Metabolism and elemental composition of four oncaeid copepods in the western subarctic Pacific

Respiration rates and elemental composition (carbon and nitrogen) were determined for four dominant oncaeid copepods (Triconia borealis, Triconia canadensis, Oncaea grossa and Oncaea parila) from 0–1,000 m depth in the western subarctic Pacific. Across the four species of which dry weight (DW) varied from 2.0 to 32 μg, respiration rates measured at in situ temperature (3°C) increased with DW, ranging from 0.84 to 7.4 nl O₂ individual⁻¹ h⁻¹. Carbon (C) and nitrogen (N) composition of the four oncaeid species ranged from 49–57% of DW and 7.0–10.3% of DW, respectively, and the resultant C:N ratios were 4.8–8.3. The high C contents and C:N ratios were reflected by large accumulation of lipids in their body. Specific respiration rates (SR, a fraction of body C respired per day) ranged between 0.5 and 1.3% day⁻¹. Respiration rates adjusted to a body size of 1 mg body N (i.e. adjusted metabolic rates, AMR) of the four oncaeid species [0.6–1.1 μl O₂ (mg body N)^−0.8 h⁻¹ at 3°C] were significantly lower than those (1.7–5.1) reported in the literature for oithonid and calanoid copepods at the same temperature. The present results indicate that lower metabolic expenditure due to less active swimming (pseudopelagic life mode) together with rich energy reserve in the body (as lipids) are the characters of oncaeid copepods inhabiting in the epi- and mesopelagic zones of this region.     

In situ grazing rate of the copepod population in the western subarctic North Pacific during spring

Time series sampling with a multi-layer plankton sampler was carried out in the western subarctic North Pacific during spring 1991. Neocalanus cristatus, N. flemingeri and Eucalanus bungii dominated and accounted for 88.5% of the copepod population in volume. Neocalanus spp. were distributed in the upper mixed layer, while E. bungii was mainly distributed between 120 and 300 m throughout the day and night. In contrast, Metridia pacifica, Pleuromamma scutullata and Gaetanus simplex showed clear diel vertical migration. Grazing activities were estimated simultaneously by gut fluorescence. Nocturnal grazing was observed for diel migrating species. Neocalanus spp. did not have a diel feeding rhythm and had relatively low gut fluorescence. E. bungii was considered to be dormant during the observation period. The estimated grazing rate of the copepod population on phytoplankton was 1.4 to 2.0% of the primary production while the metabolic requirement was 8.3 to 12.4% of the primary production. These facts suggest that the copepod population was unimportant as primary consumers and that microzooplankton plays a much more important role in sustaining low standing stock of phytoplankton and a high nutrient concentration in the western subarctic Pacific Ocean.     

Vertical distribution,abundance and community structure of oncaeid copepods in the Oyashio region,western subarctic Pacific

The vertical distribution, abundance and community structure of oncaeid copepods were investigated in the Oyashio region, western subarctic Pacific. Samples were collected with a 0.10 mm mesh closing type net from five discrete layers down to a maximum depth of 2,000 m in September and December 1996 and in April and October 1997. The copepods were widely distributed from epipelagic to bathypelagic zones, and showed prominent peaks of abundance above the thermocline and/or between 250 and 1,000 m depth. Standing stocks of total oncaeid copepods ranged from 1.5 to 2.5×10⁵ inds m^–2 at 0–2,000 m in the water column, which are the same order of magnitudes as those reported in tropical, subtropical and polar regions by previous workers. A total of 38 species and two forms belonging to the genera Oncaea, Triconia, Spinoncaea, Conaea and Epicalymma, and two provisionally classified species of the family Oncaeidae were identified in this study. Of these, 14 species have already been recorded from the eastern subarctic Pacific. Several warm-water species were also found in December 1996 and/or October 1997, when the effect of warm-core rings originating from the Kuroshio Current was evident in the epipelagic zone. Dominant species throughout the whole water column were T. borealis, T. canadensis, O. grossa, O. parila, O. rimula, O. lacinia, Epicalymma spp. and Oncaea sp. A, and these eight species together always made up about half of the total oncaeid numbers. Community analysis revealed that species composition in epipelagic and upper mesopelagic zones varied temporally due to the changes of hydrographic conditions, whereas those in lower mesopelagic and bathypelagic zones were relatively stable, reflecting the constancy of deep environments.Communicated by O. Kinne, Oldendorf/Luhe     

