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.     

Vertical distributions of Calanus finmarchicus and C. helgolandicus (Crustacea: Copepoda)

The vertical distributions of the spring populations of Calanus finmarchicus (Gunnerus) and C. helgolandicus Claus are described and compared. The differences we observed between the two species have probably confused the understanding of the vertical distribution and development of the populations of Calanus spp. in the shelf seas around the United Kingdom where the species occur together. The results imply that these two congeneric species have different behaviour patterns which minimise interspecific competition where the species have sympatric distributions. C. finmarchicus has its younger development stages overlying the older stages in the water column. In C. helgolandicus the converse is true; i. e., the majority of the populations of Stage I and II copepodites of the first spring generations are found below the thermocline. It is also suggested that the different behaviour patterns lead to different feeding regimes and strategies.JONSDAP Contribution No. 52     

Vertical distribution and nocturnal migration of zooplankton in relation to the development of the seasonal thermocline in Patraikos Gulf

Vertical distribution and nocturnal migration of zooplankton species in relation to the development of the seasonal thermocline in the shallow waters (90 m) of Patraikos Gulf (Ionian Sea, Greece) were investigated using a WP-2 closing net. Juvenile and adult copepods accounted for a mean of 91% of the total collected in three sampling periods, i.e. May, July and September 1985.Ctenocalananus vanus, Paracalanus parvus andOithona plumifera were the dominant copepods. The majority of the zooplankton tend to aggregate at the thermocline layer. Among copepods the two congeneric speciesClausocalanus pergens andC. furcatus exhibited different migratory responses to the development of the thermocline.C. pergens occurred in the lower part of the thermocline andC. furcatus in the upper region or above. The diel vertical migration of all species could be divided into four types: (1) no vertical migration; (2) upward migration at night; (3) occasional migration; and (4) reverse migration (down at night). In July when the strongest thermocline developed, most zooplankters rose close to the surface at night. For most species, temperature discontinuity did not limit their diel migration.Please address all correspondence and requests for reprints to Dr J.J. Lykakis     

Calanoid copepod eggs in sea-bottom muds. II. Seasonal cycles of abundance in the populations of several species of copepods and their eggs in the Inland Sea of Japan

The seasonal cycles of abundance of populations of dominant calanoid copepods in the water column and of their eggs recovered from the bottom sediment in the central part of the Inland Sea of Japan are described. The numbers of both copepods and eggs fluctuated markedly with season in an essentially similar pattern among the 6 species studied (Tortanus forcipatus Giesbrecht, Calanopia thompsoni A. Scott, Acartia erythraea Giesbrecht, A. clausi Giesbrecht, Centropages abdominalis Sato, C. yamadai Mori). The density of eggs in the sea bottom was highest shortly before the population of adults and late copepodids disappeared from the plankton; the numbers of eggs then gradually decreased until the appearance of the next planktonic population.     

Variations in the abundance of Centropages typicus and Calanus helgolandicus in the North Sea: deviations from close relationships with temperature

The interannual variations in abundance of the calanoid copepod Centropages typicus (Krøyer) in the North Sea are significantly positively correlated with sea-surface temperature. The same is true of Calanus helgolandicus Claus. In both cases there was a deviation from the close relationship with temperature for several years in the early to mid-1980s. Centropages typicus was less abundant than expected from 1981 to 1984, but Calanus helgolandicus recovered from a period of lower than expected abundance from 1980 to 1982, to become more abundant than expected from 1983 to 1987. The difference between the responses of the two species to climate-driven changes in seasonal cycles of phytoplankton is interpreted in relation to their biology. Geographical variations of the interannual variability and seasonal cycles of Centropages typicus in the North Sea are analysed. The centre of the population of this species in the North Sea is in the German and Southern Bights, but the variations in abundance may be partly influenced by advection from areas to the west of Britain.     

