Impact of a burrowing deposit-feeder, Monoporeia affinis, on viable zooplankton resting eggs in the northern Baltic Sea

We investigated the effect of different densities of the burrowing deposit-feeding amphipod Monoporeia affinis on the potential for recruitment of zooplankton from benthic resting eggs. Hatching of resting eggs was induced in the laboratory on sliced and resuspended 1-cm depth-sections of sediment cores, collected at six stations in an archipelago area of the Gulf of Bothnia, Baltic Sea. The uppermost 5 cm of the sediment was studied. The most common species that hatched was Eurytemora affinis (Copepoda). Individuals from another copepod genus, Acartia, hatched in significant numbers only in the cores from two stations with low amphipod abundance. Cores from stations with high amphipod densities showed a deeper distribution of emerging E. affinis nauplii compared with stations with few amphipods; the oxidised sediment layer was also deeper at high M. affinis densities than at low. Total (0 to 5 cm strata pooled) number of hatched E. affinis nauplii was independent of amphipod density. This indicates that the effect of M. affinis on E. affinis eggs involves deeper burial due to bioturbation, rather than predation. Decreased benthic recruitment of zooplankton at localities with high M. affinis density is suggested, since more deeply positioned eggs are less likely to hatch. When hatching was induced in intact, non-sliced cores from one station, the number of E. affinis nauplii that hatched was on average 43% of the number that hatched in the upper centimetre of the sliced cores from the same station. This fraction (43%), if applied to the other stations, implied a potential for benthic recruitment of up to 80 000 ind m⁻² for E. affinis. Due to its high abundance, M. affinis is likely to greatly reduce benthic recruitment of zooplankton in this system. Received: 17 September 1999 / Accepted: 23 February 2000     

Nodularin concentrations in Baltic Sea zooplankton and fish during a cyanobacterial bloom

Toxic cyanobacterial blooms, dominated by Nodularia spumigena, are a recurrent phenomenon in the Baltic Sea during late summer. Nodularin, a potent hepatotoxin, has been previously observed to accumulate on different trophic levels, in zooplankton, mysid shrimps, fish as well as benthic organisms, even in waterfowl. While the largest concentrations of nodularin have been measured from the benthic organisms and the food web originating from them, the concentrations in the pelagic organisms are not negligible. The observations on concentrations in zooplankton and planktivorous fish are sporadic, however. A field study in the Gulf of Finland, northern Baltic Sea, was conducted during cyanobacterial bloom season where zooplankton (copepod Eurytemora affinis, cladoceran Pleopsis polyphemoides) and fish (herring, sprat, three-spined stickleback) samples for toxin analyses were collected from the same sampling areas, concurrently with phytoplankton community samples. N. spumigena was most abundant in the eastern Gulf of Finland. In this same sampling area, cladoceran P. polyphemoides contained more nodularin than in the other areas, suggesting that this species has a low capacity to avoid cyanobacterial exposure when the abundance of cyanobacterial filaments is high. In copepod E. affinis nodularin concentrations were high in all of the sampling areas, irrespective of the N. spumigena cell numbers. Furthermore, nodularin concentrations in herring samples were highest in the eastern Gulf of Finland. Three-spined stickleback contained the highest concentrations of nodularin of all the three fish species included in this study, probably because it prefers upper water layers where also the risk of nodularin accumulation in zooplankton is the highest. No linear relationship was found between N. spumigena abundance and nodularin concentration in zooplankton and fish, but in the eastern area where the most dense surface-floating bloom was observed, the nodularin concentrations in zooplankton were high. The maximum concentrations in zooplankton and fish samples in this study were higher than measured before, suggesting that the temporal variation of nodularin concentrations in pelagic communities can be large, and vary from negligible to potentially harmful.     

Community-level and species-specific responses of coastal phytoplankton to the presence of two mesozooplanktonic grazers

The response of phytoplankton to the presence of two mesozooplanktonic grazers was studied in a 44-h laboratory experiment from the phytoplankton perspective. The <45-μm-filtered plankton community, as well as females of the copepod Eurytemora affinis and nauplii of the barnacle Balanus improvisus originated from the northern Baltic Sea. The phytoplankton community was dominated by the dinoflagellate Heterocapsa triquetra, which was preferred as food by the mesozooplanktonic grazers, especially E. affinis. Cryptophytes were eaten by the B. improvisus nauplii, while heterotrophic nanoflagellates increased in the presence of B. improvisus nauplii, and colonial cyanobacteria increased in the presence of E. affinis. Because of the strong selective feeding on the dominant, large-sized phytoplankton species, the negative effect of the mesozooplanktonic grazers on total chlorophyll a was stronger than any cascade effect releasing phytoplankton from protozoan grazing.     

