Does Acartia clausi (Copepoda: Calanoida) use an area-restricted search foraging strategy to find food?

The swimming behavior of Acartia clausi within small aquaria containing phytoplankton was videotaped, along with unfed controls. From these videos, aspects such as swimming speed, distance traveled, and 2D (horizontal, X, and vertical, Z) headings were then measured from over 75000 X Z positional data in order to quantify their swimming functional response. There was generally no significant difference in swimming behavior between different food levels, although there was a large difference between feeding and unfed controls. Because feeding bouts moved the copepods a short distance at a slow speed, net vertical displacement decreased as feeding-bout frequency increased. Non-feeding individuals showed longer periods of sinking, interspersed with longer-distance, high-speed jumps, which displaced them farther distances than filter-feeding copepods over the same time interval. For both fed and unfed control copepods, a 1D unbiased random walk was adequate to describe their displacement over these very short time periods (10 s). This behavior is consistent with an area-restricted search foraging strategy. The net vertical displacements predicted by combining the experimental data with a 1D random walk model suggest that A. clausi would be retained within even small 10 cm phytoplankton patches long enough to fill their gut. However, copepods outside of phytoplankton patches must rely on other means to find patches of food greater than 50 cm apart. Potentially, knowledge of a copepod's swimming functional response could enable predictions about the typical spatial and temporal patchiness of its food in situ.     

Food and feeding of aquatic larvae of the midge chaoborus flavicans (meigen) (diptera : chaoboridae) in the laboratory

Laboratory observations on the feeding of fourth instar larvae of Chaoborus flavicans (Meigen) show that features relating to the prey, such as swimming behaviour, size and shape, play a major role in determining the kind of food eaten by the predator.In tests with two interacting limnetic cyclopoid copepods, Cyclops abyssorum out-competed Mesocyclops leuckarti when Chaoborus was absent but did not do so when Chaoborus was present.     

Resisting flow–laboratory study of rheotaxis of the estuarine copepod Pseudodiaptomus annandalei

Rheotaxis is a ubiquitous phenomenon among aquatic animals and thought to be an adaptation to maintain populations in flowing waters. While many estuarine copepods can retain their populations in estuaries with net seaward flow, rheotaxis of individual copepods has not been reported before. In this study, the behavior of a calanoid copepod Pseudodiaptomus annandalei in flow was examined in a recirculating laboratory flume. This estuarine copepod displayed different responses to ambient flow fields while swimming in the water column or attaching to the flume bed (walls). Copepods in the water column showed vigorous countercurrent swimming by occasional bounding when flow velocity was increased up to 2.1 cm s^?1, but none of the individuals in the water column were retained in the flume when flow speeds were higher than 4 cm s^?1. This indicates P. annandalei profits little from rheotaxis to withstand flow when they were swimming in the water column. Instead, more individuals attempted sinking downwards to the slow flow region near the flume bed (walls) and showed active substrate attachment to avoid being flushed out by the high-velocity channel flow. The results suggest that P. annandalei benefits from rheotaxis and association with the substrate which allows them to hold position well at ambient flow velocities up to 3 cm s^?1. These adaptive responses might be important for population maintenance.     

Effects of animal density,volume, and the use of 2D/3D recording on behavioral studies of copepods

Studies on the behavior of copepods require both an appropriate experimental design and the means to perform objectively verifiable numerical analysis. Despite the growing number of publications on copepod behavior, it has been difficult to compare these studies. In this study, we studied two species of copepods, Eurytemora affinis and Pseudodiaptomus annandalei, and employed recently developed scaling and non-scaling methodology to investigate the effects of density and volume on the swimming behavior of individual organisms in still water. We also compared the results of two- and three-dimensional projections of the swimming tracks. A combination of scale-dependent and scale-independent analysis was found to characterize a number of behavioral observations very effectively. We discovered that (i) density has no effect except to increase the time spent in the swimming state of “breaking”, (ii) smaller volumes resulted in more complex trajectories, and larger volumes, like density, increased the time spent in the swimming state “breaking”, and (iii) three-dimensional projections gave a more accurate estimation of speed and the time spent cruising. When only a vertical 2D projection was used, “cruising” could be confused with “sinking”. These results indicate that both experimental conditions and the selection of 2D or 3D projection have important implications regarding the study of copepod behavior. The development of standardized procedures with which to compare the observations made in different studies is an issue of particular urgency.     

