.05) than during daytime by factors of between 1.5 and 2.7. There were no significant differences between the gut evacuation rate constants determined during the day and the night, and initial gut pigment content had no effect on the value of gut evacuation rate constants. The instantaneous ingestion rates of individual copepods calculated from gut pigment and the mean value of gut evacuation rate constants followed the same diel rhythms as gut pigment contents. Copepod daily ingestion rates were higher than the daily requirements for respiration during both bloom periods. Estimated daily ration was 40 to 91% of body carbon during the fall bloom, and 17 to 28% during the spring bloom. The higher daily rations during fall were probably due to the difference in in situ temperature (ca. 14°C).     

In situ feeding rhythms of herbivorous copepods,and the effect of starvation

In situ diel variations in gut pigment contents of neritic (Acartia omorii andPseudocalanus minutus) and oceanic copepods (Calanus plumchrus andC. cristatus) were analyzed.A. omorii andP. minutus were sampled in Onagawa Bay on the east coast of Japan in May and August 1987, andC. plumchrus andC. cristatus were sampled in the Bering Sea in June 1986. Gut pigments were generally high at night, and bimodal feeding rhythms were observed in all species. The first peak of gut pigments occurred between sunset and midnight and was followed by a midnight decrease in gut pigment levels, resulting in eventual evacuation of the gut. The second peak was observed a few hours after sunrise. Incubation experiments indicated that ingestion rates of starved copepods were higher than those of acclimated copepods. This phenomenon was most notable at high food concentrations. Gut pigments of starved copepods rapidly increased after exposure to high concentrations of culturedThalassiosira decipiens. These findings suggest that in situ feeding behavior of herbivorous copepods includes periods of cessation or reduction in feeding during the night, and consequently, feeding activity is periodically enhanced with starvation. Starvation enhanced feeding behavior is most obvious in the large oceanic species,C. plumchrus andC. cristatus and is not distinct in small coastal species such asA. omorii.     

Spatial and temporal feeding patterns of copepods from the North Pacific centra gyre

Spatial and temporal feeding patterns (determined from an index of gut fullness) are described for 10 typical species of calanoid copepods collected from the North Pacific central gyre (September 1968 to June 1977), an area where the zooplankton is food limited and there were a-priori reasons to suspect that feeding and competition for food were important in regulating zooplankton community structure. Over 100 samples from 11 cruises to the eastern part of the gyre were examined, and patterns of gut fullness were related to environmental variables and the copepod species structure. The copepods studied all tended to be omnivores and food generalists. Males had lower indices of gut fullness than females but both males and females of a species had similar spatial and temporal feeding patterns. Guts were usually fuller at night than during the day, even in nonmigrating species; however, within nighttime depth distributions, no depths were preferred for feeding. There were also differences between species in mean gut fullness, but different species tended to have similar spatial and temporal feeding patterns. There was considerable spatial variability, and locales could be identified in which most species had higher indices of gut fullness. The copepods were not necessarily more abundant in these locales, nor did these tend to be areas of above average chlorophyll concentration. These patterns were consistent with relatively nonselective feeding, and there was no evidence that these species separate their niches by feeding at differing places or times.     

Energetic costs of swarming behavior for the copepod Dioithona oculata

The cyclopoid copepod Dioithona oculata forms dense swarms within shafts of sunlight that penetrate the mangrove prop-root habitat of islands off the coast of Belize. Previous studies, based on in situ video recordings and laboratory studies, have shown that D. oculata is capable of maintaining fixed-position swarms in spite of currents of up to 2 cm s⁻¹. The purpose of this study was to examine the energetic costs of maintaining these swarms, in terms of increased metabolic costs of maintaining position in currents and in terms of reduced feeding rates in densely packed swarms during the day. Using a sealed, variable-speed flow-through chamber, the respiration rates of D. oculata were measured while swarms maintained position in different current speeds. The results indicate that active metabolism (swimming at maximum speed to maintain the swarm in a current) is approximately three times greater than routine metabolism (normal swimming speeds in the absence of currents), indicating a significant metabolic cost of maintaining swarms in the presence of currents. In addition, gut-pigment analysis indicated that feeding rates of these copepods were often reduced in swarms during the day compared to when the copepods were dispersed at night. Given the high “cost” of swarming, the adaptive value of swarming in terms of reduced predation, increased opportunities for mating, and reduced dispersal, must be substantial. Received: 4 June 1997 / Accepted: 18 September 1997     

Diel vertical migration and feeding patterns of Mysis mixta (Crustacea,Mysidacea) in the Baltic Sea