Zooplankton of the Bristol Channel and Severn Estuary. The distribution of four copepods in relation to salinity

The seasonal variations in distribution and abundance of the common zooplankton species in the Bristol Channel and Severn Estuary were related to the salinity regimes observed over the period November 1973 to February 1975. The dominant constituents in all regions were the calanoid copepods, which reached maximum densities in July: approximately 100 times their winter levels. Four zooplankton assemblages were recognised using an objective classification program which computed similarity coefficients and used group-average sorting. The assemblages existed along the salinity gradient observed from the Severn Estuary to the Celtic Sea. The assemblages were classified as true estuarine, estuarine and marine, euryhaline marine and stenohaline marine and were characterized by the copepods Eurytemora affinis (Poppe) (<30S), Acartia bifilosa var. inermis (rose) (27 to 33.5S), Centropages hamatus (Lilljeborg) (31 to 35S) and Calanus helgolandicus (Claus) (>33S), respectively.     

Vertical distribution,and seasonal and diurnal migration of Calanus helgolandicus in the Celtic Sea

The vertical distribution and migration (seasonal, diel and ontogenetic) of Calanus helgolandicus are described from the shallow (100 m) shelf-seas to the south-west of the British Isles. In 1978 and 1979, the overwintering population of C. helgolandicus consisted primarily of Stage V copepodites and adults. By late winter/early spring the copepodites had moulted to adult females (>90%), which matured and bred the first cohorts of the year, prior to onset of the spring phytoplankton bloom in April/May. C. helgolandicus reached a peak of numerical abundance in August of 20x10³ copepodites m^-2 (over the depth range sampled -0 to 70 m), which was 200 times the population in winter. The seasonal peak of abundance occurred 4 mo after the peak of the bloom of phytoplankton in spring. The yearly development of the copepod was not always out of phase with the diatom bloom, as seen when the data from 1978 was placed in the context of a longer time-series collected at 10 m over 22 yr (1960–1981, inclusive). Large vertical migrations were observed in the younger copepodites (CI and II) in May from below to above the thermocline. In the remainder of the year, the CI and CII stages behaved differently and were located above the thermocline within the euphotic zone. The largest vertical displacements of biomass were seen in the summer months due to the migrations of the CV stages and adults, which had developed from the spring cohorts. It was contended that the seasonal and vertical migrations of C. helgolandicus are part of a more complex pattern of inherent behavior than has been reported previously and that, however difficult this is to discern in the natural populations, it always expresses itself.     

Live discrimination of Calanus glacialis and C. finmarchicus females: can we trust phenological differences?

Two key players in the Arctic and subarctic marine ecosystem are the calanoid copepods, Calanus finmarchicus and C. glacialis. Although morphologically very similar, these sibling species have different life cycles and roles in the Arctic pelagic marine ecosystem. Considering that the distribution of C. glacialis corresponds to Arctic water masses and C. finmarchicus to Atlantic water masses, the species are frequently used as climate indicators. Consequently, correct identification of the two species is essential if we want to understand climate-impacted changes on Calanus-dominated marine ecosystems such as the Arctic. Here, we present a novel morphological character (redness) to distinguish live females of C. glacialis and C. finmarchicus and compare it to morphological (prosome length) and genetic identification. The characters are tested on 300 live females of C. glacialis and C. finmarchicus from Disko Bay, western Greenland. Our analysis confirms that length cannot be used as a stand-alone criterion for separation. The results based on the new morphological character were verified genetically using a single mitochondrial marker (16S) and nuclear loci (six microsatellites and 12 InDels). The pigmentation criterion was also used on individuals (n = 89) from Young Sound fjord, northeast Greenland to determine whether the technique was viable in different geographical locations. Genetic markers based on mitochondrial and nuclear loci were corroborative in their identification of individuals and revealed no hybrids. Molecular identification confirmed that live females of the two species from Greenlandic waters, both East and West, can easily be separated by the red pigmentation of the antenna and somites of C. glacialis in contrast to the pale opaque antenna and somites of C. finmarchicus, confirming that the pigmentation criterion is valid for separation of the two species.     