Microbial food partitioning by three species of benthic copepods

Using radioactively labeled bacteria and photoautotrophs in undisturbed sediment cores, we show that three cooccurring species of benthic copepods feed on different microbial food sources in their natural environment. Specifically, Thompsonula hyaenae feeds on photoautotrophs, Halicyclops coulli feeds on bacteria, and Zausodes arenicolus feeds on both photoautotrophs and bacteria. Species of benthic copepods feed differently from one another in the field, and meiofaunal species' distributions could be influenced by distributions of their preferred microbial food.     

Vertical distribution,population structure and life cycles of four oncaeid copepods in the Oyashio region,western subarctic Pacific

Vertical distribution and population structure of four dominant oncaeid copepods (Triconia borealis, Triconia canadensis, Oncaea grossa and Oncaea parila) were investigated in the Oyashio region, western subarctic Pacific. Seasonal samples were collected with 0.06 mm mesh nets from five discrete layers between the surface and 2,000 m depth at seven occasions (March, May, June, August and October 2002, December 2003 and February 2004). The depth of occurrence of major populations of each species differed by species; the surface–250 m for T. borealis, 250–1,000 m for T. canadensis, 250–500 m for O. grossa and 500–1,000 m for O. parila. The ontogenetic vertical migration characterized by deeper occurrence of early and late copepodid stages, and shallower occurrence of middle copepodid stages was observed in T. canadensis and O. parila. Of the four oncaeid copepods, almost all copepodid stages occurred throughout the study period, suggesting that their reproduction continues throughout the year in the region. Nevertheless, a clear developmental sequence of stage-to-stage was traced for T. canadensis and O. grossa copepodids, implying their generation time to be 1 year. For T. borealis and O. parila copepodids, no clear seasonal succession was observed thus estimation of their generation time was uncertain. The present comprehensive results of vertical distribution and life cycle features for T. borealis, T. canadensis, O. grossa and O. parila are compared with the few published data on oncaeid species distributing in high latitude seas.     

Sexual reproduction of three coral species from the Mexican South Pacific

Our understanding of the reproductive biology of corals from the Mexican southeastern Pacific is limited, and consequently, the role of reproduction in structuring coral communities is unclear. As a first attempt to understand the importance of sexual reproduction in structuring and maintaining of the coral communities from this region, we documented the reproductive cycles over 2 years (2003–2004) in three main reef-building corals species in the region. Pocillopora damicornis was shown to be hermaphroditic with asynchronous gamete development observed only in 2004; P. gigantea was characterized as both gonochoric and cosexually hermaphroditic. Absence of mature gametes was documented in both species, and an observation may be attributed to the 2003 El Niño Southern Oscillation event, which may have inhibited reproductive maturation via thermal stress. Porites panamensis was gonochoric with asynchronous development, and planulae were generally brooded. The presence of mature gametes and planulae in P. panamensis polyps suggests that this species is an important contributor to local and, likely, regional recruitment of this species. Further research should seek to identify important source populations for these coral recruits and document the exchange of larvae between coral populations of the Mexican Pacific.     

Comparative life histories of three species of planktonic calanoid copepods in Dabob Bay,Washington