Seasonal growth of the benthic amphipodsPontoporeia affinis andP. femorata in a Baltic archipelago in relation to environmental factors

An intensive field study of three depth zones in 1980 revealed different seasonal patterns of growth in the deposit-feedingPontoporeia affinis Lindström andP. femorata Kröyer (Crustacea, Amphipoda) in the northern Baltic Sea, SW coast of Finland. The study areas were situated in the outer archipelago at depths of 21 to 25 m and 26 to 30 m and in the sea zone at 46 m. InP. affinis growth rates decreased with increasing depth, but the same seasonal pattern was evident at all depths: growth began in spring, increased to a maximum in early or midsummer, and decreased again or ceased altogether by autumn. In the archipelago the 1 + age group attained its maximum growth rate in May–June and the 0 + age group clearly later, in June–July. InP. femorata the growth rates of all age groups were more even, having only slight peaks in summer and autumn. The seasonal variation in growth ofPontoporeia spp. was not directly dependent on sedimentation of the spring phytoplankton bloom, but was more closely connected with seasonality of temperature and total benthic metabolism. Because estimates of annual food consumption byPontoporeia spp. exceeded bacterial and meiofaunal production, these species must have derived the majority of their energy demands from dead organic detritus. BothPontoporeia species are probably strictly food-limited in the area. The cessation of growth seemed to coincide with the approach of a plateau of total biomass, suggesting food competition. Different timing of growth between the age groups ofP. affinis suggests differences in sources or utilization of food; differences betweenP. affinis adP. femorata may indicate interspecific competition or different food.     

Stoichiometry of mesozooplankton in N- and P-limited areas of the Baltic Sea

The Baltic Sea is a very suitable site for stoichiometric studies, since its subbasins differ in their concentration of elemental components, and primary production can therefore be either nitrogen or phosphorus limited. To reveal if the nutrient limitation of mesozooplankton mirrors that of the primary producers, carbon, nitrogen and phosphorus content of both seston and grazers (Acartia sp., Centropages hamatus, Daphnia cristata, Eurytemora affinis, Limnocalanus macrurus, Temora longicornis) were measured in midsummer in the Baltic proper, the Gulf of Finland and the Gulf of Bothnia. The mineral ratios of the different taxa were equal, apart from L. macrurus with notably higher C:P and N:P ratios. Molar C:N ratios were relatively stable (5.1-6.3), whereas C:P and N:P ratios fluctuated more (41-144 and 6.6-24). However, zooplankton elemental composition and limitation did not depend on the limiting nutrient of the phytoplankton, the seston mineral ratio or the sea area. Both the seston-zooplankton elemental imbalance and the food threshold ratio indicated phosphorus limitation of most of the grazers. While L. macrurus may be C or N limited, the possible P deficiency of the other studied taxa suggests that the Baltic Sea zooplankton may act as a potential phosphorus sink, as the freshwater secondary producers do.     

Electrophoretic survey of genetic variation in Pecten maximus,Chlamys opercularis,C. varia and C. distorta from the Irish Sea

Food selection by young larvae of the gulf menhaden (Brevoortia patronus) was studied in the laboratory at Beaufort, North Carolina (USA) in 1982 and 1983; this species is especially interesting, since the larvae began feeding on phytoplankton as well as microzooplankton. When dinoflagellates (Prorocentrum micans), tintinnids (Favella sp.), and N1 nauplii of a copepod (Acartia tonsa) were presented to laboratory-reared, larval menhaden (3.9 to 4.2 mm notochord length), the fish larvae ate dinoflagellates and tintinnids, but not copepod nauplii. Larvae showed significant (P<0.001) selection for the tintinnids. Given the same mixture of food items, larger larvae (6.4 mm notochord length) ate copepod nauplii as well as the other food organisms. These feeding responses are consistent with larval feeding in the northern Gulf of Mexico, where gulf menhaden larvae between 3 and 5 mm in notochord length frequently ate large numbers of dinoflagellates (mostly P. micans and P. compressum) and tintinnids (mostly Favella sp.), but did not eat copepod nauplii. As larvae grew, copepod nauplii and other food organisms became important, while dinoflagellates and tintinnids became relatively less important in the diet. Since the tintinnids and nauplii used in the laboratory feeding experiments were similar in size as well as carbon and nitrogen contents, the feeding selectivity and dietary ontogeny that we observed were likely due to a combination of prey capturability and larval fish maturation and learning.Contribution No. 5575 of the Woods Hole Oceanographic Institution     

