Distribution and abundance of ctenophores, and their zooplankton food in the Black Sea. I. Pleurobrachia pileus

The distribution of Pleurobrachia pileus Müller, 1776 in the Black Sea was determined using plankton samples collected above the anoxic zone (maximum of 200 m) in the winter, spring, and summer of 1991 to 1995. The summer samples were collected in 1991 to 1993 (for a previous) and are included in this paper for comparative purposes. High concentrations of P. pileus were found at the northern edges of anticyclonic eddies along the southern coastal regions. The biomass and abundance of P. pileus increased from winter through spring to a peak in summer. The highest mean wet weight during a sampling period was 250 g m⁻², while the maximum wet weight was 1429 g m⁻². P. pileus was mostly found in a layer extending from the lower parts of the thermocline down to the anoxic zone, where the temperature was <8 °C. The vertical distribution of P. pileus biomass had two clear maxima at night: an upper maximum at 20 to 40 m was less pronounced than the lower maximum at 90 to 120 m depth. Mean body length of P. pileus did not exceed 12 mm. Smaller individuals (9 to 10 mm length) occurred in winter. P. pileus had two length classes in early spring (March 1995) and late summer (August 1993), indicating the presence of both newly hatched and larger individuals. Overall, the stomach contents of P. pileus consisted mainly of Copepoda (90%), Cladocera (1%), Mollusca (1%), fish eggs and larvae (1%), and other taxa (7%). The preferred food of P. pileus (frequency of occurrence) was: Calanus euxinus (39%), Pseudocalanus elongatus (30%), Acartia clausi (28%), Oithona similis (2%), and Paracalanus parvus (1%). The endoparasite Hysterothylacium aduncum was commonly found in P. pileus. Abundances of Mnemiopsis leidyi and P. pileus were either negatively correlated (r = −0.5 to −0.7) or positively correlated at a low significance level (r = 0.25 to 0.3) with abundance of A. clausi in different months of the year. Aurelia aurita abundance was correlated mainly with the abundance of C. euxinus from June 1991 to March/April 1995. Over the same period the abundance of P. pileus was significantly correlated with the abundance of P. elongatus, an important prey species. Received: 1 November 1997 / Accepted: 30 August 1999     

Distribution and abundance of moon jellyfish (Aurelia aurita) and its zooplankton food in the Black Sea

The distribution of moon jellyfish (Aurelia aurita Linnaeus, 1758) in the Black Sea was determined from plankton samples collected above the anoxic zone (maximum depth 200 m) in the summer, winter and spring during 1991–1995. Distribution was patchy. Average biomass ranged from 98 to 380 g m⁻², and abundance varied from 2 to 14 individuals m⁻². Biomass and abundance peaked in late spring and summer. The distribution was correlated with hydrographic features in the Black Sea, with higher concentrations occurring at the peripheries of anticyclonic eddies. Centers of the two main cyclonic gyres generally had a low biomass of A. aurita. From July 1992 to March 1995, the populations were largely concentrated in offshore regions. A. aurita were confined to the upper part of the mixed layer. Smaller A. aurita (≤1 cm) were present in early spring (March), and individuals reached maximum size in the summer. Release of the epyhrae occurred in spring on the northwestern shelf of the sea when the seawater temperature was 11–12 °C. Microscopic analysis of stomach contents showed that copepods and mollusks form their main diet. Received: 3 September 2000 / Accepted: 29 September 2000     

Shifting abundance of the ctenophore Mnemiopsis leidyi and the implications for larval bivalve mortality

Along the mid-Atlantic coast of the US, the ctenophore Mnemiopsis leidyi (Agassiz) appears to be increasing in abundance and undergoing shifts in its historical seasonal distribution. We provide new data on shifting ctenophore abundance in Long Island estuaries and its implications for top-down control of the plankton community. Peak mean biovolume estimates of M. leidyi in Long Island estuaries in 2006 revealed ctenophore abundance values that were a factor of two to five times greater than previous studies conducted two decades ago. Furthermore, peak M. leidyi densities in 2006 occurred 2–3 months earlier than previously documented, suggesting a shift in the seasonal maxima of M. leidyi. Application of daily ingestion rates to zooplankton abundance indicates that, at its highest densities M. leidyi can remove an overall average of 20–89% per day of bivalve veligers and other zooplankton taxa, including adult copepods, nauplii, and tintinnids. Increasing ctenophore abundance, especially during a period when they were not historically abundant (i.e., June) may have significant consequences for species which spawn at this time. For example, current populations of M. leidyi represent a major source of larval mortality for bivalves which may inhibit recovery of shellfish populations and reinforce their low abundance state in Long Island estuaries.     