Diel and seasonal vertical distribution of meiobenthic copepods in muddy sediments of a eutrophic lagoon (fish ponds of Arcachon Bay)

The vertical distribution of meiobenthic copepods was investigated within muddy sediments of a eutrophic lagoon (fish ponds of Arcachon Bay, France). The aim of the study was to determine if in muddy sediments, as previously established in sandy sediments, meiobenthic copepods migrate vertically according to the seasons or diel periods. Two experimental approaches were used, viz: a three-season comparison was made of the diel vertical distribution of the harpacticoid Canuella perplexa T. & A. Scott (1893) and secondly the depth distribution of a meiobenthic copepod assemblage was followed for a 24 h period, in shallow water subtidal locations. The harpacticoid C. perplexa vertically migrated through the top three centimeters of the sediment, showing diel and seasonal variations in depth distribution. The differential vertical distributions shown by the dominant meiobenthic populations suggest that emergence into the water column may mainly concern surface dwelling copepods. The physical and biological factors affecting seasonal and diel changes in the copepod assemblage of the fish ponds are discussed.     

Latitudinal differentiation in the effects of the toxic dinoflagellate Alexandrium spp. on the feeding and reproduction of populations of the copepod Acartia hudsonica

Blooms of the dinoflagellate Alexandrium spp. increase in their frequency, toxicity and historical presence with increasing latitude from New Jersey (USA) to the Gaspé peninsula (Canada). Biogeographic variation in these blooms results in differential exposure of geographically separate copepod populations to toxic Alexandrium. We hypothesize that the ability of copepods to feed and reproduce on toxic Alexandrium should be higher in copepods from regions that are frequently exposed to toxic Alexandrium blooms. We tested this hypothesis with factorial common environment experiments in which female adults of the copepod Acartia hudsonica from five separate populations ranging from New Jersey to New Brunswick were fed toxic and non-toxic strains of Alexandrium, and the non-toxic flagellate Tetraselmis sp. Consistent with the hypothesis, when fed toxic Alexandrium we observed significantly higher ingestion and egg production rates in the copepods historically exposed to toxic Alexandrium blooms relative to copepods from regions in which Alexandrium is rare or absent. Such differences among copepod populations were not observed when copepods were fed non-toxic Alexandrium or Tetraselmis sp. These results were also supported by assays in which copepods from populations both historically exposed and naïve to toxic Alexandrium blooms were fed mixtures of toxic Alexandrium and Tetraselmis sp. Two-week long experiments demonstrated that when copepods from populations naïve to toxic Alexandrium were fed a toxic strain of Alexandrium they failed to acclimate, such that their ingestion rates remained low throughout the entire two-week period. The differences observed among populations suggest that local adaptation of populations of A. hudsonica from Massachusetts (USA) to New Brunswick (Canada) has occurred, such that some populations are resistant to toxic Alexandrium.     

The copepod Centropages abdominalis as a carrier of the stalked ciliate Zoothamnium

Adult specimens of the calanoid copepod Centropages abdominalis collected from February through April 1986 in Shinhamako, a saline lake connected to Tokyo Bay, were heavily infested with the stalked ciliate Zoothamnium. The number of copepods and those infested varied during the infestation period; in February when the copepod population was large, the incidence of infestation was low, whereas in March when the copepod population was small, copepods infested with Zoothamnium were proportionally more abundant. Numerical variations of infested copepods may correspond to changes in the Zoothamnium population. No specimens of Acartia omorii, which coexists with C. abdominalis, were infested in 1986, although a small number of A. omorii were infested with peritrich ciliates in 1985. Some shrimp larvae were also covered with the same peritrichs as copepods. Nevertheless, as a whole, the relationship between the phoront and its carrier seems to be specific. The highest incidence of infested copepods was approximately 30% similar to that in 1985. The sex ratio of C. abdominalis changed on each sampling date but in general females were more numerous than males. The percentage of infested males was higher than that of infested females, suggesting that the former can be attached to more easily than the latter.     

Factors affecting feeding selectivity of visual predators on the copepod Acartia tonsa: locomotion,visibility and escape responses

Visual predation by fish on copepods involves prey encounter, attack and capture; during any of these processes prey selection can occur. Developmental changes in copepods, including increases in swimming speed, size and image contrast increase the encounter rate and distance at which they can be detected by predators. Copepods compensate for this increase vulnerability with age through diel vertical migration and improved escape capabilities. This study quantifies the changes in swimming speed and movement pattern with developmental stage of the copepod Acartia tonsa, using a video-computer system for motion analysis. Changes in visible size and image contrast with developmental stage were quantified under simulated natural illumination conditions using a video based image analysis system. The escape responses of the naupliar stages of the copepod Acartia tonsa were quantified in response to a stationary pipette sucking in water at a constant speed. Accurate quantification of the parameters that affect feeding selectivity of planktivorous fish will provide the basis for evaluation of their relative importance in future studies.     

Diel changes in vertical distribution of Oithona similis (Cyclopoida) and Oncaea curvata (Poecilostomatoida) under sea ice in mid-summer near Syowa Station, Antarctica

The diel vertical distributions of two small copepods, Oithona similis and Oncaea curvata, were investigated at 4-h intervals over a 24-h period under fast ice near Syowa Station during continuous daylight conditions in the Antarctic mid-summer, December 1993. Oithona similis and O. curvata exhibited small-scale diel vertical migrations during the study period, in a way opposite to what is expected, i.e., remaining mostly in the upper layer during the day and moving into deeper layers at night. The nighttime descent of both species coincided with the time of disappearance of a high algal concentration at the ice–water interface during the day and an increase of the algal concentration in the mid-water layer at night. This suggests the migration behavior of the copepods was responsible for the change of food availability. The daily grazing impact of these copepods was estimated to remove one-third of the algae daily released from ice during mid-summer at Syowa Station.     

Annual changes in the population size of the salmon louse Lepeophtheirus salmonis (Copepoda: Caligidae) on high-seas Pacific Salmon (Oncorhynchus spp.), and relationship to host abundance

The population size of the salmon louse, Lepeophtheirus salmonis, was monitored annually in the summers of 1991–1997 by examining six species of Pacific salmon (Oncorhynchus spp.) caught by surface long-lines in oceanic offshore waters of the North Pacific Ocean and Bering Sea. The annual copepod population size on all salmonids caught was estimated by combining the calculated number of copepods carrying on each salmonid species. The copepod population fluctuated markedly from year to year, which resulted largely from marked annual changes in abundance of pink salmon (O. gorbuscha). Since pink salmon were most frequently and heavily infected and since their abundance changed every year, the copepod population was high in the years when this salmonid species was abundant, but low when it was rare. On the contrary, chum salmon (O. keta) did not show high prevalence and intensity of infection, but the annual abundance of this host species was consistently high, i.e. chum salmon carried many copepods every year. Copepods on other salmonid species (sockeye salmon O. nerka, coho salmon O. kisutch, chinook salmon O. tshawytscha, and steelhead trout O. mykiss) constantly formed a small percentage of the total copepod population. Both chum and pink salmon are the most important hosts in terms of their substantial contribution to support the copepod population, but the importance as hosts of each species is definitely different between the species. Chum salmon is a stable important host supporting the copepod population at a relatively high level every year, while the number of copepods on pink salmon annually exhibits marked fluctuations, and this salmonid species is regarded as an unstable important host.     

Effect of body mass and water temperature on the standard metabolic rate of juvenile yellow perch, Perca flavescens (Mitchill)

Fish respiration rates that are presumed to represent standard metabolic rates (SMR) may sometimes include an unspecified energy expenditure associated with activity and digestion. This situation may introduce a bias in bioenergetics models because standard metabolism, digestion, and activity may not be affected by the same environmental conditions. The aim of this study was to (1) develop a SMR model for juvenile yellow perch, Perca flavescens (Mitchill), that represent the minimum energy expenditure required to maintain life and (2) compare the results of this study with published perch metabolic rates and bioenergetics models. SMR was estimated for yellow perch over a range of body␣mass (4.4–24.7 g) and water temperature (12–20°C). The intercept of the relationship between fish respiration and swimming velocity obtained during forced swimming experiments was used to determine SMR. SMR estimated by the present study were comparable to values presented by two published studies on Eurasian perch, Perca fluviatilis L. However, estimated SMR were 4.1–20.9 times lower than values of a third respirometry study and predictions of bioenergetics models for perch. The present study suggests that published SMR models may sometimes include a significant fraction of energy expenditures (39.2–75.9%) associated with digestion and activity. This may complicate the implementation and the interpretation of fish bioenergetics models. The present study indicates that the intercept of respiration-velocity relationships and long-term respiration rates during starvation experiments may provide similar and reliable SMR values.     

Diel vertical migration by males,females, copepodids and nauplii in a limnetic population of diaptomus (Copepoda)

Diel vertical migration (DVM) by the sexes and life history stages of the copepod Diaptomus novamexicanus (Herrick) was investigated during two summers in a subalpine lake (Castle Lake, California, USA). Migratory behavior was quantified repeatedly by 1300 and 21–2200 hr sampling in the epilimnion. Population density and vertical distribution were estimated routinely at 1300 hr and at 2200 hr on selected dates. Males had greater night to day biomass differentials and ratios than females. The sexes showed comparable DVM participation (% of total biomass migrating) and amplitudes of migration (distance of ascent). Copepodids displayed higher participation, night/day ratios and amplitudes than younger copepods, but adult participation differed only from that of nauplii. Active participation in high amplitude ascents thus increased with age and was greatest in adult males. Analysis suggests that such relative behaviors probably occur in most limnetic Diaptomus, and that the magnitude of ascent into the epilimnion is associated with daytime vertical distribution.     

Toxicity of blooms of the cyanobacterium Trichodesmium to zooplankton

The marine filamentous bloom-forming cyanobacteria Trichodesmium thiebautii and T. erythraeum were collected at locations in the Carribean during Jan.–Feb. 1991. They were screened for toxicity using Artemia salina and several species of copepods, which were harpacticoid grazers, filter-feeding calanoids, or cyclopoid copepods. Approximately 50% of the 89 T. thiebautii samples caused> 50% lethality of A. salina, though none of the 16 T. erythraeum samples caused> 25% lethality. The T. thiebautii bloom samples were toxic to the calanoid and cyclopoid copepods and non-toxic to the harpacticoid grazers. In contrast the T. erythraeum bloom samples were not toxic to any copepods tested.     

Swimming Mode Inferred from Skeletal Proportions in the Fossil Pinnipeds Enaliarctos and Allodesmus (Mammalia, Carnivora)

Swimming modes are crucial for understanding evolutionary transitions from land to sea, because locomotion affects many aspects of an animal’s life. The modern pinniped families Otariidae (fur seals and sea lions), Phocidae (true seals), and Odobenidae (walruses) are thought to share a common origin, but each differs in its primary mode of aquatic locomotion. Previous studies of locomotor evolution in pinnipeds suggested: (1) forelimb swimming was ancestral; (2) hind limb swimming evolved once at the base of the clade including Phocidae, Odobenidae, and the extinct Desmatophocidae; and (3) reversal to forelimb swimming occurred in the odobenid subfamily Dusignathinae. The oldest and most basal pinnipedimorph Enaliarctos mealsi has been portrayed as a forelimb swimmer, and the desmatophocid Allodesmus kelloggi has been portrayed as a hind limb swimmer. These interpretations have been questioned by others and are tested here. Principal components analysis of trunk and limb measurements from 58 modern semiaquatic mammals demonstrates that Enaliarctos is most similar in skeletal proportions to hind limb-dominated swimmers, whereas Allodesmus is most similar to forelimb-dominated swimmers. Principal components and discriminant function analyses of trunk and limb measurements from 24 modern pinniped species demonstrate that Enaliarctos is most similar to hind limb-swimming phocids, while Allodesmus is most similar to forelimb-swimming otariids. These interpretations complicate previous portrayals of swimming evolution in pinnipeds and can paint a very different picture of how this behavior evolved when viewed in the context of alternative phylogenetic hypotheses.     

Relationship between feeding of blueback herring and the zooplankton community of a Texas reservoir

Following the introduction of blueback herring (Alosa aestivalis), we examined herring food habits and the crustacean zooplankton community in Lake Theo, a 30-hectare reservoir located on a tributary of the Red River in north Texas. Prior to the introduction in spring, 1982, the reservoir contained an established fish community dominated by centrarchids. Blueback herring stomachs and zooplankton were sampled quarterly from summer, 1982 through winter, 1985. Cladocerans accounted for 89.4% of the zooplankton consumed by herring. Median lengths of cladocerans and copepods in the herring diet were significantly greater than those in the reservoir. No appreciable changes in lengths of cladocerans and copepods in the reservoir were observed after blueback herring introduction, but the zooplankton community shifted from cladoceran to copepod domination.     

Mobiline peritrich riders on Australian calanoid copepods

Calanoid copepods from billabongs near Wodonga, Victoria, Australia were found to be infested with a disc-shaped mobiline peritrich ciliate belonging to the genus Trichodina. Biometrical data, including mean body diameter (43.6 m), mean adhesive disc diameter (35.9 m), mean denticle ring diameter (20.1 m), modal denticle number (17), modal number of radial pins per denticle (9), and denticle shape and dimensions, indicate that the species is T. diaptomi Basson & Van As (1991). This is the first record of T. diaptomi from Australasia. Adults of Boeckella fluvialis and B. minuta were infected, but adult Calamoecia lucasi were not. Levels of infestation were higher on B. fluvialis (67.5% of females, 54.5% of males) than B. minuta (47.4% of females, 33.3% of males). Mean numbers of Trichodina per adult B. fluvialis were 7.68 (range 0–78) for females and 4.06 (range 0–43) for males, and differed significantly between sexes. Although calanoid copepods were present in the plankton from January to early December 1991, Trichodina infestations occurred during October and November only, peaking during late November just before the copepods disappeared from the plankton. Whether the infestation was a cause of the copepod decline, developed because the copepods were rendered susceptible to infection by some other cause, or was determined by physical and/or chemical environmental factors is unknown.     

Feeding,egg production,and egg hatching success of the copepods Acartia tonsa and Temora longicornis on diets of the toxic diatom Pseudo-nitzschia multiseries and the non-toxic diatom Pseudo-nitzschia pungens

In 1987, there was an episode of shellfish poisoning in Canada with human fatalities caused by the diatom Pseudo-nitzschia multiseries, which produced the toxin domoic acid. In order to examine whether domoic acid in this diatom serves as a grazing deterrent for copepods, we compared feeding rates, egg production rates, egg hatching success and mortality of the calanoid copepods Acartia tonsa and Temora longicornis feeding on unialgal diets of the toxic diatom P. multiseries and the similarly-sized non-toxic diatom Pseudo-nitzschia pungens. Copepods were collected in summers of 1994, 1995 and 1996 from Shediac Bay, New Brunswick, Canada, near Prince Edward Island, the site of the 1987 episode of domoic acid shellfish poisoning. Rates of ingestion of the toxic versus the non-toxic diatom by A. tonsa and T. longicornis were similar, with only one significantly different pair of values obtained in 1994, for which A. tonsa had a higher mean rate of ingestion of the toxic than the non-toxic diatom. Thus, domoic acid did not appear to retard grazing. Analyses of copepods with high performance liquid chromatography (HPLC) revealed that copepods accumulated domoic acid when feeding on P. multiseries. Egg production rates of copepods when feeding on P. multiseries and P. pungens were very low, ranging from 0 to 2.79 eggs female^–1 d^–1. There did not appear to be differential egg production or egg hatching success on diets of the toxic and non-toxic diatoms. Mortality of females on the toxic diet was low, ranging from 0 to 20%, with a mean of 13%, and there was no apparent difference between mortality of copepods feeding on toxic versus non-toxic diatoms. Egg hatching success on both diets, although based on few eggs, ranged between 22% and 76%, with a mean percentage hatching of 45%. Diets of the non-toxic diatom plus natural seawater assemblages supplemented with dissolved domoic acid, revealed similar rates and percentages when compared to previous experiments. In summary, none of the variables measured indicated adverse effects on copepods feeding on the toxic compared to the non-toxic diatom.     