Omnivorous feeding behavior of the Antarctic krill Euphausia superba

Feeding experiments were conducted at Palmer Station from December 1985 to February 1986 to examine the potential role of copepod prey as an alternative food source for Euphausia superba. Copepod concentration, copepod size, phytoplankton concentration, the duration of krill starvation and the volume of experimental vessels were altered to determine effects on ingestion and clearance rates. Krill allowed to feed on phytoplankton and copepods in 50-litre tubs showed greatly increased feeding rates relative to animals feeding in the much smaller volumes of water traditionally used for krill-feeding studies. Clearance rates on copepods remained constant over the range of concentrations offered, but clearance rates on phytoplankton increased linearly with phytoplankton concentration. Feeding rates increased when larger copepods were offered and when krill were starved for two weeks prior to experiments. Clearance rates of krill feeding on copepods were higher than, but not correlated with, their clearance rates on phytoplankton in the same vessel. E. superba may have a distinct mechanism for capturing copepods, perhaps through mechanoreception. Although our observed clearance rate of 1055 ml krill^-1 h^-1 indicates that krill can feed very efficiently on copepod prey, such feeding would meet less than 10% of their minimum metabolic requirements at the typical copepod concentrations reported for Antarctic waters. However, substantial energy could be gained if krill fed on the patches of high copepod concentrations occasionally reported during the austral summer, or if krill and copepods were concentrated beneath the sea ice during the winter or spring months. Our results, indicating efficient feeding on zooplankton and higher clearance rates on phytoplankton than previously believed, represent a step towards balancing the energy budget of E. superba in Antarctic waters.     

Variation in cadmium uptake by estuarine phytoplankton and transfer to the copepod Eurytemora affinis

The pathways of cadmium (Cd) uptake and transfer within an estuarine planktonic community from the Patuxent River, Maryland, USA, were investigated using an assemblage of natural phytoplankton and the copepod Eurytemora affinis Poppe. The experiment was carried out in October 1992 in replicated 500-liter, flow-through, fiberglass tanks. Growth rate, species composition, and Cd loading affected the accumulation of Cd by the phytoplankton. Uptake of Cd by phytoplankton was proportional to the amount of Cd available in the water column. Partition coefficients (K _d) for phytoplankton uptake averaged 4.4 × 10⁴. As metal loading rates and phytoplankton species composition changed during the 12-d experiment, Cd partitioning declined. Transfer of Cd to E. affinis occurred from Cd-laden phytoplankton, with levels in the copepods being approximately the same as, or somewhat less, than in the phytoplankton. Some Cd uptake occurred in copepods exposed to dissolved Cd only; however, the uptake was considerably less than that seen from food. Thus, Cd content of higher trophic levels, such as copepods, can be affected by the degree of Cd incorporation in their food source, and by ecological factors regulating phytoplankton ingestion. Received: 13 September 1995 / Accepted: 29 October 1998     

Pathways of arsenic uptake and incorporation in estuarine phytoplankton and the filter-feeding invertebrates Eurytemora affinis,Balanus improvisus and Crassostrea virginica

Arsenic uptake from water and from phytoplankton was followed in the copepod Eurytemora affinis and the barnacle Balanus improvisus collected from the Patuxent River estuary, Chesapeake Bay, eastern coast of the USA in 1987, and in the oyster Crassostrea virginica obtained from a hatchery on the shore of Chesapeake Bay in 1987. Dissolved arsenic was readily taken up by phytoplankton and by shell material of B. improvisus and C. virginica; however, no dissolved arsenic was incorporated into the invertebrate tissues. When E. affinis, B. improvisus and C. virginica were fed phytoplankton containing elevated arsenic contents, significant arsenic incorporation occurred. Juvenile B. improvisus incorporated relatively more arsenic than adults of all three species. Compared to the 100 to 200% increase in arsenic content by phytoplankton exposed to dissolved arsenic, the 25 to 50% increase in these invertebrate species via trophic transfer is relatively small. Even though the trophic pathway for arsenic transfer is the major one for higher trophic levels within an ecosystem, the potential for direct arsenic impact to trophic levels other than phytoplankton appears to be minimal.     