Geographical range and taxonomic divergence in North Atlantic Calanus (C. helgolandicus,C. finmarchicus and C. glacialis)

The known distribution of Calanus helgolandicus in the North Atlantic Drift is difficult to explain in the absence of a reproductively active population inhabiting continental waters off eastern North America. New evidece indicates that this population, overlooked in the past, does in fact exist. The species has been found in a study of zooplankton samples from a number of MARMAP (Marine Resource Monitoring, Assessment, and Prediction Program) cruises surveying ichthyoplankton between Cape Hatteras and the New York Bight. Sexual activity in these stocks of C. helgolandicus was indicated by the ripeness of ovaries, the frequency of males, the presence of sperm in the females' seminal receptacles and the appearance of a female bearing a Calanus spermatophore. The new records provide a likely origin for the presence of the species in the vicinity of the Labrador Grand Banks as well as further east in the North Atlantic Drift. C. finmarchicus, similarly sexually active, was found to be sympatric with C. helgolandicus in the MARMAP collections which were taken from a region contiguous with the southernmost, known distribution, of C. glacialis. The distribution of integumental organs (i.e., pore signature patterns) was examined in the three species to determine whether they would be taxonomically useful. Strikingly different patterns were found on the female urosome. Pore signature differences between the polar species C. glacialis and the temperate C. helgolandicus proved to be as pronounced as those between the boreal C. finmarchicus and its two neighboring species. The successively overlapping ranges and the distinctive differences in pore signature patterns suggest that divergence from the generic pattern of integumental organ distribution has been a product of selection against hybridizing among the three species. If this is in fact the case, the reproductive range of C. helgolandicus has overlapped with those of C. finmarchicus and C. glacialis for appreciable periods in the history of the three species.     

Simple molecular method to distinguish the identity of Calanus species (Copepoda: Calanoida) at any developmental stage

Diagnostic morphological characteristics of copepods of the genus Calanus are restricted largely to minor variations in secondary sex characteristics. This presents a persistent problem in the identification of individuals to species level, especially for immature stages. We have developed a simple molecular technique to distinguish between the North Atlantic Calanus species (C. helgolandicus, C. finmarchicus, C. glacialis and C. hyperboreus) at any life stage. Using the polymerase chain-reaction (PCR), the mitochodrial large subunit (16S) ribosomal RNA (rRNA) gene was amplified from individual copepods preserved in ethanol. Subsequent digestion of the amplified products with the restriction enzymes DdeI and VspI, followed by electrophoretic separation in 2% agarose (Metaphor, FMC Ltd), produced a characteristic pattern for each species. The versatility of the method is demonstrated by the unambiguous identification to species of any life stage, from egg to adult, and of individual body parts. Received: 11 May 1998 / Accepted: 5 August 1998     

Suspended particulates in the Irish Sea and feeding conditions for fish larvae

The size distributions (2 to 160 m equivalent spherical diameter) of suspended particulate material sampled on two cruises along a transect in the Irish Sea in 1988 are described in relation to hydrographic conditions, chlorophylla concentration and carbon to nitrogen ratios. Particulates were more abundant and larger size modes in the distribution were more evident, in the upper mixed layer of stratified water than in areas where the water column was fully mixed. The detrital content was estimated at 52% of total particulate matter above the thermocline in stratified regions and at around 97% at mixed water sites. In stratified regions the predominance of larger sized phytoplankton and lower levels of detritus is argued to support a more direct and efficient transfer of energy to fish larvae via larger sizes of copepods. Conversely, in mixed areas of high detrital loading the smaller size spectrum of particulates incorporates a less efficient transfer of energy through bacterial cycling and smaller copepods.     

Large-scale oceanic distribution and population structure of Calanus finmarchicus, in relation to physical environment, food and predators