The life history patterns of Calanus marshallae Frost, C. pacificus Brodsky, and Metridia lucens Boeck were determined in Dabob Bay, Washington, USA, during 1973 and October 1981–September 1982. C. marshallae emerged from diapause and moulted to adults in January–February. One major generation was observed, having been produced mainly during early to mid-March. Most surviving individuals spawned in March, arrested development at the C5 stage, and were in diapause by late May. However, some individuals developed to adults by late April and produced a very minor second generation. C. pacificus emerged from diapause and moulted to adults in February–March. The first generation produced during the spring lagged slightly behind the C. marshallae generation. C. pacificus produced two additional generations, one in late spring and one in the fall. The production of the three generations coincided with or closely followed phytoplankton blooms. Some C. pacificus arrested development at the C5 stage and entered diapause during the summer, but the majority of the population did not do so until fall. M. lucens did not appear to enter a diapause state. The fall and winter population was chiefly adult females which mostly remained at depth and were reproductively immature. Nevertheless, naupliar M. lucens were found on all sampling dates, indicating that reproduction never ceased as it did for the two Calanus species. M. lucens produced generations in late winter/early spring, late spring, and late summer/early fall. Interspecific comparisons between the life histories observed in Dabob Bay and intraspecific comparisons between the life histories in Dabob Bay and other locations are made. The advantages or disadvantages conferred to the copepods by their differing life histories are also discussed.     

Zooplankton responses to hypoxia: behavioral patterns and survival of three species of calanoid copepods

Seasonally recurrent and persistent hypoxic events in semi-enclosed coastal waters are characterized by bottom-water dissolved oxygen (d.o.) concentrations of < 2.0 ml l⁻¹. Shifts in the distribution patterns of zooplankters in association with these events have been documented, but the mechanisms responsible for these shifts have not been investigated. This study assessed interspecific differences in responses to hypoxia by several species of calanoid copepods common off Turkey Point, Florida, USA: Labidocera aestiva (Wheeler) (a summer/fall species), Acartia tonsa (Dana) (a ubiquitous year-round species), and Centropages hamatus (Lilljeborg) (a winter/spring species). Under conditions of moderate to severe hypoxia 24-h survival experiments were conducted for adults and nauplii of these species from August 1994 to October 1995. Experiments on adults used a flow-through system to maintain constant d.o. concentrations. Adults of A. tonsa showed no decline in survival with d.o. as low as 1.0 ml l⁻¹, sharp declines in survival at d.o. = 0.9 to 0.6 ml l⁻¹, and 100% mortality with d.o. = 0.5 ml l⁻¹. Adults of L. aestiva and C. hamatus were more sensitive to oxygen depletion: both species experienced significant decreases in survival for d.o. = 1.0 ml l⁻¹. Nauplii of L. aestiva and A. tonsa showed no significant mortality with d.o. = 1.1 to 1.5 ml␣l⁻¹ and d.o. = 0.24 to 0.5 ml l⁻¹, respectively. In addition, experiments investigating behavioral avoidance of moderate to severe hypoxia were carried out for adults of all three species. None of the three species effectively avoided either severely hypoxic (d.o. < 0.5 ml l⁻¹) or moderately hypoxic (d.o. ≈ 1.0 ml l⁻¹) bottom layers in stratified columns. These results suggest that in␣nearshore areas where development of zones of d.o. < 1.0 ml l⁻¹ may be sudden, widespread, or unpredictable, patterns of reduced copepod abundance in bottom waters may be due primarily to mortality rather than avoidance. Received: 31 August 1996 / Accepted: 24 September 1996     

Small-scale horizontal distributions of zooplankton taxa

Small-scale (100 to 2 400 m) horizontal distributions of major taxonomic categories (class and order) of zooplankton were measured at a depth of 90 m with an opening-closing plankton net over a 3 d period in October 1978 in the California Current. Some zooplankton categories showed evidence of diurnal vertical migration, while others had long-period temporal changes in mean abundance. Variance-to-mean ratio for large copepods and euphausiids was higher at night than during the day, while the opposite was true for chaetognaths and pteropods. Within a given category, the variance-to-mean ratio generally increased with a category's abundance. Spatial abundance variations were characterized by trends (i.e., fluctuations larger than length of the net hauls) in some taxonomic categories. No consistent differences in scales of variability were found as a function of animal size or from day to night. Correlation analysis of taxonomic counts implied that significant biological interactions occurred. The proportions of counts of taxonomic groups showed no large changes over the time-space scales sampled. However, the proportions of biomass in taxonomic groups differed from day to night due to the large variability of euphausiids. Comparisons of wet weight biomass to taxonomic counts indicated that biomass was usually less variable than taxonomic counts.     