Population development of the invader ctenophore Mnemiopsis leidyi, in the Black Sea and in other seas of the Mediterranean basin

In the last two decades of the twentieth century, the ctenophore Mnemiopsis leidyi (A. Agassiz) has invaded the Black, Azov, Marmara and Aegean Seas, and, recently, the Caspian Sea. Here, we compare its spatial and temporal distribution, seasonal dynamics and the time and duration of reproduction. We also discuss factors that control its abundance throughout its invasive range and its effect on ecosystems. Observations are based on the long-term field data collected by three research institutes. An analysis of the effects of temperature, salinity, prey (zoo- and ichthyoplankton) availability and predation (by ctenophores of the genus Beroe) on M. leidyi population size, and the effects of M. leidyi on zoo- and ichthyoplankton, and on fish populations in the Black and Azov Seas is also provided. With the Black Sea current, M. leidyi spreads to the upper layers of the Sea of Marmara, where it now occurs around the year. At regular intervals, the Black Sea current also takes it to the northern Aegean Sea. In contrast, it has to re-invade the Sea of Azov every spring or summer, dying out during winter when the temperature drops below 4°C. The warm summer and mild winter temperatures, relatively low salinity and abundance of prey in the Black Sea are close to optimal for M. leidyi, while they are suboptimal in the northern Aegean Sea, where salinity and temperature are often too high. In the Black Sea the absence of gelatinous and other predators led to an enormous ctenophore abundance for a decade, but with the appearance of Beroe ovata in 1999, M. leidyi abundance greatly decreased. Analysis of seasonal dynamics of M. leidyi in the Black Sea and in other seas of the Mediterranean basin indicates similarities in the timing of maximum abundance and biomass, in spite of some differences in the initiation and duration of reproduction. A peak biomass and density occurred in 1989 in the Black and Azov Seas and in 1990 in the other seas. The M. leidyi invasion negatively affected the ecosystems of the Black Sea and the Sea of Azov. The zooplankton, ichthyoplankton and zooplanktivorous fish stocks all underwent profound changes. Similar effects, but less pronounced, were recorded in the Sea of Marmara. Effects on Mediterranean food chains have, so far, remained insignificant. Salinity is probably supraoptimal here, and several predators prevent M. leidyi from reaching outbreak levels.     

Escape of the ctenophore Mnemiopsis leidyi from the scyphomedusa predator Chrysaora quinquecirrha

The ctenophore Mnemiopsis leidyi A. Agassiz, 1865 is known to be eaten by the scyphomedusan Chrysaora quinquecirrha (Desor, 1948), which can control populations of ctenophores in the tributaries of Chesapeake Bay. In the summer of 1995, we videotaped interactions in large aquaria in order to determine whether M. leidyi was always captured after contact with medusae. Surprisingly, M. leidyi escaped in 97.2% of 143 contacts. The ctenophores increased swimming speed by an average of 300% immediately after contact with tentacles and 600% by mid-escape. When caught in the tentacles of C. quinquecirrha, the ctenophores frequently lost a portion of their body, which allowed them to escape. Lost parts regenerated within a few days. The striking ability of M. leidyi to escape from C. quinquecirrha may be critically important in maintaining ctenophore populations in situ. Received: 14 November 1996 / Accepted: 4 December 1996     

Distribution and abundance of finless porpoise in the Inland Sea of Japan

Aerial sighting surveys were conducted in 2000 to evaluate the distribution and abundance of finless porpoise Neophocaena phocaenoides in the Inland Sea of Japan. We flew 60 north–south transects (2,218 km in total) at intervals of 6.43 km over the study area between 131°3′ and 134°59′ E. In total, 148 groups were detected by two observers. The average group size was 1.56 individuals. The effective strip half-width of each observer was estimated to be 107 m (coefficient of variation [CV] = 8.26%). Porpoise abundance was estimated at 7,572 individuals and the density was as low as 0.506 individuals/km² (CV = 17.3%). The sea is among the Japanese waters with the lowest density in spite of its favorable topographical conditions. In the western stratum of the study area, where the estimated density was 1.31 individuals/km², we observed a relatively regular distribution. In the central-eastern stratum, which had an estimated density of 0.208 individuals/km², we confirmed a clumped distribution that was restricted to inshore waters or near islands. No individuals were observed in waters between 132°51′ and 133°11′E, and between 133°43′ and 133°59′E, where sand dredging and other human activities have been active, suggesting that habitat fragmentation has occurred.     