The investigation was carried out from 62°N to 73°N and from 14°E to 11°W in the Norwegian Sea during 19 June-12 July 1997. Regional differences in the phase of the seasonal development of the plankton community were evident, most pronounced across the Arctic front. In the Coastal and eastern Atlantic domains, post-bloom conditions prevailed, characterised by low chlorophyll a (chl a) levels and a phytoplankton assemblage dominated by coccolithophorids and small flagellates. During the study period, egg production rates of Calanus finmarchicus were low (<10 eggs female^-1 day^-1), older copepodite stages dominated, and the seasonal descent to deeper waters had started. In the Arctic domain, bloom conditions were evident by high chl a levels and a high abundance of large diatoms. Egg production rates were higher (a maximum of 29 eggs female^-1 day^-1), but the dominance of stages CI-CIII indicated that considerable spawning had already occurred prior to the spring bloom. The seasonal descent had barely started. Both invertebrate and fish predators were most abundant in the Coastal and eastern Atlantic domains, with abundance strongly decreasing north-westwards. No tight relationship between total abundance of invertebrate or fish predators and that of C. finmarchicus was apparent. However, a weak, but significant, relationship between abundance of young stages of chaetognaths and Euchaeta spp. versus young stages of C. finmarchicus was found, indicating that these invertebrate predators develop parallel to the development of the new cohort of C. finmarchicus. In early summer, C. finmarchicus had reached overwintering stages, and had started to accumulate in deeper waters in areas with the highest abundance of horizontally migratory planktivorous fish.     

Biosynthesis of trimethylamine oxide in calanoid copepods. seasonal changes in trimethylamine monooxygenase activity

Live copepods, Calanus finmarchicus (Gunnerus) and C. hyperboreus (Krøyer), exposed to dissolved ¹⁴C-labeled trimethylamine (TMA) in seawater, oxidized TMA to trimethylamine oxide (TMAO), which accumulated in the organisms. The amount of TMAO synthesized was dependent on the time of exposure to TMA and the concentration of TMA in the seawater. It was inferred that copepods can produce TMAO by oxidation of TMA found in their plant food. Choline and methionine did not appear to be of importance as precursors of TMAO. There were large seasonal changes in TMA monooxygenase activity of both copepods. The activity was high in spring, and decreased through summer and autumn to a winter low in October to March. The changes in TMAO content of C. finmarchicus were, in comparison, small (this aspect was not tested for C. hyperboreus).     

Short-term effects of sucralose on Calanus finmarchicus and Calanus glacialis in Disko Bay,Greenland

The potential effects of sucralose on the Arctic copepods Calanus finmarchicus and Calanus glacialis were studied in Disko Bay, Greenland. Sucralose is a non-calorie sweetener and chlorine derivate of sucrose containing three chlorine atoms. Scandinavian screening studies of sucralose in 2007, revealed sucralose in all effluent samples. To investigate whether sucralose is harmful to the Arctic aquatic ecosystems, possible short-term effects were investigated on egg production, hatching rate, food intake and mortality of two species of Arctic copepods. The copepods were exposed to six different concentrations (0–50,000 ng · L^?1) of sucralose, which spans the range of concentrations found in the screening studies. Exposure led to no mortality among the copepods. Food intake by C. glacialis increased with increasing concentrations of sucralose. In C. finmarchicus, food intake did not differ with increasing concentrations. No effect of sucralose was observed on egg production of C. finmarchicus. Despite increased food intake with increasing concentrations of sucralose, C. glacialis did not increase its egg production. The results show that both species responded weakly to sucralose, but with C. glacialis being possibly slightly more sensitive to sucralose than C. finmarchicus.     

Molecular systematics of six Calanus and three Metridia species (Calanoida: Copepoda)

The discrimination of species of the copepod genus, Calanus (Copepoda; Calanoida), is problematical-especially in regions of sympatry. Although the species of Calanus exhibit exceptional morphological similarity, they are quite distinct in genetic character. The DNA base sequences of the mitochondrial large subunit (16S) ribosomal RNA (rRNA) gene unambiguously discriminated C. finmarchicus (Gunnerus 1765), C. glacialis (Jaschnov 1955), C. marshallae (Frost 1974), C. helgolandicus (Claus 1863), C. pacificus (Brodsky 1948), C. sinicus (Brodsky 1965), and C. hyperboreus (Kroyer 1838). Sequence differences among Calanus species for this gene portion range from 7.3% (between C. glacialis and C. marshallae) to 23.9% (between C. glacialis and C. sinicus). Differences among conspecific individuals were approximately 1 to 2%. [These sequence data were determined between April and November 1993; the sequenced domain is similar to that published previously in Bucklin et al. (1992) but are derived from analysis of additional individuals.] Statistical analysis of the sequence data using a variety of tree-building algorithms separated the taxa into one group of species corresponding to the C. finmarchicus group (C. finmarchicus, C. marshallae, and C. glacialis) and another ungrouped set of species corresponding to the C. helgolandicus group (C. helgolandicus, C. pacificus, and C. sinicus). The C. helgolandicus group may be older than the C. finmarchicus group, making the tree topology less reliable in this area. Calanus hyperboreus was an outlier; Nannocalanus minor (Claus 1863) was the outgroup. Similar analysis of Metridia species confirmed that M. lucens (Boeck 1864) and M. pacifica (Brodsky 1948) are distinct species; M. longa (Lubbock 1854) was still more divergent. These sequence data will allow the design of simple, molecular tools for taxonomic identifications. Diagnostic characters, assayed by rapid molecular protocols, will enable biological oceanographers to answer important questions about the distribution and abundance of all life stages (as well as patterns of reproduction) of morphologically similar species, such as those of Calanus.     