Succession and growth limitation of phytoplankton in the Gulf of Bothnia (Baltic Sea)

A one year field study of four stations in the Gulf of Bothnia during 1991 showed that the biomass was ca. two times, and primary productivity ca. four times, lower in the north (Bothnian Bay) than in the south (Bothnian Sea) during the summer. Nutrient addition experiments indicated phosphorus limitation of phytoplankton in the Bothanian Bay and the coastal areas in the northern Bothnian Sea, but nitrogen limitation in the open Bothanian Sea. A positive correlation between the phosphate concentration and the production/biomass ratio of phytoplankton was demonstrated, which partly explained the differences in the specific growth rate of the phytoplankton during the summer. Differences in photosynthetic active radiation between the stations also showed a covariation with the primary productivity. The relative importance of nutrient or light limitation for photosynthetic carbon fixation could not, however, the conclusively determined from this study. Marked differences in phytoplankton species composition from north to south were also observed. The number of dominating species was higher in the Bothnian Sea than in the Bothnian Bay. The distribution of some species could be explained as due to nutrient availability (e.g. Nodularia spumigena, Aphanizomenon sp.), while salinity probably limits the distribution of some limnic as well as marine species. The potentially toxic phytoplankton N. spumigena, Dinophysis acuminata and Chrysochromulina spp. were common in the Bothnian Sea but not in the Bothnian Bay. The pico- and nanoplankton biomass during late summer was higher than previously reported due to a revised carbon/volume ratio.     

Zooplankton feeding ecology: contents of fecal pellets of the copepods Temora turbinata and T. stylifera from continental shelf and slope waters near the mouth of the Mississippi River

In-situ feeding habits of the copepods Temora turbinata and T. stylifera were investigated by scanning electron microscope examination of fecal pellets, the contents of which reflected copepod gut contents upon capture. Pellet contents were compared with assemblages of available phytoplankton in the water column at the times of zooplankton sampling. Samples were collected in continental shelf and slope waters of the Gulf of Mexico near the mouth of the Mississippi River. Both species ingested a wide size range and taxonomic array of phytoplankters, and to a lesser extent, other crustaceans. Fecal pellets contained primarily the remains of the phytoplankters that were most abundant in the water at times of collection. There was considerable overlap in the food items ingested by adult females of both copepod species, or two stages of T. turbinata copepodites. Thus, T. turbinata and T. stylifera are omnivores, but primarily opportunistic herbivores.     

Growth and development of Calanus spp. (Copepoda) during spring phytoplankton succession in the North Sea

Experiments were carried out to determine growth and development rates of the herbivorous copepod Calanus finmarchicus (Gunnerus) under natural conditions during the phytoplankton spring bloom in the northern North Sea. From 28 April to 25 May 1983 copepodite stages I, IV and V were incubated for a 3-d period on board a ship in vessels with naturally occurring phytoplankton or cultured algae as food. Highest rates of growth and development were achieved while the diatom Chaetoceros sp. was the dominant phytoplankton organism. These rates decreased considerably when this chain-forming diatom was succeeded after one week by the small-celled diatom Thalassiosira conferta. Again one week later, during the bloom of the succeeding colonial microflagellate Corymbellus aureus, copepodite stage IV still managed to maintain moderate rates of growth and development, but these rates dropped to almost zero in CV, suggesting the start of a resting stage. Nevertheless, brood collected from this generation and from Calanus helgolandicus (Claus) was raised in the laboratory to the adult stage at high speed. Since temperature and the total phytoplankton concentration in the sea remained almost constant it seems that the retardation and arrestment of growth and development were an immediate response to a qualitative change of the food composition related to the successive blooms of different algal species.     

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.     

Analysis of the biological cycle of Acartia clausi (Copepoda) in a meso-oligotrophic coastal area of the eastern Mediterranean Sea using time-series analysis

We evaluated the duration of Copepodite Stages C1 to C6, the biological cycle and the number of annual generations of the planktonic copepod Acartia clausi in a meso-oligotrophic area of the eastern Mediterranean Sea (Saronikos Gulf, Greece). The results were based on 95 zooplankton samples collected during the period November 1988 through June 1990, at intervals of 1, 2, 7 and 15 d, the sampling intervals being dependent on the abundance of A. clausi. Time-series analysis (cross-correlation) of fluctuations in the comparative abundance (percentages) of the copepodite stages present was used to determine the duration of the development stages and generation length. This methodology could significantly contribute to the identification of cohorts, and hence to the estimation of stage duration, from field data for a given copepod species. The development of A. clausi stages was not isochronal; duration of the first copepodite stage was shorter than that of the last three stages. The mean generation length estimated (28.6 d) is among the highest recorded in the literature for A. clausi at the range of temperatures prevailing in the area (13 to 25°C). Throughout the year there were four or five generations. The possible limiting role of food availability on the duration of each stage and hence on generation length is also discussed.     