Feeding habits and food requirements of some amphipods in the Black Sea

Food composition and food requirements of four amphipod species of the Black Sea — Dexamine spinosa (Mont.), Amphithoe vaillanti Lucas, Gammarellus carinatus (Rathke), Gammarus locusta L. — were studied and an attempt made to assess the quantitative composition of the food, daily rhythms of feeding, and daily food rations. The composition of the food of the above-named amphipods is similar and consists primarily of seaweeds. Daily feeding intensity is characterized by a distinct increase during the dark period (D. spinosa, A. vaillanti); however, in G. locusta feeding intensity increases in the morning. The daily food rations of amphipods vary greatly (from 1.7 to 360% of the specimens' body weight), depending upon factors such as the kind of food, age, sex, the physiology of specimens, and the water temperature. The total quantity of food eaten by the populations of these amphipods in the coastal zone of the Black Sea amounts to 4000 g organic matter per square metre within 1 year.     

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     

Distribution and bioaccumulation of organochlorine pesticides along the Black Sea coast

Sediment, mussel, and seawater samples were collected three times during 2001–2003 at nine sampling stations along the mid-Black Sea coast of Turkey. The samples were analyzed with GC-ECD for contents of various organochlorine pesticides (OCPs) in the environment. DDT and its metabolites were detected at concentrations significantly above the detection limits. The highest concentrations of DDT metabolites measured in the sediment and mussel samples were 35.9 and 14.0 ng/g wet weight respectively. Considerable levels of aldrin, dieldrin, endrin, heptachlor epoxide, lindane, endosulfan sulphate, and HCB were also detected in the sediment, mussel, or seawater samples. Although these persistent toxic compounds have been banned for some years in Turkey, they may still be used illegally in some regions, contributing to their significant levels in the environment. The biota–sediment accumulation factor (BSAF) estimated for DDT and its metabolites in mussels was 2.9, which is nearly two times higher than the benchmark of 1.7. In spite of such high BSAF values observed for these toxic compounds, their levels in mussels were significantly below the international legal limits recommended by the Food and Agriculture Organization of the United Nations. Edible biota from the waterbodies examined may thus still be considered safe for human consumption at this time. However, as pollutants can biomagnify through the food chain over time, further routine sampling and analysis of biota along the Black Sea coast are warranted in order to better assess the threat of OCPs to public health in the region.     

Distribution and ecology of the pelagic hydromedusae,siphonophores and ctenophores of the Barents Sea,based on perennial plankton collections

The evaluation of more than 10,000 samples of plankton, taken at standard stations from 1953 to 1960 in the Barents Sea, made it possible to supplement considerably the species list of medusae, siphonophores, and ctenophores, and to revise some ecological and biogeographical characteristics of many of them. Quantitative data, assembled over many years, on the occurrence of the above-mentioned groups, demonstrated the impossibility of using the vast majority of representatives of this fauna as biological indicators. During the very warm year of 1960, reproductively and vegetatively cold-adapted forms produced the same population increase as the thermophilic forms. The change in thermal conditions influenced these animals indirectly via a favourable biotic situation due to a prolongation of the nutrition period. The capability to vary the number of eggs, timing, rates and types of reproduction, to withstand long periods of starvation, etc., facilitates a flexible synchronization of the population dynamics of these pelagic predators with the quantity of plankton used for nutrition. The same properties stipulate the possibility of seasonal and annual meandering in regard to the boundaries of the water areas occupied.     