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.     

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).     

Variability in dry weight and vertical distributions of decapod larvae in the Irish Sea and North Sea during the spring

Accumulation of waterborne cadmium in Littorina littorea, Mytilus edulis and Carcinus maenas (collected in 1988 and 1989 around the island of Funen, Denmark) was investigated in a matrix of salinities (10 to 30) and calcium concentrations (2.9 to 8.9 mM Ca⁺⁺). Cadmium accumulation rates in soft parts of L. littorina, soft parts and shells of M. edulis and whole bodies and exoskeletons of C. maenas decreased with increasing salinity. Changes in the calcium concentrations accounted for 72% of the salinity effect on cadmium accumulation rates in L. littorina, whereas calcium concentrations had little or no effect on cadmium accumulation in M. edulis. Cadmium accumulation in the whole body of C. maenas was affected equally by calcium concentrations and total salinity, whereas accumulation in the exoskeleton was mainly affected by changes in total salinity. Individual variability in cadmium accumulation in the organs of C. maenas was greater than the variation attributable either to changes in ambient calcium concentrations or total salinity. An appreciable amount of the inter-individual variability in the cadmium accumulation in all three species was correlated with wet:dry weight ratios of the tissues and size of the organisms.     

Feeding four species of pelagic copepods under experimental conditions

Different qualitative and quantitative aspects of feeding with varied compositions of diets have been studied in 4 species of pelagic copepods: Calanus helgolandicus, Centropages typicus, Temora stylifera and Acartia clausi. By feeding copepods different algal concentrations, it was shown that when food concentration increases grazing rate decreases; the ingestion rate remains fairly constant in the lower range of concentrations, but then increases, reaching a plateau at higher algal concentrations. There is a significant correlation between daily food intake and fecal pellet production. On a pluri-algal diet, selective grazing is observed: larger phytoplankton cells are more efficiently removed than smaller ones. Using Artemia nauplii, it is shown that the copepods studied are also able to eatch and ingest animal prey. Increased daily food intake affects respiration and oviposition. Metabolic requirements, gross growth-efficiency, and food assimilation have been ealculated in Calanus helgolandicus, Centropages typicus and T. stylifera for a large range of algal concentrations.     

Water-soluble vitamins in natural plankton (copepods) during two consecutive spring blooms compared to vitamins in Artemia franciscana nauplii and metanauplii

Weight-specific contents of asorbic acid, thiamin, riboflavin, niacin, pantothenic acid, vitamin B₆, biotin, folate and vitamin B₁₂ were analysed in natural zooplankton collected from a sea water lagoon in a halibut fry production plant. The samples of natural zooplankton were collected from April/May to the end of June during two consecutive spring blooms. For comparison, the same vitamins were measured in nauplii of Artemia franciscana, just after hatching and after enrichment for 24 and 48 h with a commercial diet. The changes in vitamin content in zooplankton, which consisted mainly of Temora longicornis, during the two spring blooms were <2.5-fold for each specific vitamin. The large number of high quality halibut fry produced using the plankton as a dietary source suggests that the plankton contained sufficient amounts of all water-soluble vitamins. With the exception of thiamin and ascorbic acid, Artemia– both newly hatched and enriched for 24 and 48 h – contained higher levels of water-soluble vitamins than natural zooplankton. Enrichment feeding of Artemia for 48 h stabilised or increased the level of the water-soluble vitamins. Received: 8 March 1999 / Accepted: 21 February 2000     

Production of tropical copepods in Kingston Harbour, Jamaica: the importance of small species