The Baltic Sea spring phytoplankton bloom in a changing climate: an experimental approach

The response of the Baltic Sea spring bloom was studied in mesocosm experiments, where temperatures were elevated up to 6°C above the present-day sea surface temperature of the spring bloom season. Four of the seven experiments were carried out at different light levels (32–202?Wh?m^?2 at the start of the experiments) in the different experimental years. In one further experiment, the factors light and temperature were crossed, and in one experiment, the factors density of overwintering zooplankton and temperature were crossed. Overall, there was a slight temporal acceleration of the phytoplankton spring bloom, a decline of peak biomass and a decline of mean cell size with warming. The temperature influence on phytoplankton bloom timing, biomass and size structure was qualitatively highly robust across experiments. The dependence of timing, biomass, and size structure on initial conditions was tested by multiple regression analysis of the y-temperature regressions with the candidate independent variables initial light, initial phytoplankton biomass, initial microzooplankton biomass, and initial mesozooplankton (=copepod) biomass. The bloom timing predicted for mean temperatures (5.28°C) depended on light. The peak biomass showed a strong positive dependence on light and a weaker negative dependence on initial copepod density. Mean phytoplankton cell size predicted for the mean temperature responded positively to light and negatively to copepod density. The anticipated mismatch between phytoplankton supply and food demand by newly hatched copepod nauplii occurred only under the combination of low light and warm temperatures. The analysis presented here confirms earlier conclusions about temperature responses that are based on subsets of our experimental series. However, only the comprehensive analysis across all experiments highlights the importance of the factor light.     

The influence of temperature and salinity on<Emphasis Type="Italic"> Acartia</Emphasis> (Copepoda: Calanoida) nauplii survival

Adult Acartia congeners, A. bifilosa, A. clausi, A. discaudata and A. tonsa, have distinct seasonal and spatial distribution patterns in Southampton Water (UK), reflecting patterns of temperature and salinity, respectively. The effect of these factors on other life stages, hatch success and naupliar survival was investigated by exposing the congeners to a range of salinity (15.5–33.3) and of temperature (5–20°C). A. clausi is known to prefer more saline waters, and showed highest hatch success at 33.3 salinity. A. tonsa is most tolerant to dilution, and at 15.5 salinity it had the highest hatch success of all the congeners. Hatch success in both A. bifilosa and A. discaudata was similar over the range of salinities investigated, confirming that they are intermediate species in terms of spatial distribution. The nauplii of all species survived well at the higher salinities and best at 33.3, which allows for differential transport of the poorly swimming nauplii to the mouth of the estuary until size and swimming ability increase, after which they can then return to regions of preferred salinity. The summer species, A. clausi and A. tonsa showed higher hatch success at 20°C, whereas A. discaudata, which is present in the water column all year round, showed no significant temperature-related differences in hatch success. A. bifilosa, which diapauses over summer, showed significantly higher hatch success at 10°C than at 20°C. The physiological relationship between temperature and development time was clear; naupliar survival of all species was highest at 20°C and all congeners reached the first copepodite stage (CI) significantly faster at 20°C. However, no consistent pattern was seen for salinity. It would appear that the adult Acartidae in Southampton Water remain in regions of their preferred salinity and lay eggs there which hatch well. However, because the nauplii are not good swimmers, they are swept towards the mouth of the estuary and into areas of higher salinity, where they remain and develop into more advanced stages before moving back up the estuary to take up their adult distribution pattern.Communicated by J.P. Thorpe, Port Erin     

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.     