Influences of abundance,behavior, species composition,and ontogenetic stage on active emergence of meiobenthic copepods in subtropical habitats

The emergence of meiobenthic copepods in subtidal sand and seagrass sites in Tampa Bay, Florida (USA), was investigated on nine dates from March 1983 to August 1984. Numbers of emerging copepods were dependent on habitat, date and diel sampling period. Greater numbers typically emerged in seagrass sites, and differences between habitats may be related to lower copepod abundances in sand sites. Both copepod abundance and behavior significantly affected numbers emerging among dates. An increase in emergent behavior alone resulted in consistently greater numbers emerging during postsunset periods. Over 30 species in 15 families were found to emerge. Total numbers of emerging copepods were affected by the sporadic presence and seasonal behavior of certain species. Numbers of the predominant speciesParadactylopodia brevicornis, Tisbe furcata, Parategastes sp.,Mesochra pygmea, Laophontid sp., andHarpacticus sp. emerging depended on differences in both species abundance and behavior. The emergent behavior ofParategastes sp.,Harpacticus sp., andT. furcata also was influenced by total copepod densities in the sediment. Juvenile copepods constituted only a small proportion of the total numbers emerging. The consistent postsunset entry of between 10³ to 10⁵ copepods m^?2 into the water column in subtidal sand and seagrass habitats will contribute to the increased probability of copepod dispersal and reassortment of the benthic community, while providing a pathway for benthic-pelagic exchange.     

Winter studies on zooplankton in Arctic seas: the Storfjord (Svalbard) and adjacent ice-covered Barents Sea

Zooplankton was sampled in the Storfjord and ice-covered Barents Sea during March 2003. Environmental conditions represented a typical winter situation with low air temperatures, close pack ice, and extremely low chlorophyll concentrations. Polar water dominated the hydrographic regime in the upper layers. Zooplankton distribution reflected spatial variability of hydrography. The copepods Pseudocalanus spp., Oithona similis, Microsetella norvegica together with gastropod larvae were most numerous. Biomass averaged for the entire water column varied from 3.3 to 14.3 mg dry mass m⁻³, Calanus glacialis and Parasagitta elegans contributed most, followed by C. finmarchicus, Oithona similis, and Pseudocalanus spp. Various holoplankters showed reproductive activity, especially cyclopoid copepods and chaetognaths. A few C. glacialis females laid eggs in situ, but when fed diatom cultures rapidly increased their egg production. Meroplankton including larvae of nudibranchia, bivalvia, ophiuroida, polychaeta, and bryozoa were also present. Our data demonstrate that the pelagic community of the seasonally ice-covered Barents Sea was not in a “sleeping” state at the end of the winter, but in addition to dormant stages, a portion of mainly omnivorous and several carnivorous species was reproducing.     

Continuous plankton records: overwintering and annual fluctuations in the abundance of zooplankton

A study of seasonal variations in the pattern of year-to-year changes, for 1948 to 1982, in the abundance of four zooplankton taxa (Pseudocalanus elongatus Acartia clausi, Calanus finmarchicus and Hyperiidea) for the North Sea suggests that an element of the variations, represented at least in part by a quasi-linear downward trend, has its origins in winter and its occurrence through the year is a function of inherent persistence. A downward linear trend is the dominant pattern in the annual fluctuations in abundance of many species of zooplankton throughout the north-east Atlantic and the North Sea. It is argued that it is reasonable to extrapolate from the limited data set included in the detailed study and to assume a winter origin for the pattern wherever it occurs. Some implications for ideas about the dynamics of the plankton ecosystem are considered.     

