The cladocerans of the Aegean Sea: Occurrence and seasonal variation

The little-known cladoceran population of the Aegean Sea was studied. One hundred and seventy-six planktonic samples were collected during 8 cruises in the Aegean Sea (4 in the northern part, 4 in the southern part). From these samples, the following 4 species of Cladocera were determined: Evadne spinifera Muller, Evadne tergestina Claus, Penilia avirostris Dana, Podon intermedius Lilljeborg. All species were found both in the north and in the south Aegean Sea, with the exception of P. avirostris, which was found only in the northern Aegean Sea. Its absence from the southern part is attributed to the higher salinities and the pelagic character of the area. With regard to abundance, E. spinifera is by far the commonest in this area; P. avirostris and E. tergestina could be characterised as frequent forms; P. intermedius is scarce. P. avirostris, E. spinifera and E. tergestina are thermophilic; their occurrence is more or less restricted to the warm season; P. intermedius is psychrophilic and appears only in winter.     

Distribution of marine cladocerans in the Indian Ocean

Cladocera of the Indian Ocean have been studied, on the basis of samples collected during the International Indian Ocean Expedition (IIOE). Out of a total of 1927 samples, 552 contained Cladocera. Although the distribution of this group is mainly coastal, one species, Evadne tergestina, has been found in oceanic waters also, where it may establish seasonal populations. Evadne spinifera was recorded off the Australian and South African coasts in the southern hemisphere, and in the Northeastern part of the Arabian Sea coastal waters in the northern hemisphere. Podon polyphemoides was reported from a single station in South African waters. Penilia avirostris was found in coastal waters throughout the Indian Ocean. Short-term distribution patterns of E. tergestina and P. avirostris are discussed. The possible value of this group as food for pelagic fishes has been indicated. This work makes clear that adequate sampling in space and time is required along the Continental Shelf of the Indian Ocean in order to obtain more comprehensive information on this group.Visiting Scientist at Indian Ocean Biological Centre, Cochin, India under UNESCO contract SC/2077/70-20579.     

Seasonal contrasts in abundance and reproductive parameters of Penilia avirostris (Cladocera, Ctenopoda) in a coastal subtropical area

We studied the population dynamics and the reproductive biology of Penilia avirostris during three consecutive years on the inner shelf off Ubatuba, Brazil. Penilia avirostris individuals and its eggs and embryos were counted, measured, and classified into stages. The species occurred throughout the studied period, in a wide temperature range (14.8–28.2°C). Cladoceran densities were usually higher (>2,000 ind m⁻³) in warm seasons, when the water column was stratified as a consequence of bottom intrusions of the cold- and nutrient-rich South Atlantic Central Water. Juveniles, non-reproducing females, and parthenogenic females were the dominant developmental stages. Males and gamogenic females were rare and only occurred when females reached peak abundances. This suggests that in tropical and subtropical coastal seas gamogenesis in P. avirostris is not as common as in temperate seas, but may play a significant role in the density-dependent control of the population preceding unfavourable periods.     

Longevity of subitaneous and diapause eggs of Centropages hamatus (Copepoda: Calanoida) from the northern Gulf of Mexico

Copepod resting eggs are abundant in the seabed of many bays and estuaries where they provide a potential source of recruits for growth of planktonic populations. In the northeastern Gulf of Mexico the copepod Centropages hamatus (Lillejeborg) occurs in the water column only during the late fall, winter and early spring. The species produces subitaneous and diapause eggs, and both egg types have been found in the seabed. We determined the longevity of these two egg types to ascertain their potential for contributing to the growth of the planktonic population and for sustaining a persistent egg bank. Eggs were collected from females and incubated in the laboratory under temperature and oxygen conditions chosen to simulate field conditions. The diapause eggs were also exposed to sulfide. The total hatching success of subitaneous eggs in two experiments declined from highs of 78 and 97% to zero after 60 and 90 d of exposure to anoxia. The total hatching success of diapause eggs that were exposed to anoxia for 90 d however was typically greater than 80%. Some diapause eggs hatched after being incubated under anoxia for 437 d. Diapause eggs survived longer at ambient field temperatures when incubated under anoxia (437 d) compared to normoxia (118 d). Exposure to sulfide did not result in greater mortality of diapause eggs compared to anoxia alone. Diapause eggs that were incubated at ambient field temperatures did not hatch when exposed to normoxia until the temperature dropped to <20 °C. The results of this study suggest that C.␣hamatus sustain a short-term reserve of subitaneous eggs in the seabed that provides recruits for the current year's population. The greater longevity of diapause eggs suggests that they sustain the seasonal reappearance of the species year after year in the northeastern Gulf of Mexico. However, the contribution of diapause eggs of C. hamatus from the Gulf of Mexico to a persistent egg bank is questionable since hatching ceased after 437 d. Received: 30 July 1997 / Accepted: 18 January 1998     