Seasonal grazing of Pseudocalanus minutus on particles

Grazing by Pseudocalanus minutus on naturally occurring partieles has been investigated over a 2 year period at 5 m depth, in a small coastal embayment in Nova Scotia, Canada. Large variations in the standing stock of particulate matter in the water from 5 m depth were associated with seasonal changes in phytoplankton, and at times with runoff from rivers. Each size group displayed seasonal changes, with high amplitudes occurring in the large-particle size ranges. P. minutus consumption was associated with seasonal changes in total particle concentration as well as with the concentration in each size group. The food uptake was correlated (P>0.01) with the particle concentration in all particle categories, except in the size range below 3.57 . Frequency of positive electivity indices increased with particle sizes up to 57 and then decreased. Observations on particle spectra revealed considerable seasonal variability in both particle sizes and copepod feeding patterns. P. minutus adapted to seasonal variations within the particle spectrum by shifting its grazing pressure from one size range to another. By taking advantage of every particle peak concentration, P. minutus revealed a strong opportunistic feeding behaviour, and a very efficient utilization of the standing stock. Maximum consumption was recorded in early spring, when P. minutus fed on large-size particles. Feeding took place in the medium and small-size ranges during the summer and part of fall. Food uptake was rarely less than 2.26% of body weight during the winter, and reached up to 55% and sometimes more in spring. The unselective feeding patterus demonstrated by P. minutus suggest certain ecological implications of feeding pressure on standing stock.     

Physioecology of zooplankton. II. Effects of phytoplankton concentration,temperature, and body size on the development and molting rates of Calanus pacificus and Pseudocalanus sp.

Developmental time and stage duration for Calanus pacificus Brodsky and Pseudocalanus sp. and the rate of loss of body carbon by molting for C. pacificus were estimated for copepodite stages cultured under various combinations of phytoplankton concentration and temperature. Mean development time and stage duration for C. pacificus decreased hyperbolically with increasing food concentration, and the minimum time required for reaching a given stage decreased logarithmically with a logarithmic increase in temperature. Low temperature retarded the development of early stages proportionally more than that of late stages, and stage duration increased logarithmically with increasing body weight. Therefore, copepodite development was not isochronal. The rate of loss of body carbon by molting was small, ranging from 0.2 to 2% day^-1. This rate increased hyperbolically with food concentration and was linearly related to the growth rate. The critical food concentration for the rates of development and molting increased with temperature and stage of development, but these rates were less dependent on food concentration than the growth rate. The development rate of Pseudocalanus sp. was higher than that of C. pacificus, and was less influenced by changes in food concentration and temperature. It is postulated that the inverse relationship between temperature and body size results from a differential effect of temperature and body size on the rates of growth and development. That is, with increasing body size the growth rate tends to become temperature-independent, but the development rate remains proportional to temperature. Thus, copepodites growing at low temperature can experience a greater weight increment between molting periods than individuals growing at high temperature, because the growth rate is similar at all temperatures but stage duration is longer at low temperature.Contribution No. 1128 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Physioecology of zooplankton. I. Effects of phytoplankton concentration,temperature, and body size on the growth rate of Calanus pacificus and Pseudocalanus sp.

Changes in dry weight and in weight-specific growth rates were measured for copepodite stages of Calanus pacificus Brodsky and Pseudocalanus sp. cultured under various combinations of phytoplankton concentration and temperature. Mean dry weight of early copepodites was relatively unaffected by either food concentration or temperature, but mean dry weight of late stages increased hyperbolically with food concentration and was inversely related to temperature. The food concentration at which maximum body weight was attained increased with increasing temperature and body size, and it was considerably higher for C. pacificus than for Pseudocalanus sp. This suggests that final body size of small species of copepods may be determined primarily by temperature, whereas final body size of large species may be more dependent on food concentration than on temperature. Individual body weight increased sigmoidally with age. The weight-specific growth rate increased hyperbolically with food concentration. The maximum growth rate decreased logarithmically with a linear increase in body weight, and the slope of the lines was proportional to temperature. The critical food concentration for growth increased with body size proportionally more at high than at low temperature, and it was considerably higher for C. pacificus than for Pseudocalanus sp. Because of these interactions, early copepodites optimized growth at high temperature, even at low food concentrations, but under similar food conditions late stages attained higher growth at low temperature. The same growth patterns were found for both species, but the rates were significantly higher for the larger species, C. pacificus, than for the smaller one, Pseudocalanus sp. On the basis of findings in this study and of analyses of relationships between the maximum growth rate, body size, and temperature from other studies it is postulated (1) that the extrapolation of growth rates from one species to another on the basis of similarity in body size is not justified, even for taxonomically related species; (2) that the allometric model is inadequate for describing the relationship between the maximum weight-specific growth rate and body size at the intraspecific level; (3) that the body-size dependence of this rate is strongly influenced by temperature; and (4) that species of zooplankton seem to be geographically and vertically distributed, in relation to body size and food availability, to optimize growth rates at various stages of their life cycles.Contribution No. 1127 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA