Sinking rates of natural copepod fecal pellets

Many pure samples of natural fecal pellets have been collected from mixed small copepods and from the pontellid copepod Anomalocera patersoni in the Ligurian Sea, using a specially designed pellet collection device. Sinking rates of fresh pellets and pellets aged up to 33 days have been determined at 14°C, the mean temperature of the essentially isothermal water column in the Ligurian Sea. Sinking rates of pellets collected during calm sea states increased with increasing pellet volume, but sinking rates of pellets collected during rough sea (Beaufort scale 6) showed little correlation with pellet size. Much of the variability in the sinking rate-pellet size relationships was the result of different pellet composition and compaction, but not pellet age. Pellets produced from laboratory diets of phytoplankton and phytoplankton-sediment mixes showed the expected wide variability in sinking rates, with sediment-ballasted pellets sinking much faster than pellets produced from pure algal diets; thus determination of vertical material fluxes in the sea using laboratory-derived fecal pellet sinking rates is unwarranted. Natural pellet sinking data for small copepods and A. patersoni have been combined with similar data for euphausiids, to yield sinking rates of roughly two orders of magnitude over three orders of magnitude in pellet volume. Pellets from small copepods sank at speeds too slow to be of much consequence to rapid material flux to the deep sea, but they undoubtedly help determine upper water distribution of materials. Recalculation of fecal pellet mass flux estimates from the literature, using our sinking rate data for natural small copepod pellets, yielded estimates about half those of previously published values. Earlier studies had concluded that small fecal pellets were of lesser significance to total material flux than fecal matter; our recalculation strengthens that conclusion. Pellets from large copepods and euphausiids, however, have the capability to transport materials to great depths, and probably do not substantially recycle materials near the surface. The fact that the majority of pellets which had previously been collected in deep traps by other workers were of a size comparable to pellets from our large copepods supports the contention that these larger pellets are the main ones involved in vertical flux.     

Sinking rates of fecal pellets from the marine copepod Pontella meadii

Sinking rates of fecal pellets produced by the marine copepod Pontella meadii, grazing on 4 different phytoplankton diets, ranged from 15 to 153 m/day, with a mean of 66 m/day. Sinking rates, in general, were directly related to fecal pellet volumes, but unrelated to the diets used to produce the fecal pellets. There were two-to-threefold variations in sinking rates between fecal pellets of the same volumes, often produced on the same diets. Twenty repetitions of timed sinking of a single fecal pellet revealed sinking rates varying from 33 to 79 m/day, as well as variations in sinking rates within the course of individual descents. It is suggested that copepod fecal pellets are of such small volumes and densities that their sinking rates are subject to microstructural variations in the most carefully controlled water columns. Scanning electron microscope observations revealed lack of structural damage to some of the diatom frustules in the fecal pellets, suggesting that superfluous feeding may have occurred. Thus, the accelerated sinking rates of copepod fecal pellets may provide a mechanism for nutritional enrichment of the deep-sea ecosystem with organic parcels containing incompletely-assimilated plant material.     

210Po and 210Pb in zooplankton fecal pellets

²¹⁰Po and ²¹⁰Pb concentrations in fecal pellets from the zooplanktonic euphausiid Meganyctiphanes norvegica are reported. The ²¹⁰Po:²¹⁰Pb activity ratio is 2.2±0.3, a value in good agreement with that found in suspended particulate matter in surface seawater. Estimates of ²¹⁰Po and ²¹⁰Pb removal times from the mixed layer by fecal pellets alone yield values which are of the same order of magnitude as the removal times for these nuclides by all routes. It is suggested that there is a high probability that zooplanktonic fecal pellets play a significant role in the removal of both these nuclides from the surface layers of the ocean.     

Sinking rates and dissolution of midwater fish fecal matter

The sinking rates of fecal matter from 7 southern California midwater fish species were investigated. Feces were obtained from 162 specimens of Stenobrachius leucopsarus, Triphoturus mexicanus, Leuroglossus stilbius, Lampanyctus ritteri, Argyropelecus affinis and Parvilux ingens, which were collected in the Santa Barbara and San Clemente Basins between 1977 and 1979. In addition, feces obtained from 6 laboratory-maintained specimens of the midwater zoarcid Melanostigma pammelas were used for repeated sinking-rate measurements. The mean of the measured sinking rates for all species was 1.19 cm s^-1 (1 028 m d^-1), which is much higher than the known descent rates of euphausiid and copepod fecal pellets and of most other particulate organic detritus. Dissolution characteristics were also investigated for fecal matter from 4 species collected by the same series of net hauls: S. leucopsarus, T. mexicanus, A. affinis, and Sternoptyx obscura. The release of dissolved organic compounds from this material is low and does not represent a significant output during the relatively short time required to sink through the water column. These findings suggest that midwater fish fecal matter may represent a major source of organic transfer between the pelagic community and the benthos.     

