Reaction times and force production during escape behavior of a calanoid copepod, Undinula vulgaris

Effective escape behavior contributes to the success of copepods in planktonic communities. The kinematics of escape were studied in tethered Undinulavulgaris (Calanoida) by analyzing the timing and magnitude of their power strokes to a precisely controlled, sudden mechanical perturbation in the surrounding water. Copepods responded with rapid swims to water velocities of 36 to 86 μm s⁻¹. Reaction times were under 2.5 ms following stimulus onset. The time course of force exerted was complex, but reproducible from stimulus to stimulus. Multiple power strokes (“kicks”) were frequently observed in response to single stimuli. Time intervals of 5 ms were observed between the end of one escape kick and the beginning of the next. U. vulgaris developed maximum forces of 40 to over 100 dynes (dyn) during a rapid swim. The behavioral reaction times and intervals between multiple responses observed in this calanoid are among the shortest reported in aquatic invertebrates. Received: 25 November 1997 / Accepted: 8 October 1998     

Spatial and temporal variability in egg production rates of the calanoid copepod Acrocalanus inermis

Egg production rate of the copepod Acrocalanus inermis was measured at 3 stations in south Kaneohe Bay, Hawaii in August 1977 and July–October 1979. The egg production rates were significantly variable spatially, but temporal variability over a time scale of weeks was larger than spatial variability. In three experiments, egg production rates were correlated with ambient concentrations of particulate matter, a crude estimate of food concentration. Most values for egg production rates ranged between 5 and 16.9 eggs female^-1 d^-1.     

Effects of suspended sediments on egg production of the calanoid copepod Acartia tonsa

The estuarine copepod Acartia tonsa was collected on several occasions between 4 April and 14 August 1985 from Terrebonne Bay, Louisiana (29°08N; 90°36W) and the effects in its diet of suspended sediments, collected from the same area, were measured at five different concentrations of sediment (100 to 1 000 ppm) and six phytoplankton concentrations (500 to 13 000 cells ml^-1 Thalassiosira weissflogii). Egg production rate was used as an index of diet quality. At low phytoplankton concentrations (500 cells ml^-1), and at intermediate phytoplankton concentrations (2 000 cells ml^-1) for previously starved copepods, egg production was reduced by up to 40% at a sediment concentration of 250 ppm and further reduced at higher sediment concentrations. At higher food concentrations (4 000 to 13 000 cells ml^-1), suspended sediment had no effect on egg production rates at sediment concentrations up to 500 ppm. Rates were reduced only at the highest sediment concentration of 1 000 ppm. Under most natural conditions, suspended sediment would not significantly affect egg production rates in A. tonsa.     

Annual biomass and production of the oceanic copepod community off Discovery Bay,Jamaica

Monthly samples were collected in oceanic waters off Discovery Bay, Jamaica, in 60- and 200-m vertical hauls, using 200- and 64-m mesh plankton nets, from June 1989 to July 1991. Length-weight regressions were derived for twelve genera of copepods (R²=0.79 to 0.97). For eight occasions spanning the study period, biomass estimates generated from these length-weight regressions differed by only 3% from direct weight determinations. The mean ash content of copepods was 7.1%, and the energy density was 20.8 kJ g^-1 ash-free dry weight (AFDW). Mean annual biomass of the total copepod community in the upper 60 m was 1.83 mg AFDW m^-3 (range 1.14 to 2.89 mg AFDW m^-3), and for the 200-m water column was 0.96 mg AFDW m^-3 (range 0.12 to 1.99 mg AFDW m^-3). Estimates of generation times for five common taxa ranged from 16.1 to 33.4 d. None of the taxa investigated displayed isochronal development; in general, stage duration increased in later copepodite stages. Weight increments showed a significant decrease in later copepodite stages, but with strong reversal of the trend from stage 5 to adult female in most species. Daily specific growth rates also declined in later copepodite stages, and ranged from 1.49 d^-1 in stage 1–2 Paracalanus/Clausocalanus spp. to 0.04 in stage 5-female of Oithona plumifera. Progressive food limitation of somatic copepodite growth and egg production is postulated. Naupliar production was 50.4 to 59.5% of copepodite production, and egg production was 35.1 to 27.7% of copepodite production in the 60-and 200-m water columns, respectively. Total annual copepod production, including copepodites, nauplii, eggs and exuviae, was 160 kJ m^-2 yr^-1 for the upper 60 m and 304 kJ m^-2 yr^-1 for the upper 200 m. Secondary production of the copepod community in oceanic waters off Discovery Bay approaches 50% of the corresponding value in tropical neritic waters.     