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.     

Are direct developers more locally adapted than planktonic developers?

The hypothesis that populations of direct developers exhibit greater geographic differentiation in life history features than populations of planktonic developers, was tested with two species of grazing snails of the genus Littorina from 1986 to 1987. Littorina sitkana (direct developer) and L. scutulata (planktonic developer) coexist on sun- and wave-sheltered beaches from Alaska to Oregon, USA. Seasonal patterns in growth, survival and reproduction were monitored for samples from four geographically separated populations of each species grown in population cages at a common site, Friday Harbor, Washington, USA. The environmental and population effects on growth in the two species were determined in a four-way reciprocal transplant experiment with the same populations. Both the direct and planktonic developers exhibited geographic differentiation in life history features. Differentiation in the direct developer occurred over distances shorter than 30 km, while differentiation in the planktonic developer occurred over the 500 km distance examined (greater than their larvae would likely travel).     

Mesozooplankton resting eggs in the Baltic Sea: identification and vertical distribution in laminated and mixed sediments

Benthic resting eggs of rotifers, cladocerans and calanoid copepods were studied at two sites with different hydrographical and sediment properties off the SW coast of Finland, the northern Baltic Sea between 1991 and 1993. Vertical distribution of the resting eggs in the sediment was studied down to 10 cm depth, and hatching of the eggs extracted from different 1-cm thick sediment layers was experimentally tested. The larvae of eight mesozooplankton taxa emerged from the eggs: Calanoida (mostlyAcartia bifilosa), cladoceransBosmina spp.,Daphnia spp.,Evadne nordmanni, andPodon spp. (mostlyP. polyphemoides), and rotifersSynchaeta spp.,Keratella spp. andAsplanchna priodonta. Some calanoid and cladoceran eggs hatched after 7 mo storage of the sediment at 3°C. A few calanoid and cladoceran eggs hatched from the 8 to 10 cm deep sediment layer. Their age, estimated by means of¹³⁷Cs analysis, was ca. 8 to 10 yr. Vertical distribution of the benthic eggs reflected the different sediment structures of the study sites. The eggs of all zooplankton taxa were more stratified in the laminated sediment than in the disturbed sediment. We suggest that the intensity of deep water mixing is the key factor for regulating the vertical distribution and hatching probability of mesozooplankton resting eggs, since it largely determines the stratification of the sedimenting material, distribution of oxygen within the sediment, and the living conditions of benthic animals causing bioturbation.     

Calanoid copepod eggs in sea-bottom muds. V. Seasonal changes in hatching of subitaneous and diapause eggs of Tortanus forcipatus

A warm-water neritic calanoid copepod, Tortanus forcipatus, has been found to lay two physiologically different types of eggs (subitaneous and diapause eggs), which are separately spawned in response to different seasonal environmental conditions. Subitaneous eggs are produced when the planktonic populations are at their numerical maxima, and are only shed until late summer. After peak spawning, true resting eggs are spawned which undergo diapause for 1 to 3 months. The eggs recover from their diapause around mid-winter, but hatching does not take place until the water temperature at the sea bottom exceeds 15°C in early summer.     

Effects of various environmental factors on egg development of several species of Acartia in Southern California

The hatching of the eggs of 3 Acartia species (A. tonsa, A. clausi, and Acartia sp. I) spawned in the laboratory and the eggs of Acartia (species undetermined) isolated from natural bottom sediments was examined under various environmental conditions (temperature, salinity and temperature combinations, oxygen concentration, illumination, and the presence of bottom mud). The range of temperature and salinity in which eggs hatched differed from one species to another. Egg hatching was inhibited under certain environmental conditions for all species, although egg viability under such conditions varied from one species to another. In relation to these findings, habitat separation of planktonic Acartia populations and the role of dormant eggs during their seasonal absence in the plankton of Southern California coastal waters are discussed.     

Uncommon diversity in developmental mode and larval form in the genus <Emphasis Type="Italic">Macrophiothrix</Emphasis> (Echinodermata: Ophiuroidea)

Development mode in the ophiuroid genus Macrophiothrix includes an unusual diversity of planktonic larval forms and feeding types. The modes of development for seven congeners that coexist in coral reef habitats at Lizard Island, Australia were compared using larvae generated from crosses over several reproductive seasons from 1999 to 2003. Three species (Macrophiothrix koehleri Clark, Macrophiothrix longipeda Lamarck, Macrophiothrix lorioli Clark) develop from small eggs (<170 μm) into typical obligately feeding planktonic (planktotrophic) pluteus larvae with four larval arm pairs. The remaining four species develop from larger eggs (≥230 μm) into either facultatively-feeding or non-feeding (lecithotrophic) larval forms. The facultative planktotroph (Macrophiothrix rhabdota Clark) retains the ability to digest and benefit from food but does not require particulate food to complete metamorphosis. Among the lecithotrophic species, Macrophiothrix caenosa Hoggett retains the pluteus morphology with four pairs of larval arms, but is incapable of feeding, depending instead on maternal provisions for larval development. The remaining two lecithotrophs have simplified larval morphologies with only a single pair of full length (Macrophiothrix nereidina Lamarck) or highly reduced (Macrophiothrix belli Doderlein) larval arms and no functional mouth or gut. This genus includes the first example of facultative planktotrophy in ophiuroids, the first example in echinoderms of a complete pluteus morphology retained by a lecithotrophic larva, and three degrees of morphological simplification among lecithotrophic larval forms. Egg volume varies 20-fold among species and is related to variation in feeding mode, larval form, and development time, as predicted for the transition from planktotrophic to lecithotrophic development.     

Subitaneous, diapause, and delayed-hatching eggs of planktonic copepods from the northern Gulf of Mexico: morphology and hatching success

Experiments were conducted to examine the morphology and hatching success of eggs, either spawned by freshly caught planktonic copepods or recovered from bottom sediments in the northern Gulf of Mexico. Collections were made between August 1992 and September 1995. Eggs of nine species were described and these differed in their diameter, color and surface attributes. Three types of eggs were distinguished: subitaneous, diapause, and delayed-hatching. Three species, Labidocera aestiva Wheeler, Acartia tonsa Dana, and Centropages velificatus (Oliveira) produced only subitaneous eggs. Hatching success varied greatly among these species. Two species, Labidocera mirabilis Fleminger and Centropages hamatus (Lilljeborg) produced diapause eggs and subitaneous eggs. The length of the refractory phase of the diapause eggs differed greatly both within and between these two species. A third type of dormant egg, delayed-hatching, was recognized in Labidocera scotti Giesbrecht and Pontella meadi Wheeler. The existence of delayed-hatching eggs may be an adaptive response of subtropical species to less seasonal fluctuation. Based upon morphological characteristics of the eggs and rearing of nauplii to an identifiable stage, benthic dormant eggs of eight species of calanoid copepods were also identified. Received: 9 September 1996 / Accepted: 1 October 1996     

Seasonal study of planktonic copepods and their benthic resting eggs in northern California coastal waters