Concentrations and vertical fluxes of zooplankton fecal pellets on a continental shelf

The contribution of fecal pellets to the benthos of the southeastern shelf of the USA is investigated through an analytic model which considers pellet production by different stage groups of the genus Paracalanus. Model results indicate that the concentration and vertical flux of pellets is a function of producer size and consumer size and abundance. Nauplii and adults, respectively, produce daily on the average 50 and 13% of total pellet mass, yet contribute 4 and 63%, respectively, to the daily pellet flux. Most of the pellets produced are consumed or degraded in the water column, with only 0.2% of the average daily primary production reaching the seafloor (35 m) as fecal pellets. This contributes to an impoverished benthos, such as that found on the southeastern continental shelf.     

A comparison of carbon based ammonia excretion rates between gelatinous and non-gelatinous zooplankton: Implications and consequences

About 560 literature data on weight-specific ammonia excretion rates of gelatinous zooplankton (cnidarians, ctenophores and salps) and non-gelatinous zooplankton (mainly crustacea) were converted to carbon based units to enabel a better comparison between both groups. If carbon is used as body-mass unit ammonia excretion rates of gelatinous zooplankton are in the same range as values obtained for other zooplankton taxa, indicating a similar nitrogen output per unit of organic matter in both groups. These results suggest nutrient regeneration potential to be the same in gelatinous and non-gelatinous zooplankton, and that nutrient regeneration within the pelagic system depends more upon the carbon biomass ratio between gelatinous and other zooplankton than on physiological differences.     

Distribution and trophic links of gelatinous zooplankton on Dogger Bank,North Sea

The ecology of small, gelatinous zooplankton is not integrated into management of Dogger Bank (54° 00′ N, 3° 25′ E to 55° 35′ N, 2° 20′ E). In pursuit of this goal, gelatinous zooplankton and their potential prey were sampled along a transect across the bank on June 10–16, 2007. Eleven species of small medusae and ctenophores were collected, with six abundant taxa occurring in greater numbers below the thermocline and in the shallower, southeastern portion of the bank. There were no statistically significant diel changes in distribution. In contrast, potential prey were distributed more evenly across the bank and throughout the water column. Isotopic analyses revealed that gelatinous zooplankton fed on both smaller (100–300 μm) and larger (>300 μm) mesozooplankton, but also potentially on each other. These ecological insights suggest that small medusae and ctenophores should be integrated into sustainable management of Dogger Bank.     

Bioluminescence spectra of shallow and deep-sea gelatinous zooplankton: ctenophores, medusae and siphonophores

We have examined the variability and potential adaptive significance of the wavelengths of light produced by gelatinous zooplankton. Bioluminescence spectra were measured from 100 species of planktonic cnidarians and ctenophores collected between 1 and 3500 m depth. Species averages of maximal wavelengths for all groups ranged from 440 to 506 nm. Ctenophores (41 species) had characteristically longer wavelengths than medusae (34 species), and the wavelengths from siphonophores (25 species) had a bimodal distribution across species. Four species each produced two different wavelengths of light, and in the siphonophore Abylopsistetragona these differences were associated with specific body regions. Light from deep-dwelling species had significantly shorter wavelengths than light from shallow species in both ctenophores (p = 0.010) and medusae (p = 0.009). Although light production in these organisms was limited to the blue-green wavelengths, it appears that within this range, colors are well-adapted to the particular environment which the species inhabit. Received: 27 April 1998 / Accepted: 27 October 1998     

Production,composition and sedimentation of salp fecal pellets in oceanic waters

Rates of fecal pellet production have been recorded from seven species of oceanic salps feeding on natural diets. Expressed as g C defecated per mg salp body C per hour, the values range between 3.7 and 27.7. Carbon: nitrogen ratios of the salp fecal pellets average 11.4; the organic matter of the pellets is mainly protein and carbohydrate. Sinking velocities of the pellets are very high, ranging from 320 to 2 238 m d^-1 for pellets from three species. However, the pellets sink slower than would be predicted from extrapolation of rates for crustacean pellets, probably due to the shape of the pellets and their density. The high rates of defecation, large size and rapid sedimentation of salp fecal pellets make them likely mechanisms for rapid transport of small particulate matter from surface waters to deep water and the benthos.     