Density-dependent growth in the Atlantic surfclam, Spisula solidissima, off the coast of the Delmarva Peninsula, USA

Spisula solidissima (Dillwyn, 1817) is a large, suspension-feeding bivalve, whose range extends from Nova Scotia to South Carolina. This species is harvested commercially. Shell length and age data were collected for this species from 1980 to 1994 during surveys of population size and structure. These data were used to examine the relationship between the growth rate of S.␣solidissima and intraspecific density. The null hypothesis was that density (represented by number of individuals per tow) would have no effect on rate of growth. A negative relationship would support the alternative hypothesis, that intraspecific competition had taken place. This analysis focused on the surfclam population offshore from the Delmarva Peninsula, USA because: (1) a major recruitment event occurred in 1977, (2) clam fishermen had reported “stunted” surfclams in that area, (3) a wide range of local densities were available to examine, and (4) the existence of a closed area within the study area set up an interesting contrast with areas left open to harvesting. Maps of surfclam abundance across the Delmarva region demonstrate that areas of highest density have generally remained in the same location through time. The results suggested that intraspecific competition has been important in structuring this population. Based on data from 1980 to 1992, shell length was significantly reduced at high density, and a significant interaction between age and density was observed. Growth modeling indicated decreased asymptotic lengths and growth rates with increasing density. In nine out of ten pairwise randomization tests, fitted von Bertalanffy growth curves, representing different densities, were significantly different from each other. High densities of clams have persisted in the area that was closed to harvesting for 11 years (1980 to 1991). In 1994, length at age was significantly less in this closed area compared to that in the surrounding area. This effect was apparent in clams from 3 to 17 years of age, and most pronounced in the cohort that recruited to the Delmarva region in high numbers in 1977. Lower growth rates within the closed area have management implications for the optimal duration of closures. Received: 16 May 1997 / Accepted: 7 October 1997     

Occurrence,viability and significance of resting eggs of the calanoid copepod Labidocera aestiva

Field and laboratory observations on the occurrence and viability of eggs of Labidocera aestiva Wheeler show that eggs of this temperate species which are laid in late fall remain viable on the bottom and hatch in late spring. These eggs thereby provide the means of repopulating temperate areas with this species after its winter disappearance from the plankton.     

Spatial and temporal variations in the population size structure of three lanternfishes (Myctophidae) off Oregon,USA

Size-frequency distributions were determined for 3 common lantern-fishes (Stenobrachius leucopsarus, Diaphus theta, and Tarletonbeania crenularis) off Oregon in the summer. The fishes were caught mainly in sound-scattering layers by a large pelagic trawl with 5 opening-closing nets. Changes in depth distribution and diel vertical migration with growth were evident for all 3 species. The size of S. leucopsarus increased markedly with depth both at 0 to 90 m at night and 250 to 500 m during the day. Larger D. theta were also found deeper during the day (between 250 and 450 m), but neither D. theta nor T. crenularis demonstrated size segregation in the upper 90 m at night. Large D. theta and small T. crenularis did not appear to migrate into surface waters at night. Age-Group O (15 to 20 mm) S. leucopsarus were most abundant in deep water (400 to 480 m) in the daytime and did not migrate into near-surface waters at night. Age-Group I (30 to 40 mm) S. leucopsarus were common at about 300 m by day and within the upper 30 m at night. Age-Group II–III (50 to 60 mm) apparently followed the evening ascent of Age-Group I fish and most resided at 75 to 90 m at night, beneath Age-Group I fish. Age-Group III+fish (70 to 80 mm) were associated with Age-Group O at 400 to 480 m by day and usually did not migrate above 200 m at night. The size structure of S. leucopsarus differed among the nets of a single tow at one depth, or between two tows that fished the same depths on successive nights, indicating horizontal patchiness in age structure. D. theta demonstrated low within-tow variability in size composition which indicated a spatially more uniform age structure on a scale of kilometers. The size structures of these 3 lanternfishes were different in the same area and the same season during two different years, suggesting variable survival of year classes or horizontal patchiness of age composition in the area sampled.     

The effect of low concentrations of phenol and ammonia on egg production rates,fecal pellet production and egg viability of the calanoid copepod Acartia clausi

Estimates of daily fecundity, hatching success and fecal pellet production are reported for Acartia clausi females exposed for 10 d to low levels of phenol and ammonia. Copepods were collected in 1991 and 1992 from a southern coastal area of the Mar Grande of Taranto (southern Italy). A reduction in egg numbers and fecal pellet production was observed for females after 8 d of exposure to 500 g l^-1 phenol concentration. Ammonia (120 g l^-1) produced a significant increment in egg production, but hatching success was reduced by about 50% after nine exposure days. A. clausi was more sensitive to ammonia than phenol at high concentrations (24-h LC₅₀ phenol-32.26 mg l^-1; 24-h LC₅₀ ammonia=0.91 mg l^-1). At low concentrations, only long-term exposure to phenol determined a reduction in fecundity.     

Lunar influence on distribution of a calanoid copepod in the water column of a shallow,temperate estuary

Moonlight had a significant influence on the distribution of the copepodPseudodiaptomus hessei in the water column. Copepod samples were taken between March 1986 and March 1987 in the Sundays River estuary. South Africa. Late copepodid stages and adults showed vertical differences in abundance on moonlit nights, with more animals in bottom waters, but not on dark nights. Numbers of adults present in the water column were markedly lower on full moon nights compared with new moon nights. The strongest decrease in abundance was displayed by ovigerous females. These observed distribution patterns support published predation hypotheses.     

Discriminate feeding of the calanoid copepod Acartia clausi in mixtures of phytoplankton and inert particles

Adult female Acartia clausi were allowed to feed in the laboratory on each of three algae, the small green alga Dunaliella tertiolecta (5.6 m ESD), the diatoms Thalassiosira decipiens (13.4 m ESD) and T. nordenskioldii (17.7 m ESD), singly or in mixtures with polystyrene beads (15.7 m ESD). The ingestion rate on beads was much lower than that on cells, even when the supply of beads was one order of magnitude higher than that of cells. Beads offered singly were not ingested. In the experiments with T. nordenskioldii, the more cells adult females ingested, the higher the ingestion rate on beads. In addition, a linear regression equation describing the relationship between bead concentration and ratio of ingested beads to ingested cells was highly significant. These results suggest that beads are ingested by chance in feeding bouts on T. nordenskioldii. The bead interference on the ingestion rate on cells was observed in the experiments with D. tertiolecta and T. decipiens, but not with T. nordenskioldii. In a mixture of beads and T. decipiens, juvenile A. clausi discriminated beads from cells more successfully than adults. The presence of beads did not affect the ingestion rate of juveniles on cells. Starved adult females tended to ingest more beads than well fed individuals. The possible effects of body size and hunger on the discriminate feeding of copepods are discussed.     