Seasonal changes in zooplankton abundance in the lower Rhine during 1987–1991

Zooplankton in the River Rhine was surveyed for five years at the Dutch sampling stations, Lobith (German/Dutch border) and Maassluis (at the point of discharge of the river into the North Sea). The zooplankton abundance showed an apparent seasonal pattern at both stations, characterized by low densities during the winter period, and higher densities during the summer period, with a spring peak. Zooplankton was dominated by rotifers at both stations, although during the winter periods the contribution of copepods was considerable. The rotifers were dominated byBrachionus angularis, B. calyciflorus, Keratella cochlearis andK. quadrata; the copepods by cyclopoid nauplii; the cladocerans by small-sized species mainly belonging toBosmina. At Maassluis the relative contribution of copepods was higher than at Lobith. Furthermore, the zooplankton at Maassluis included the speciesEurytemora affinis, characteristic for estuarine conditions. In spring, the rotifer density and water temperature and rotifer density and chlorophylla concentration were positively correlated. Furthermore, both rotifer density and chlorophylla were inversely correlated with discharge. The possible role of environmental factors (water temperature, chlorophyll content, discharge and biotic factors) controlling the river zooplankton dynamics is indicated.     

Responses of the photosynthetic flagellate,Euglena gracilis,to hypergravity

Motility and orientation has been studied in the unicellular photosynthetic flagellate, Euglena gracilis, using real time image analysis capable of tracking up to 200 cells simultaneously in the slow rotating centrifuge microscope (NIZEMI) which allows one to observe the cells' swimming behavior during centrifugation accelerations between 1 g and 5 g. At 1 g the cells show a weak negative gravitaxis, which increases significantly at higher accelerations up to about 3 g. Though most cells were capable of swimming even against an acceleration of 4.5 g, the degree of gravitaxis decreased and some of the cells were passively moved downward by the acceleration force; this is true for most cells at 5 g. The velocity of cells swimming against 1 g is about 10% lower than that of cells swimming in other directions. The velocity decreases even more drastically in cells swimming against higher acceleration forces than those at 1 g. The degree of gravitactic orientation drastically decreases after short exposure to artificial UV radiation which indicates that gravitaxis may be due to an active physiological perception rather than a physical effect such as an asymmetry of the center of gravity within the cell. Offprint requests to: D.-P. Häder     

Swimming behavior and ultrastructure of sperm of<Emphasis Type="Italic"> Lygodium japonicum</Emphasis> (Pteridophyta)

Swimming behavior of the sperm of Lygodium japonicum (Pteridophyta) and the associated ultrastructure of the flagellar apparatus were studied by video microscopy, transmission electron microscopy (TEM) and scanning electron microscopy (SEM). The sperm has approximately 70 flagella that emerge from a sinistrally-coiled flagellar apparatus, and swims forward by ciliary beat of these flagella. Backward swimming was not observed even after sperm collided with obstacles. Video microscopy showed that the flagella of the swimming sperm are oriented laterally and oblique-anteriorly. TEM and SEM observations revealed that the basal bodies of these flagella are arranged in at least two rows and oriented in the same directions as observed by video microscopy. These basal bodies (flagella) are categorized into two types according to their orientation: group I (laterally directed) and group II (oblique-anteriorly directed). The directionality of the basal bodies appears to be fixed by electron-dense material around their base. The outer dynein arms of the flagellar axoneme are entirely absent. These morphological characteristics of basal bodies (flagella) may relate to the lack of backward swimming behavior of the sperm. Based on these results, the evolution of swimming behavior in the archegoniates is discussed in connection with lack of backward swimming in a distantly related green alga, Mesostigma viride, and the Streptophyta.