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.     

The trophic linkage between zooplankton and benthic suspension feeders: direct evidence from analyses of bivalve faecal pellets

Using radiotracer (¹⁴C) and microscopic observation, we demonstrated that mussels (Mytilus edulis and Perna viridis) could be predators of mesozooplankton (rotifer Brachionus plicatilis). At radio-labelled rotifer densities of 0.1, 0.2, 0.5, 1.0 individual ml⁻¹, faecal pellets of mussels showed different degrees of radio signals and most of the faecal pellets were expelled 4 h after pulse feeding on rotifers. The maximum gut retention time (GRT) of ¹⁴C-labelled rotifers in the digestive diverticula did not o show any significant difference between the two mussel species or the different densities of rotifers, and the averaged GRT was 43.4±3.06 h (mean ± SE). At a rotifer density of 4.5 individual ml⁻¹, rotifer lorica pieces and rotifer bodies without eggs were found in faeces of M. edulis, while in the pseudofaeces, only complete rotifer bodies were found.     

Production and fate of copepod fecal pellets across the Southern Indian Ocean

The vertical distribution of copepods, fecal pellets and the fecal pellet production of copepods were measured at seven stations across the Southern Indian Ocean from productive areas off South Africa to oligotrophic waters off Northern Australia during October/November 2006. We quantified export of copepod fecal pellet from surface waters and how much was retained. Furthermore, the potential impact of Oncaea spp. and harpacticoid copepods on fecal pellets degradation was evaluated and found to be regional substantial. The highest copepod abundance and fecal pellet production was found in the western nutrient-rich stations close to South Africa and the lowest at the central oligotrophic stations. The in situ copepod fecal pellet production varied between 1 and 1,000 μg C m⁻³ day⁻¹. At all stations, the retention of fecal pellets in the upper 400 m of the water column was more than 99% and the vertical export of fecal pellets was low (<0.02 mg m⁻² day⁻¹).     

In situ grazing rates of deep-sea benthic boundary-layer zooplankton

In situ grazing rates of mixed assemblages of deep-sea benthic boundary-layer zooplankton were measured in July 1983 at a site in the North Atlantic Ocean at 2 175 m depth using a short-term radioisotope-uptake method. Zooplankton were collected with an opening-closing net system from the bottom 1 m of the ocean and incubated in situ with mixed tritiated amino acids in special cod-end chambers. Incubations were terminated at depth by the addition of MS-222. Radioisotope uptake beyond that of dead controls was shown by both the zooplankton and particulate fractions. Grazing rates in the deep-sea experiments were surprisingly high, being comparable to wintertime Narragansett Bay zooplankton grazing rates determined in a separate series of laboratory experiments. These laboratory experiments also documented nonparticle-associated uptake of dissolved amino acids by Narragansett Bay zooplankton, but the importance of this in the deep sea is unknown. The deep-sea benthic boundary layer may be a region of elevated rates and activity because of its higher particulate concentrations, and our experiments may also have measured maximum rather than average rates.     

Digestion rates and assimilation efficiencies of siphonophores fed zooplankton prey

Digestion times and assimilation efficiencies are critical factors used in calculations of carbon and nitrogen budgets. Digestion times of natural copepod prey differed significantly among 4 genera of siphonophores (P>0.001), from a minimum of 1.6 h to a maximum of 9.6 h. Assimilation efficiencies, in contrast, were uniformly high; 87 to 94% for carbon and 90 to 96% for nitrogen. Nitrogen assimilation was consistently greater than carbon assimilation. Assimilation efficiencies calculated according to dry weight substantially underestimated assimilation of carbon and nitrogen, while calculations using ash-free dry weights and ash-free to dry weight ratios approached values for carbon assimilation. These values are appreciably higher than most of the assimilation efficiencies previously measured for a few other planktonic carnivores. These results indicate very efficient digestion of food by siphonophores in oceanic environments where prey densities are low.     