Microsatellite variation, effective population size, and population genetic structure of vermilion snapper, Rhomboplites aurorubens, off the southeastern USA

Vermilion snapper (Rhomboplites aurorubens) were collected from four sites off the Atlantic coast of the USA and one site in the Gulf of Mexico to evaluate effective population size and genetic stock structure. Previous studies had suggested geographic variation in the ratio of males to females, so this population characteristic was explored in conjunction with the genetic analysis. Sex ratio varied greatly among the five sample sites; males comprised 57% of samples in the Gulf of Mexico, while within the South Atlantic Bight they comprised between 36% (Morehead City, North Carolina) and 53% (Carolina Beach, North Carolina) of samples. No clear geographic trends in the sex ratio emerged; instead, it was found to vary with fish length, the percentage of males decreasing with increasing size. Allelic variation assessed at seven dinucleotide microsatellite loci was large; gene diversities ranged from 0.43 to 0.95 and allelic counts from 7 to 39. Estimates of the effective population size ranged from 24 500 (based on the infinite-alleles model) to 150 500 (based on the stepwise-mutation model). There was evidence for excess homozygosity within samples: estimates of F _IS (the correlation of alleles within individuals) ranged from 0.01 to 0.03 among the seven loci, and three estimates were significantly greater than zero. Differentiation among localities was very weak, as estimates of F _ST (the correlation of alleles within populations) were on the order of 0.001 to 0.002 and genetic distance estimates between localities were not related to geographic distances. This suggested that vermilion snapper in the South Atlantic Bight (Cape Hatteras, North Carolina to Cape Canaveral, Florida) and Gulf of Mexico are likely to consist of one genetic stock. Despite the overall homogeneity, there were indications of a temporally dynamic local structure that would bear further examination. Received: 6 July 1998 / Accepted: 9 February 1999     

Bacterial chitinase in the stomachs of marine fishes from Yaquina Bay,Oregon, USA

The occurrence of chitinase in the stomach contents of Enophrys bison and Platichthys stellatus was investigated. The highest chitinase activity and the greatest percentage of chitinoclastic bacteria in relation to the total bacterial flora were associated with fish whose stomach content was composed primarily of chitinous animals. Stomach contents lacking visible chitin possessed low or no detectable levels of chitinase activity and few chitinoclastic bacteria. Juvenile E. bison treated with chloramphenicol to remove their indigenous bacterial flora had no detectable level of chitinase in their stomach contents while non-treated juveniles showed inducible chitinase activity, indicating the bacterial origin for the chitinase.Published as Technical Paper No. 4435, Oregon Agricultural Experiment Station. This research was supported by NIH training grant 5TO1 GM-00704 and NSF grant DES73-06611 AO2.     

Growth and production of the red alga Gelidium sesquipedale off the Basque coast (northern Spain)

Several aspects of the growth of the agarophyte Gelidium sesquipedale Clemente (Thuret) were studied in sublittoral populations of the Basque coast (northern Spain) from 1981 to 1986. The standing stock of the populations varied greatly with season. During two consecutive years, 60 to 70% of summer biomass had been lost by the end of the unfavourable period of year to growth (autumn and winter). The decline in biomass was mainly due to partial losses of branches, with more branches being lost from larger fronds. Yearly variations in the standing stock were also recorded. Production estimates obtained by a destructive sampling method were not satisfactory because of the spatial heterogeneity of the populations at the three locations studied. Indirect estimation of production by measuring the growth of marked fronds in a control quadrat gave more reliable estimates. Production values of 622±124, 735±147 and 615±123 g dry wt m^-2 yr^-1 were obtained at the three locations, respectively. Frond elongation rates of 7.5, 6.1, 7.5 and 7.9 cm yr^-1, respectively, were recorded during four consecutive years. In winter elongation rates of fronds were rather low. Elongation did not seem to be affected by frond length, the self-shading effect of the canopy, or changes in depth. Cuts resulting from grazing were frequent on fronds through the year, decreasing significantly with increasing depth. Sheared fronds regenerated promptly, independent of season or height at which the cut was made. For commercial purposes the best season for harvesting is summer, which is the period of maximum standing stock, infertility, and calm warm waters that provide easier working conditions for divers.     

Phytoplankton sinking rates in oligotrophic waters off Hawaii,USA

The sinking rates in two size fractions of natural phytoplankton were measured over much of the photic zone in a subtropical environment. At 24, 40 and 71 m, sinking rates (± SD) of the 3 to 20 m fractions were 0.72±0.05, 0.83±0.05, and 0.34±0.04 m · d^-1, respectively, and rates for the 20 to 102 m fraction were 1.50±0.21, 1.65±0.14, and 0.95±0.22 m · d^-1, respectively. At all depths sampled, the 20 to 102 m size fraction was observed to sink significantly (P<0.01) faster than the 3 to 20 m fraction. considering vertical variability, both size fractions were observed to have significantly (P<0.01) lower sinking rates at 71 m than at more shallow depths. The finding of lower sinking rates in the region which lies just above the subsurface chlorophyll maximum provides empirical support for the hypothesis that variations in phytoplankton buoyancy may be related to the maintenance of this feature which is typical in oceanic environments. Analysis of sinking rate traces describes the distributions of specific sinking velocities for each size fraction and their variation.Oceanic Institute Contribution No. 162     

Persistent blooms of surf diatoms along the Pacific coast,USA

Productivity was studied in two diatom species, Chaetoceros armatum T. West and Asterionella socialis Lewin and Norris, which form persistent dense blooms in the surf zone along the Pacific coast of Washington and Oregon, USA. Past observations have shown that surf-diatom standing stock usually declines in summer along with concentrations of nitrate and ammonium. Using the ¹⁴C method, photosynthetic rates in natural surf samples were measured monthly for one year (October 1981 through September 1982) at a study site on the Washington coast. Also measured were temperature, salinity, dissolved nutrients, particulate carbon and nitrogen (used as estimates of phytoplankton C and N), and chlorophyll a. Assimilation numbers (P _max) were higher in summer (5 to 8 g C g^-1 chl a h^-1) than in winter (3 to 4gC). Specific carbon incorporation rates (µ_max) showed no obvious seasonality, mostly falling within the range of 0.09 to 0.13 g C g^-1 C(POC) h^-1. The discrepancy between the seasonal trends for chlorophyll-specific and carbon-specific rates reflects a change in the carbon-to-chlorophyll ratio. Because of seasonal differences in daylength and light intensity, daily specific growth rates () are thought to be higher in summer than in winter. Neither ammonium enrichment assays nor particulate carbon-to-nitrogen ratios provided convincing evidence for nitrogen limitation during summer, and the observed changes in diatom abundance cannot be explained on this basis. Both the high diatom concentrations and their seasonal variations probably are due mainly to factors other than growth rates; two factors considered important are diatom flotation and seasonal changes in wind-driven water transport. C. armatum usually dominates the phytoplankton biomass in the surf zone, and evidence suggests that this species is strongly dominant in terms of primary production.Contribution No. 1391 of the School of Oceanography, University of Washington, Seattle, Washington, USA     

Comparison of the chemical composition of particulate material and copepod faecal pellets at stations off the coast of Labrador and in the Gulf of St. Lawrence

Faecal pellets were collected in 1988 from copepods which had fed in situ or in laboratory experiments, using screened natural seawater as food, at two stations off the coast of Labrador and one in the Gulf of St. Lawrence. The chemical composition of the pellets and of particulate material in profiles and in laboratory food were measured in terms of particulate carbon, carbohydrate (soluble and insoluble), protein and lipid. Faecal pellet composition was somewhat similar in all experiments at the first two stations, where the compositions of particulate material in situ and copepod species assemblages were also similar. At the third station the compositions of faecal pellets and particulate material were slightly different from those at the other stations and the copepod species composition varied between sampling times. Faecal pellets at the first two stations had very low levels of soluble carbohydrate, while concentrations in the food were generally high, suggesting that it was efficiently metabolized by copepods, although it might have been absent because of sloppy feeding or release, after passage through the gut, in soluble form or from faecal pellets. Comparisons of POC: biogenic silica ratios in food and faecal pellets, calculated using data presented elsewhere (Head 1992; Mar. Biol. 112: 583–592), suggested that at these stations, where food concentrations were high (chlorophyll concentrations>8 gl^-1), copepods may have been assimilating carbon rather inefficiently.     

Calanus marshallae,a new species of calanoid copepod closely allied to the sibling species C. finmarchicus and C. glacialis

Species of the copepod genus Calanus frequently dominate the marine zooplankton in boreal and arctic waters. Up to now there have been no operational means of identifying several species closely related to Calanus finmarchicus. Reanalysis of these taxa, using material from plankton samples collected throughout Northern Hemisphere polar and coreal waters, shows that there are 3 sibling species which have been previously combined under the names C. finmarchicus and C. glacialis. Several new taxonomic characters permit unequivocal identification of C. finmarchicus, C. glacialis, and a new species, C. marshallae. Claims that C. finmarchicus and C. glacialis are subspecies are refuted; there is no evidence that the two species continuously intergrade either where they co-occur or where they are allopatric, nor is there evidence that the two species hybridize. C. finmarchicus is basically restricted to the North Atlantic Ocean and C. marshallae to the North Pacific Ocean and Bering Sea; C. glacialis is primarily an Arctic species, but its geographical distribution slightly overlaps those of the other two species. Taxa closely related to C. finmarchicus and C. helgolandicus probably represent two separate, but closely linked evolutionary lineage; species of these two lineages are placed in one of two species groups, the finmarchicus group and the helgolandicus group.     

Annual structure of the copepod community and its associated pelagic environment off Discovery Bay,Jamaica

Monthly samples were collected in oceanic waters off Discovery Bay, Jamaica, in 60- and 200-m vertical hauls, using 200- and 64-m mesh plankton nets, from June 1989 to July 1991. Sixty-nine species of copepods were identified: nauplii, copepodites and adults were separately enumerated. Total copepod abundances (all stages) ranged from 695 to 4120 m^-3 in the upper 60 m, and from 483 to 3319 m^-3 in the 200-m water column, without any clear seasonal pattern. With the exception of temperature, no seasonal variations in physico-chemical (chlorophyll a, S, particulate organic carbon, particulate protein) or biological variables were evident. Nauplii, adults and copepodites of selected taxa, and two chaetognath species, showed no significant variations in body length. Significant variantions in reproductive index were detected for several species, but without seasonal trends; many species appear to be continuous or intermittent breeders. There was no evidence of seasonal pattern in overall community composition or diversity, or evidence of changes due to water mass advection. The copepod community can be divided into a recurrent group of 13 (at 60 m) to 17 (at 200 m) perennial species, present year-round, and associated ephemeral groups of 1 to 3 species, present randomly for 1 to 4 consecutive months. The most plausible explanation of these patterns is that broad areas of the Caribbean Sea are dominated by the community of perennial species, while the ephemeral species represent the superimposed influence of local mesoscale gyres.     

Some particulate and soluble agents affecting the relationship between metal toxicity and organism survival in the calanoid copepod Euchaeta japonica

Particulate and water-soluble agents were tested to determine their ability to affect the relationship between metal toxicity and the survival of Euchaeta japonica (Copepoda, Calanoida). Clay minerals and diatoms were two types of particles capable of affecting this relationship. Ascorbic acid, sewage effluent, and water extracts of humic acid and two types of soils exhibited the same capability. The ability of the water-soluble agents was compared with that of a known chelating agent in an attempt to quantify the activity of the agents.