The copepod community observed during an 18-month period at the mouth of eutrophic Kingston Harbour, Jamaica, was dominated by small species of Parvocalanus, Temora, Oithona, and Corycaeus. Mean copepod biomass was 22.1 mg AFDW m⁻³ (331 mg m⁻²). Annual production was 1679 kJ m⁻², partitioned as 174 kJ m⁻² naupliar, 936 kJ m⁻² copepodite, 475 kJ m⁻² egg and 93 kJ m⁻² exuvial production. All nauplii, most copepodites and many adults, equivalent to half of the biomass and production, were missed by a standard 200-μm plankton net, emphasizing the importance of nauplii and small species in secondary production estimates. The evidence suggests that growth rates and production are generally not food limited, and we speculate that size-selective predation shapes the structure of the harbour community. Biomass and production are higher than previous estimates for tropical coastal waters, but comparable to other eutrophic tropical embayments and many productive temperate ecosystems. Far from being regions of low productivity, tropical zooplankton communities may have significant production and deserve greater research attention than they currently receive. Received: 19 September 1997 / Accepted: 21 October 1997     

Egg production and feeding by copepods prior to the spring bloom of phytoplankton in Fram Strait,Greenland Sea

Reproductively activeCalanus hyperboreus (Krøyer) andC. glacialis Jaschnov were captured in the upper 100 m of Fram Strait (77° to 79°N) in late winter 1987. There was no evidence of a phytoplankton bloom; chlorophylla concentrations were uniformly low (<0.1 mg m^–3), and nitrate concentrations were uniformly high (>11.3 mg-at m^–3). Gut-fullness measurements indicated that females were ingesting very little. The maturation state of gonads of bothC. hyperboreus andC. glacialis indicated that 75% of females were in a ripe condition consistent with observed egg laying. The lipid content of females laying eggs was reduced in both species compared to that of females not laying eggs. InC. hyperboreus the reduction was 39% and inC. glacialis it was 44%. All the evidence suggests that bothC. hyperboreus andC. glacialis were laying eggs in late winter by using lipids stored previously; they were not relying on ambient concentrations of phytoplankton. The daily rate of egg laying byC. glacialis using lipids in late winter exceeded the rate reported for summer when ambient food supplies have been shown to be necessary. It is suggested that individuals, spawned well in advance of the spring bloom of phytoplankton, may comprise a major portion of the annual recruitment to the entire population ofC. glacialis in this area, and that their life cycle can be completed within 1 yr. NeitherMetridia longa (Lubbock) norC. finmarchicus (Gunnerus) laid eggs during this study.     

Spatial and temporal distributions of copepods to leeward and windward of Oahu, Hawaiian Archipelago

The spatial and temporal distributions of two island-associated copepod species, Undinula vulgaris Dana and Labidocera madurae Scott, were compared to the distributions of two open ocean species, Cosmocalanus darwinii Lubbock and Scolecithrix danae Lubbock, along 28-km windward and leeward transects off the island of Oahu, Hawaii. Samples were taken in September and December 1985 and April and June 1986. A warm, low salinity pool on the leeward side was a prominent feature during all transects except December. The abundances of the two oceanic species did not change significantly between leeward and windward stations, with distance from shore, or between September 1985 and April 1986 samples. As expected, very high abundances of U. vulgaris occurred at some nearshore stations, up to 3 g dry wt m⁻² for adults alone. Calculations of respiratory loss at these densities (0.7 g C m⁻² d⁻¹) suggest a high local productivity would be required to meet these demands. L. madurae, a surface-dwelling species normally restricted to within 1 km of shore, was an effective indicator species of nearshore water movement. It was more common in offshore samples on the leeward transects, rarely being found offshore on the windward side, consistent with prevailing currents and the presence of the leeward warm, low salinity pool. The occurrence of a strong mixing event in April 1986 resulted in L. madurae being distributed throughout the upper 100 m of the water column. The presence of oceanic species close to shore on the windward side also coincided with this wind-driven event. The primary environmental influence on vertical distributions was daytime cloud cover, with U. vulgaris tending to be found shallower on cloudy days. Of the two oceanic species, S. danae exhibited the most pronounced vertical migration, however, vertical distributions were not significantly correlated with environmental factors for either species. The abundant nearshore U. vulgaris population cannot be explained by differences in vertical distribution between it and the two oceanic species that might allow a physical mechanism to concentrate U. vulgaris. A high population growth rate is likely necessary to explain U. vulgaris' dominance. Received: 26 June 1998 / Accepted: 31 March 1999