Morphology and sinking velocities of fecal pellets of copepod, molluscan, euphausiid, and salp taxa in the northeastern tropical Atlantic

Morphological characteristics and sinking velocities of naturally occurring fecal pellets of copepods, euphausiids, salps, and pelagic mollusks collected in the northeastern tropical Atlantic were investigated during the period of May-June 1992. The fecal pellets of copepods and euphausiids were cylindrical and distinguished only by their size. Those of salps were, in general, rectangular, and slight differences were noted according to the species. The fecal pellets of the molluscan pteropod Clio sp. were conical, while those of the molluscan heteropod Carinaria sp. were spiral. The sinking velocities ranged from 26.5 to 159.5 m day^-1 for copepod fecal pellets, from 16.1 to 341.1 m day^-1 for euphausiid pellets, from 43.5 to 1167.6 m day^-1 for salps' pellets (Cyclosalpa affinis, Salpa fusiformis, Iasis zonaria, and two unidentified species), from 65 to 205.7 m day^-1 for Clio sp. pellets, and from 120.3 to 646.4 m day^-1 for Carinaria sp. fecal pellets. The measured sinking velocities were compared with estimates predicted using the equations of Komar et al. (1981; Limnol Oceanogr 26:172-180), Stokes' law, and Newton's second law, using either a constant density of fecal pellets (1.22 g cm^-3) or densities estimated with the three different equations. When a constant density was used, the three equations overestimated the sinking velocities; Stokes' law resulted in the largest overestimation, and Newton's second law, the smallest. At the taxa level, the overestimation was greatest for euphausiid 1 fecal pellets and smallest for copepod fecal pellets. When the three equations were used to estimate fecal pellet density, the density estimated using the equation of Komar et al. was the greatest, and that using Stokes' law, the smallest, resulting in over- and underestimation of sinking velocities, respectively. Newton's second law resulted in an intermediate density and gave the closest estimate of sinking velocities. We propose that measurement of sinking velocities of a portion of the fecal pellets might guide in choosing an appropriate equation to be used for a reasonable interpretation of vertical mass flux.     

Resting eggs of zooplankton (Copepoda and Cladocera) from the Kiel Bay and adjacent waters (southwestern Baltic)

Resting eggs of four species of calanoid copepods and three species of cladocerans were collected from sediments up to 5cm depth from the Kiel Bay and adjacent waters in the southwestern Baltic Sea during April–May 1994. All but one species of cladoceran was successfully hatched/reared in the laboratory. In the Kiel Bay, egg abundances varied from 1.8x10⁵ to 7.4x10⁵ m^-2. Hatching success of copepod eggs collected from all depths was high (49 to 94%), but was 0 to 79% for cladoceran eggs. Darkness did not seem to affect hatching. Eggs found in the 4 to 5 cm layer of sediment were estimated to be about 15 yr old, showing the presence of an egg bank in the Baltic. Formation of resting eggs may be a genetic trait acquired during the ice ages.     

Ecological significance of salp fecal pellets collected by sediment traps in the eastern North Pacific

Five sediment traps were moored at depths of 740, 940, 1 440, 3 440 and 4 240 m for 7 d in December 1982 at Station 5 in the eastern North Pacific about 400 km from San Francisco. Dark green sinking particles enclosed in tough membrane consisted of mostly coccolithophores with some diatoms, dinoflagellates and chrysophytes. The average size of the particles was 10x5x2 mm. These characteristics indicate that the particles were fecal pellets of salp inhabiting the surface waters. Vertical fluxes of the organic carbon and nitrogen through sinking of the salp fecal pellets ranged from 6.7 to 23 mgC m^-2 d^-1 and from 0.88 to 3.2 mgN m^-2 d^-1, respectively. These values were several times higher than those determined in other oceanic areas by sediment trap experiments. Hydrocarbons consisting of short-chain n-alkanes (n-C₁₅-C₂₀) with n-C₁₇, the most predominant component, heneicosa-hexaene (n-C_21:6), br-C₂₅ alkenes and long-chain n-alkanes (n-C₂₁-C₃₀), without any odd or even carbon number predominance, were found from five depths. The presence of short-chain n-alkanes and n-C_21:6 indicated that phytoplankton in the surface waters was a primary source of organic matter in the sinking particles. Two isomers of br-C_25:3 and br-C_25:4 alkenes found in these particles also indicated that br-C₂₅ alkenes were the important biological marker of fecal pellet of zooplankton. The distribution pattern of the long-chain n-alkanes suggested that the sinking particles may be affected by bacteria to some extent. Fatty acids of the sinking particles were separated into free, triglyceride and wax ester fractions consisting of mono- and poly-unsaturated, and saturated fatty acids, with a range from C₁₄ to C₃₀. Concentrations of mono- and poly-unsaturated fatty acids decreased more rapidly toward the deep than those of saturated fatty acids, which cause low ratios of mono- and poly-unsaturated fatty acids/saturated fatty acids. This indicates that unsaturated fatty acids were more rapidly decayed by marine microbes than saturated fatty acids in the deep water, despite the fact that a significant amount of unsaturated fatty acids still remained in the sinking particles collected from the deep waters. Our results revealed that the salp fecal pellet plays an important role in supplying foods to organisms in intermediate and deep seas.     