Seasonal population structure,vertical distribution and migration of the chaetognath Sagitta elegans in the Celtic Sea

The biology of the chaetognath Sagitta elegans Verrill has been much researched, but detailed studies of population structure have generally been conducted in coastal water where dynamic tidal conditions may cause difficulty in interpretation of data. The resolution of sampling examining vertical distribution and diurnal migration has also been rather coarse. During a series of eight cruises to a seasonally thermally stratified sampling site in the Celtic Sea in 1978 and 1979, detailed vertical zooplankton profiles were taken to study the seasonal population structure, vertical distribution and migration of this species. The overwintering stock of S. elegans (22 to 52 individuals m^-2, 0 to 90 m) had a wide range of lengths (5 to 20 mm) and matured in 1978 from early March, spawning several times before dying out by late July. Young produced by the overwintering stock started to mature in July and population numbers reached their highest in August (2483 m^-2, 132.8 mg C m^-2) when sea temperature peaked (17.1°C). By October, the population of S. elegans declined (284 m^-2), which was thought to be due to a combination of lower sea-water temperature, competition for and availability of food, and predation. Because of the length range of the overwintering population (5 to 20 mm), it is assumed that reproduction continued at a low level over the winter, although eggs were not found in January and February, the coldest months of the year. In summer, the smallest S. elegans (2 to 6 mm) were found in the near-surface waters and did not migrate, but as their lengths increased they occupied deeper depth ranges and a portion of the population started to migrate diurnally. Individuals which did not migrate and stayed in the warmer surface waters, or those which migrated into it, matured faster than those remaining in the colder water below the thermocline. Migration to surface waters by mature individuals seemed to be stopped by high surface temperatures (17°C) and a sharp thermocline (3 C°). As sea temperature increased during the year from the winter minimum of 7.7°C, S. elegans matured at a progressively shorter length (14 mm in March 1978 to 10 mm in August). There are probably only three generations of S. elegans a year in the Celtic Sea.     

Seasonal variations in the densities of fecal pellets produced by Oikopleura vanhoeffeni (C. Larvacea) and Calanus finmarchicus (C. Copepoda)

Two abundant macrozooplankters, Oikopleura vanhoeffeni (Lohmann) and Calanus finmarchicus (Gunnerus) were collected from the coastal waters off Newfoundland in different seasons during 1990–1991 and incubated in natural seawater to collect freshly egested, field produced, fecal pellets. The densities of fecal pellets from O. vanhoeffeni and C. finmarchicus were measured in an isosmotic density gradient. These are the first reported seasonal measurements of fecal pellet densities from two different types of macrozooplankters, O. vanhoeffeni, a larvacean, filter feeder and C. finmarchicus, a crustacean, suspension feeder. Pellet density ranges and medians were significantly different among seasons for both species, depending primarily on the type of phytoplankton ingested and its ability to be compacted. Winter O. vanhoeffeni and fall C. finmarchicus feces filled with nanoplankters and soft bodied organisms had less open space [packing index (% open area) = 3.5 and 4% for O. vanhoeffeni and C. finmarchicus, respectively] and were more dense (1.23 and 1.19 g cm^-3) than spring feces filled with diatoms (packing index = 15 and 23%, density = 1.13 and 1.11 gcm^-3). For copepods, these results contrast with previously published density values and with the predicted copepod fecal pellet density calculated, in the present study, using the conventional mass/volume relationship. Copepod spring and summer diatom-filled feces had a calculated density of 1.12 and 1.24 gcm^-3 vs a measured median density of 1.11 gcm^-3 for both spring and summer feces; the fall feces containing nanoplankters had a calculated density of 1.08 gcm^-3 vs a measured median density of 1.19 gcm^-3. Knowledge of the seasonal variations in fecal pellet densities is important for the development of flux models.