The extent of the nocturnal vertical migration of Mysis mixta Lilljeborg varied between early July and late October (of 1985 and 1986) in a coastal area of the Baltic Sea. Migration was more restricted in early July and late October. Seasonal changes in surface light levels and transparency were sufficient to explain the observed differences. Mysids avoided light levels above 10^-4 lux throughout the study period. Smaller juveniles migrated higher up than larger juveniles and adults. A two-layered distribution with part of the population close to the bottom was observed at night. Zooplankton were more abundant in water layers above the main concentration of mysids. M. mixta fed on phytoplankton, detritus, copepods, cladocerans, rotifers and tintinnids. Diel changes in gut fluorescence indicated a higher intake of phytoplankton at night, but levels were low compared to primarily herbivorous zooplankton. Comparisons of stomach contents of mysids caught at the bottom in the evening and in the water column at night showed a higher ingestion of zooplankton at night and of detritus during the day. Mysids caught at the bottom at night had an intermediate diet. Copepods and cladocerans constituted between 90 and 100% of ingested material by weight in all mysid groups.     

Gut pigment accumulation and destruction by arctic copepods in vitro and in situ

The results presented here were obtained at six locations during three cruises in 1985 (off the coast of Labrador), 1986 (at the eastern end of Viscount Melbourne Sound) and 1988 (off the coast of Labrador). In situ chlorophyll maximum concentrations were >7 gl^-1 at depths of between 0 and 30 m in all sampling areas. In feeding experiments copepods attained higher gut pigment concentrations the longer they had been previously starved and higher concentrations when fed in the dark than when fed in the light. Community ingestion rates calculated from changes in particulate chlorophyll were higher than estimates derived from gut pigment data except when copepods had been starved for 24 h. Differences between estimates by the two methods suggested pigment destruction. In feeding experiments pigment: biogenic silica ratios in food and faecal pellets suggested that the length of starvation period affected the degree of pigment destruction differently at different stations and that feeding in the light greatly increased pigment destruction. A comparison of pigment: silica ratios in the water column, and in faecal pellets collected from copepods which had fed there, suggested that pigment destruction may occur in situ sometimes and that the degree to which it occurs may be affected by feeding history, light, diel feeding behaviour and species composition.     

Inter- and intra-specific diurnal habitat selection of zooplankton during the spring bloom observed by Video Plankton Recorder

Diel vertical migration (DVM) is a common behavior adopted by zooplankton species. DVM is a prominent adaptation for avoiding visual predation during daylight hours and still being able to feed on surface phytoplankton blooms during night. Here, we report on a DVM study using a Video Plankton Recorder (VPR), a tool that allows mapping of vertical zooplankton distributions with a far greater spatial resolution than conventional zooplankton nets. The study took place over a full day–night cycle in Disko Bay, Greenland, during the peak of the phytoplankton spring bloom. The sampling revealed a large abundance of copepods performing DVM (up during night and down during day). Migration behavior was expressed differently among the abundant groups with either a strong DVM (euphausiids), an absence of DVM (i.e., permanently deep; ostracods) or a marked DVM, driven by strong surface avoidance during the day and more variable depth preferences at night (Calanus spp.). The precise individual depth position provided by the VPR allowed us to conclude that the escape from surface waters during daytime reduces feeding opportunities but also lowers the risk of predation (by reducing the light exposure) and thereby is likely to influence both state (hunger, weight and stage) and survival. The results suggest that the copepods select day and night time habitats with similar light levels (~10^?9 μmol photon s^?1 m^?2). Furthermore, Calanus spp. displayed state-dependent behavior, with DVM most apparent for smaller individuals, and a deeper residence depth for the larger individuals.     

Comparison of gut-evacuation rates of feeding and non-feeding Calanus marshallae

Gut-evacuation rates were compared for feeding and non-feeding Calanus marshallae collected near Yaquina Bay, Oregon (44°37N; 124°04W) from June 1986 through May 1987. Evacuation rates were measured at four concentrations of Thalassiosira weissflogii from the decline of gut-pigment fluorescence following transfer of copepods to filtered seawater and from the loss of cells labelled with 68-germanium, a radioactive analog of silicon. There was no significant difference between gut-evacuation rates of feeding and non-feeding copepods over both short-term (20 min) and long-term (90 min) evacuation times. Furthermore there was no difference between rates obtained using either pigment or ⁶⁸Ge as tracers of gut food-passage. These results are discussed in the light of possible dilution of tracer in the guts of feeding copepods due to mixing with unlabelled food. Gut-evacuation rates measured at food concentrations ranging from 500 to 4000 cells ml^-1 were not significantly different, regardless of the technique employed.     

Der einfluß des lichtes auf Körperfärbung und dämmerungsverhalten des korallenfisches Chaetodon melanotus

The influence of light on colour patterns and behaviour of the coral-reef fish Chaetodon melanotus were investigated under aquarium and field conditions. Colour pattern and fin position are correlated with agonistic behaviour: they were used as criterion for analysing the behavioural patterns during morning and evening transition.Readiness for attack is correlated with a darkening of the analfin. A specimen with yellow anal fin and spread dorsal fin is more likely to flee. The colour patterns vary during day, twilight, and night time. The speed of colour change was measured experimentally; it appears to be hormonally regulated. During twilight, 5 different colour patterns are related with certain anal-fin colour-patterns and dorsal-fin positions. Swimming activity decreases with reduction in irradiance, while the readiness to flee increases. In groups, specimens become more territorial with diminishing day light. Territorial aggression can be light-dependent; no territorial aggression was observed at illuminations less than 50 lux. Behaviour and concomitant coloration-changes during the evening and morning transition periods are interpreted as adaptational protections against predators. The remarkable white eyespots in the twilight body-coloration could represent a warning signal, and could also have a somatolytic effect. The increased readiness to flee during dusk and dawn may be a behavioural mechanism for reducing predator pressure.     

Nocturnal movements of phototactic zooplankton in shallow waters

Downwardly directed light-traps were used by night, over seagrass and coral sand to investigate the existence and significance of nocturnal peaks in phototactic zooplankton. Samples each half-hour from sunset to sunrise showed that small invertebrates, mostly copepods, were trapped in large numbers once or twice a night. Each period of high catches lasted only 1 to 2 h and was often associated with periods of failing natural light (evening twilight, moonset). This pattern and its variations are explained in terms of nocturnal vertical migration, in which plankton move towards light in response to decreasing light intensity, move away from light in response to increasing light intensity, and disperse at constant light intensity. Some morning peaks are unexplained.     

Phytoplankton pigments in the gut concents of planktonic copepods from coastal waters off southern California

Phytoplankton xanthophylls in the gut contents of the copepods Calanus pacificus, Corycaeus anglicus, and Paracalanus parvus, collected from 5 stations off San Onofre, California, in June 1982, were measured by reverse phase, high-performance liquid chromatography (HPLC). The dinoflagellate pigment, peridinin, was usually the most abundant xanthophyll in the guts of all three species of copepods. Evidently, feeding was principally on dinoflagellates (which dominated the phytoplankton biomass). The level of feeding activity, rather than the class of phytoplankton ingested, seemed to differentiate the behaviors of the copepods. Xanthophyll content per unit copepod wet weight was higher in Corycaeus anglicus and Paracalanus parvus than in Calanus pacificus. Chlorophyll a fluorescence of the copepod gut contents was measured in conjunction with the analysis of gut xanthophylls. The xanthophyll content of the gut varied directly with the concentration of chlorophyll a in the gut. Xanthophyll content was not related to the concentration of pheopigments in the gut. Apparently, the xanthophylls that were detected were due to the presence of recently ingested phytoplankton biomass.     

Diel trophic interactions between vertically-migrating zooplankton and their fish predators in an eelgrass community

Diel changes in the composition of crustacean zooplankton and the diets of fish predators from an intertidal eelgrass flat were monitored concurrently. The zooplankton is characterized by two major components. The obligate zooplankters (holoplanktonic calanoid copepods and meroplanktonic decapod larvae) appear to exhibit vertical migration, being present in higher densities near the surface of the water column at night. The facultative zooplankton (amphipods and ostracods) are benthic during the day, but move up into the water column at night. Planktivorous midwaterdwelling fish consume calanoid copepods and decapod larvae during the day and cease feeding or switch their diet to amphipods at night. Benthic-dwelling fish consume some amphipods during both day and night. The factors important in prey selection by fish and the functional significance of vertical migration in both components of the zooplankton are discussed in the light of the changing patterns of fish predation.This paper is Publication No. 183 in the Ministry for Conservation of Victoria, Environmental Studies Series.     

Diel variability of feeding activity in haddock (Melanogrammus aeglifinus) larvae in the East Shetland area, North Sea

Investigations of factors affecting feeding success in fish larvae require knowledge of the scales of variability of the feeding process itself and the indices used to assess this variability. In this study, we measured short-term (diel) variability in feeding rates of wild haddock (Melanogrammus aeglifinus) larvae four times per day during a 10-d cruise in the northern North Sea. Feeding activity was evaluated using indices of gut fullness, prey digestive state and biochemical measurements (tryptic enzyme activity). The gut fullness and the enzyme activity indices indicated moderate to high rates of food consumption throughout the cruise. Time series analysis of the three indices showed significant diel variability in all indices and enabled identification of significant lags between food uptake and peak digestive enzyme activity. The typical pattern of food consumption and digestion was characterized by maximal ingestion of prey early in the evening (19:00 hrs) and peak digestive enzyme activity at 01:00  hrs. The time scale over which enzyme activities reacted to prey ingestion was ca. 6 h, and is consistent with expectations from controlled laboratory experiments with other larval fish species. Significant diel variability in tryptic enzyme activity suggests that attempts to relate this measure of feeding success to other variables (e.g. food concentrations) should take care to accommodate natural cycles in feeding activity before making statistical comparisons. Received: 29 October 1998 / Accepted: 18 June 1999     

Osmoacclimation in Enteromorpha intestinalis: long-term effects of osmotic stress on organic solute accumulation

The fate of the protease trypsin in intestines of individual herring larvae Clupea harengus L. was studied following digestion of the copepod Acartia tonsa. Trypsin was retained in the intestine during two consecutive pulses of feeding and defaecation of copepods. Quantification of herring trypsin in digested, defaecated copepods showed that ca. 1% of larval intestinal enzyme was defaecated along with 1 to 3 copepods. Following ingestion of a single meal, the level of intestinal trypsin post-ingestion declined to pre-ingestion levels within 1 to 2 d of starvation. All enzyme data thus indicated that trypsin, released in response to ingestion of a meal, was retained. In addition, analysis of fed subgroups of starved larvae clearly indicated that release of trypsin from the pancreas stopped after 6 to 8 d of starvation. As the fish still contained substantial amounts of trypsinogen, the underlying cause might be defective release mechanisms. Daily secretion of trypsin and processes responsible for enzyme retention in the gut are discussed. Assimilation efficiency in herring larvae was estimated for copepodite prey. Average carbon assimilation was 90%.     

Comparison of four methods to estimate meiobenthic copepod Amonardia normani ingestion rates

Four different methods were used in the control conditions of laboratory to estimate the ingestion rate of a female meiobenthic harpacticoid copepod Amonardia normani: (1) reduction of algal biomass, (2) the quantification of total pigments in fecal pellets, (3) the gut fluorescence method, (4) the percentage of assimilation and the total egestion rate. The food used during all experiments was the diatom Nitzschia constricta in an axenic condition at the concentration of 0.13 μg Chl-a mL^?1 at stationary growth phase. All experiments were made at 20 °C and 30 salinity. All tested methods excepted the quantification of total pigments in fecal pellets resulted in similar estimatives. The gut fluorescence method indicated that during the day gut contents are smaller than during the night but the gut passage time was faster, resulting in similar ingestion rates during the day and the night. The reduction of algal biomass and the percentage of assimilation and the total egestion rate also indicated similar ingestion rates in the day and in the night. The daily ingestion rate represents 107 % of female carbon weight per day (903 ng C cop^?1day^?1).     

Effect of microclimatic factors and tide on vertical migrations of the mangrove crab Sesarma leptosoma (Decapoda: Grapsidae)

The tree crab Sesarma leptosoma Hilgendorf migrates up the mangrove trunks twice a day, both in the morning and in the evening, leaving its refuge near the tree base to feed on fresh leaves. In spite of the 150 cm of tide excursion at the base of the trees, the timing of the migrations seems largely controlled by nontidal factors. The timing of the morning upward migration is controlled by the light level alone. The morning downward migration takes place as soon as the temperature rises and relative humidity falls, with no relationship to tidal cycle or light level. Only the evening upward flow seems primarily controlled by the tide, while the evening downward flow, which ends at twilight, seems to be related to the timing of the evening upward migration, to the tide and again to light levels. During the day, crabs abandon the canopy to avoid low relative humidity, usually coupled with the higher wind speeds, which would quickly dehydrate them. At night, crabs stop migrating, probably because their movements and orientation capability along the trunk and in the canopy are largely vision dependent. Nontidal factors have never previously been shown to influence the migratory behaviour of an intertidal crustacean species so extensively. Received: 14 June 1997 / Accepted: 17 July 1997     

Feeding in Arctic darkness: mid-winter diet of the pelagic amphipods Themisto abyssorum and T. libellula

The pelagic amphipods Themisto abyssorum and Themisto libellula represent important links between the herbivore zooplankton community and higher trophic levels of the Arctic marine food webs. Large double structured eyes of both of these hyperiid species are assumed to be used for visual prey detection. However, no information is available on the feeding strategies of these visually searching predators for the period of the polar night, a time of year with no or very low levels of daylight. Here, we report on the stomach and gut content of both Themisto species collected during a January expedition around Svalbard (78° to 81°N). Results indicate that T. abyssorum and T. libellula feed actively during the Arctic winter. The major food source of both amphipods consisted of calanoid copepods, most frequently Calanus finmarchicus.     

The feeding habits of three species of lanternfishes (family myctophidae) off oregon,USA

The common myctophids Stenobrachius leucopsarus, Diaphus theta and Tarletonbeania crenularis were found to feed primarily on the euphausiid Euphausia pacifica, the copepods Metridia lucens and Calanus spp., and the amphipod Parathemisto pacifica. The diets of these species of fishes were diverse and overlapped broadly, suggesting that they are feeding generalists. Most stomachs contained either all copepods or all euphausiids. Euphausiids were the most important prey on the basis of biomass. They comprised over one-half the weight of the stomach contents in over 40% of individuals of each of the three fishes. Stomach fullness and state of digestion of stomach contents differed over the diel period, but not enough to indicate feeding only during the nighttime. Average stomach fullness was greatest during the morning and night and well-digested material, which predominated in most stomachs, was most prevalent in the morning and afternoon. Either some feeding occurs throughout the day or digestion rates are slow.     

Observations on the nocturnal feeding of some mesopelagic decapod crustacea

Feeding in relation to temporal changes in the depth distribution of predator and prey is described for 9 species of mesopelagic decapods from an examination of 268 foreguts. Intensive nighttime feeding appears to be the rule in all species. The smaller decapods Sergestes (Sergestes) atlanticus, Sergestes (Sergestes) sargassi and Sergestes (Sergestes) pectinatus exploit the smaller prey, principally copepods and to a lesser extent ostracods. Larger decapod species Sergestes (Sergestes) henseni, Sergestes (Sergestes) curvatus, Sergestes (Sergia) grandis, Systellaspis debilis, and Acanthephyra purpurea mainly prey on macrozooplankton and micronekton, i.e., chaetognaths, euphausiids, decapods and fish, but copepods also occur in the foreguts. Gennadas valens is exceptional for the high incidence of foraminiferal remains, and a predator-prey relationship seems probable. All 9 decapod species have mixed diets, and pronounced feeding preferences are not evident. However, a high incidence of “secondary” feeding or “dietary contamination” has been deduced from the frequent occurrence of remains of the copepods Pleuromamma spp. and Oncaea spp. in the foreguts of the larger decapod species. Direct feeding cannot have occurred, since the depth distributions of these copepods and decapods are disjunct by day and night. It is concluded that the remains of Pleuromamma probably represent the food of the larger prey such as chaetognaths etc. which are eaten by the decapods. The presence of Oncaea is speculatively attributed to a possible ectoparasitic relationship with the larger prey items, but confirmatory evidence is required. These anomalies suggest that caution must be exercised in deducing predator-prey relationships simply from gut contents without consideration of distributional factors.     

Relationship between the zooplankton,phytoplankton, particulate matter and dissolved free amino acids in the Celtic Sea

Estimates of the biomass of zooplankton, phytoplankton and particulate matter collected in the Celtic Sea during mixed-water conditions (on 8 and 9 April 1983) were compared to the concentration and diversity of sixteen dissolved free amino acids (DFAA) measured in seawater and in particles. During a day profile, variations of dissolved amino acids with depth reflected the feeding activity of copepods. The relationship was not apparent in a night profile and other processes, such as heterotrophic utilization of dissolved nitrogen by microorganisms, were thought to be involved. The ratios of total DFAA concentration (nM litre^-1) in the particulate phase over the concentration in seawater ranged from 1 to 200 within the water column. Of the sixteen amino acids measured, ornithine, a decomposition product of arginine, was responsible for more than 70% of the total concentration of DFAA in seawater. In the particles, phenylalanine ranged from 30 to 88% of total DFAA. In seawater this amino acid occurred in the 20 to 40 m depths (1.3 to 9.9% of total DFAA) in the day profile and at 5 m (12.4%) and 80 m (6.4%) in the night profile. Previously it has only been found in very low concentrations (<5%) in seawater, and its presence is considered to be the result of zooplankton feeding.