Hydrolytic enzymes in the two North Sea ctenophores Pleurobrachia pileus and Beroe gracilis

Hydrolytic enzyme activities were determined in tissue extracts of the two North Sea ctenophores Pleurobrachia pileus and Beroe gracilis. The following activities were measured in field samples and characterized according to their mean specific activities and their pH optima: protease, aminopeptidase, amylase, N-acetylglucosaminidase, -glucosidase, chitinase, carboxylesterase, phosphatase. Many of the enzyme activities are correlated, indicating their control by the same factors (i.e. nutritional conditions). Differences found between the phosphatase activities and the apparent lack of the chitinolytic system in B. gracilis reflect the differences in body composition and food specialization in both ctenophores, respectively. The low amylase and -glucosidase activities in both ctenophores agree with the low carbohydrate content of their prey. In spite of high carboxylesterase activities, no lipolytic activity was detected in either of the ctenophores. In a laboratory feeding experiment with P. pileus, the enzyme activities increased but did not reach the activities found in the field samples. In B. gracilis, seasonal variations of some enzyme activities are discussed in relation to the seasonal abundance of its prey, P. pileus.     

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.     

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.     

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.     

Distribution and abundance of ophiuroids on the continental shelf and slope of the East Sea (southwestern Sea of Japan), Korea

The present study was carried out to describe the distribution of ophiuroids on the continental shelf and slope of the East Sea, off the Korean Peninsula. Three cruises were made from 1985 to 1987 in the area between 35 to 38°N and 129 to 130°E. A total of ten species was identified from the samples collected at 136 stations. Among them four species (Amphiodia craterodometa, Amphioplus macraspis, Ophiura leptoctenia and Ophiura sarsi) constituted more than 98% of abundance by number. The continental shelf off Gampo, near 35°40N, could be considered to be the southern limit of cold-water species. The abundance of dominant ophiuroids changed according to depth. Three faunal zones, namely, the inner shelf, the shelf edge and the slope zones, coincided with the topographic divisions of continental shelf and slope. The major inhabitant of the inner shelf was A. craterodometa and that of the continental slope was O. leptoctenia. The shelf edge area revealed a transitional zone where all four species occurred, contributing to the total number of individuals with almost equal proportions. The temperature of bottom waters was considered to be a factor determining zonal distribution. The sediment characteristics were not closely related to ophiuroid distribution.     

Temporal variability of deep-sea zooplankton in the Arabian Sea

Mesozooplankton samples from two stations in the Arabian Sea (WAST, 4,050 m, 16°15′N, 60°20′E; CAST, 3,950 m, 14°30′N, 64°30′E) were collected from the surface down to 20 m above bottom during three monsoon periods: the autumn intermonsoon in October 1995, the spring intermonsoon in April 1997, and the NE monsoon in February 1998. The main goal of this study is to enhance our knowledge on the effect of spatial and temporal differences in primary production and particle flux rates on the abundance and distribution of mesozooplankton, with special attention to the deep sea. Literature data indicate episodically high rates of primary production and particle flux in the region during the SW monsoon and the autumn intermonsoon. Set in this context, the zooplankton showed an in-phase coupling in biomass and abundance with the primary production in the surface 150 m. In the mesopelagic realm (150–1,050 m), the seasonal coupling was less clear. In the bathypelagic zone, below 1,050 m, there was no evidence of in-phase coupling, though temporal differences in the distribution of zooplankton abundance and biomass with depth between seasons could be shown by an analysis of covariance and an a posteriori test. The results suggest that the bathypelagic community responds to increased particle flux rates, but with longer time gaps than in the epipelagic zone. This is probably due to longer development and response times of zooplankton in the cold, deep-water environment, independent of possible lateral advection processes.     

Distribution and abundance of choanoflagellates (Acanthoecidae) across the ice-edge zone in the Weddell Sea,Antarctica

Choanoflagellates are thought to be an important component of oceanic microbial food webs, but little quantitative data exists on their abundance,, distribution, or relationship to potential food sources. In an Antarctic ice edge zone (northern Weddell Sea, March 1986), choanoflagellate abundance varied over two orders of magnitude in the upper 100 m. The lowest abundances were recorded at the bottom of the water column under ice cover and the highest abundances occurred in the upper 30 m of open water. Species that were predominantly in colonies dominated the open-water samples. Abundances of total choanoflagellates and some individual species were correlated with primary and secondary biomass and production, indicating a response to gradients in potential food sources. This suggests that choanoflagellates are tightly coupled with their food sources and supports the contention that they may an important link between bacteria-sized particles and metazoan grazers.