Few investigations have examined the occurrence of zooplankton resting eggs in the sea bed of waters deeper than 20 m. In this study the distribution and abundance of planktonic copepods and their benthic resting eggs in coastal waters off northern California, U.S.A., were determined and related to environmental parameters (temperature, salinity, depth, and sediment grain size). Sediment cores, net tows, and CTD profiles were obtained in April and October 1989, and February, April, and October 1990. Water depths in the study area ranged from approximately 60 to 120 m. The mean abundance of eggs was as high as 1.2×10⁵ m^-2 for Acartia clausi Giesbrecht and 1.9×10⁵ m^-2 for Tortanus discaudatus Thompson and Scott. These egg concentrations are comparable to those reported previously for shallower more protected regions. The abundance of eggs in the sediments decreased with increasing depth of the water column. For the region as a whole, eggs were least abundant in muddy sediments. The mean abundance of eggs in the sea bed also varied seasonally and annully. Benthic resting eggs of A. clausi were more abundant in April 1989 than in April 1990, and adults of the species were never found in the plankton samples. The lack of adults is not unusual since results of previous studies indicate that A. clausi is a cold-water species, and in this region water temperatures are colder in summer, than in winter, due to upwelling. Temperature and salinity data indicated that the upwelling season had commenced by the time of the April 1990, but not the April 1989 sampling. Thus, the reduced abundance of benthic eggs in April 1990 may have been due to egg-hatching in response to reduced temperatures. The results suggest that the presence of A. clausi in coastal waters off northern California is linked to recruitment from benthic resting eggs.     

Remarques sur la structuration des populations zooplanctoniques dans la zone de l'émissaire de Marseille-Cortiou

The impact of a polluted water-layer on zooplanktonic populations was studied in the sewage discharge zone at Marseille-Cortiou throughout 1977 and a modification of specific diversity was observed: diversity was instable near the outfall, but increased slightly and became more stable further away in non-polluted stations. The population composition did not seem to change. Species rank varied slightly and the number of species increased with increasing distance from the outfall. Specific dominance remained relatively unchanged, and essentially consisted of only 5 species: Clausocalanus acuicornis, Evadne spinifera, Oithona helgolandica, Temora stylifera and Centropages typicus. The population structure, as illustrated by Frontier diagrams, was higher during or just preceding winter. Population organisation was the same close to or far away from the outfall, although the total population counts and number of species were smaller at the most polluted stations. The taxons exhibiting the largest decreases in density were: euphausids, nauplii, larvae of various brachyurans and gastropods, fish eggs, the pteropod Cavolinia inflexa, the copepod Calanus minor and several Coryceidae. The greatest disturbance of the population structure at all stations was observed during the summer. The composition of the zooplanktonic population seems to depend much more on seasonal factors than on the level of pollution. In was not possible to distinguish a cluster of species which were sufficiently dominant to be considered as really characteristic; perhaps the rather high exposure of the sewage site to winds and waves prevents the formation of a local facies.     

Reproduction of Halicarcinus planatus (crustacea, decapoda, hymenosomatidae) in the Deseado River estuary, southwestern Atlantic Ocean

The small spider crab Halicarcinus planatus, is a common resident in the rocky coasts of the Magellanic region in South America. The H. planatus population in the Deseado River estuary differs from other populations of the same species in the spatial segregation of sexes. Between March 2002 and October 2003, samples were taken monthly from the lower intertidal and upper subtidal levels. All crabs captured were females. Additional samplings were carried out at the subtidal level to search for the presence of males. A total of 3,346 females were captured and their relative age was determined. Some of these females were maintained in the laboratory to study the terminal moult and the spawning seasons. The analysis of the ovary development of adolescents (ADO) and mature (MAT) females, and the times when females of different ages appear, when spermatophores are found inside spermathecae, and when post-spawning mortality takes place allowed us to establish the whole reproductive process. Adolescents are impregnated only once before the terminal moult. There are seven or eight spawnings after the moult, with a brief resting period between two successive reproductive seasons. The life span of mature females ranges between 15 and 22 months. Results indicate that the H. planatus population of the Deseado River estuary differs greatly from that of the Kerguelen Islands.     