Detection of elements and radioactivity in pellets from long-eared owls (Asio otus) inhabiting the city of Belgrade (Serbia)

In this study, we analysed pellets from long-eared owls (Asio otus) collected from four localities in Belgrade (Serbia). The pellets contained the remains of prey, namely voles (Arvicola terrestris) and field mice (Apodemus agrarius). The concentrations of 14 elements (Ca, P, Mg, Na, K, Fe, Zn, Sr, Ba, Mn, Ti, Cu, Si, B) were evaluated in whole pellets and in samples containing only bone tissue, which were dissected from the whole pellet. The increased levels of certain elements, including Mn, Zn, Ba, Cu and radioactive ⁴⁰K, indicate contamination of the soil by various sources, such as industrial plants and agricultural practices. From the results presented in this article, we suggest that the analysis of owl pellets may indicate the quality of the local environment.     

Ecological implications of fecal pellets produced by the Antarctic krill Euphausia superba in the Antarctic Ocean

Organic analyses and electron microscopic observations on fecal pellets produced by the Antarctic krill Euphausia superba Dana showed that krill fed on choanoflagellates, the abundant heterotrophic flagellate in the Antarctic Ocean. Two new pathways of organic materials in the Antarctic ecosystem are proposed: (1) a new food chain including non-living particulate and dissolved organics, and bacteria-choanoflagellate-krill-vertebrate, which coexists with the traditional diatom-krill-vertebrate food chain; (2) non-phytoplanktonic organic materials in surface waters are transferred into choanoflagellates and are transported to deep water as fecal pellets which are still useful as nutrition for other organisms there.     

Metabolic activity and elemental composition of krill and other zooplankton from Prydz Bay,Antarctica, during early summer (November–December)

Oxygen uptake and ammonia excretion rates, and body carbon and nitrogen contents were measured in krill (Euphausia superba) and eight other zooplankton species collected during November–December 1982 in the Prydz Bay, Antarctica. From these data, metabolic O:N ratios (by atoms), body C:N ratios (by weight) and daily metabolic losses of body carbon and nitrogen were calculated as a basis from which to evaluate seasonal differences in metabolism and nutritional condition. Comparison of the present data with mid-summer (January) data revealed that early-summer E. superba were characterized by higher metabolic O:N ratios (58.7 to 103, compared with 15.9 to 17.5 for mid-summer individuals). Higher O:N ratios of early-summer E. superba resulted largely from reduced ammonia excretion rates and, to a lesser degree, from slightly increased oxygen uptake rates. Body C:N ratios of E. superba were low in early-summer (3.8 to 4.2) compared with mid-summer krill (4.1 to 4.7) due to lowered body-carbon contents in the former (42.6 to 43.6% compared with 43.2 to 47.5% dry weight of midsummer individuals); gravid females formed an exception, since no seasonal differences in body elemental composition were detected for these. No significant changes in water content (75.3 to 81.4% wet wt) and nitrogen content (9.9 to 11.1% dry wt) in E. superba were evident between the two seasons. Seasonal differences in metabolic rates and elemental composition were less pronounced in a salp (Salpa thompsoni), but a higher metabolic O:N ratio occurred in early-summer individuals. Interspecific comparison of the seven remaining zooplankton species studied with twelve species from mid-summer zooplankton investigated in an earlier study indicated that higher metabolic O:N ratios in early-summer are characteristic not only of herbivore/filter-feeders, but also of some carnivores/omnivores. No relationship between metabolic O:N ratios and body C:N ratios was apparent either intraspecifically or interspecifically, within or between early-summer and mid-summer seasons.