Overwintering strategies of dominant calanoid copepods in the German Bight, southern North Sea

Abundance, stage composition and reproductive parameters (egg production, egg viability, proportion of spawning females) of the four copepod species Acartia clausi, Centropages hamatus, C. typicus and Temora longicornis were measured at the long term sampling station Helgoland Roads (German Bight, southern North Sea) from September 2003 to May 2004 to study their overwintering strategies. A. clausi was overwintering as females with arrested reproduction from November to January. T. longicornis, which is known to produce resting eggs in the North Sea, had a pelagic population with all developmental stages present during winter and reproductive rates closely related to food concentrations. Although their females produced eggs in response to ambient food conditions, both C. hamatus and C. typicus were rare in the pelagic. The C. hamatus population returned in May, probably from resting eggs, whereas C. typicus depended on advection. The Centropages species seemed to be less adapted to pelagic life in winter than A. clausi and T. longicornis. Sporadic occurrence of large numbers of nauplii and young copepodids of A. clausi and Centropages spp. pointed to different overwintering strategies or more successful survival in adjacent regions and advection of them into the waters around Helgoland island. While A. clausi was decoupled from environmental conditions in late autumn and winter, the other species were able to respond to variations in the food environment. Thus, egg production of T. longicornis increased during an unusual autumn diatom bloom.     

Genetic consequences of long larval life in the starfish Linckia laevigata (Echinodermata: Asteroidea) on the Great Barrier Reef

Gene flow between populations of the asteroid Linckia laevigata (Linnaeus) was investigated by examining over 1000 individuals collected from ten reefs throughout the Great Barrier Reef (GBR), Australia, for genetic variation at seven polymorphic enzyme loci. Despite geographic separations in excess of 1000 km, Nei's unbiased genetic distance (0 to 0.003) and standardised genetic variation between populations (F _ST) values (mean 0.0011) were small and not significant. Genetic homogeneity among L. laevigata populations is consistent with the long-distance dispersal capability of its 28 d planktonic larval phase, and is greater than that observed for other asteroid species, including another high-dispersal species, Acanthaster planci, which has a 14 d larval phase. Variation within populations was also higher than previously recorded for asteroids (mean heterozygosity=0.384; number of alleles per locus ranged from 5.1 to 6.0 in each population). Among asteroids, dispersal ability is positively correlated with gene flow and levels of variation, and negatively correlated with levels of differentiation.     

Comparative population genetic structure of marine gastropods (Littorina spp.) with and without pelagic larval dispersal

 Population genetic theory predicts that marine animal species with planktonic larvae will have less genetic structure than those with direct development. We compared the genetic structure of four species of littorinid snails – two with planktonic egg capsules that hatch as planktonic larvae and two with benthic egg masses that hatch as crawl-away juveniles. We used DNA sequencing and single stranded conformational polymorphism (SSCP) to assess sequence variation in a 480 bp fragment of the mitochondrial cytochrome b gene and then used an analysis of molecular variance (AMOVA) to estimate Φ_st for populations from the northeastern Pacific coast. One of the two direct-developing species, Littorina subrotundata, had a moderate amount of population structure (Φ_st=0.209) as expected but the other direct-developing species, L. sitkana, was nearly fixed for a single haplotype that made it impossible to precisely estimate Φ_st. One of the two planktonic-developing species, L. scutulata, did not show any significant population structure (Φ_st=0.004). In contrast to our expectations, the other planktonic-developing species, L. plena, showed some weak but statistically significant population structure (Φ_st=0.052). We discuss how differences in population genetic structure between species with the same type of development may reflect differences in their historical demography. Received: 22 December 1999 / Accepted: 24 July 2000     

Feeding selectivities of the marine cladocerans<Emphasis Type="Italic"> Penilia avirostris</Emphasis>,<Emphasis Type="Italic"> Podon intermedius</Emphasis> and<Emphasis Type="Italic"> Evadne nordmanni</Emphasis>

We conducted grazing experiments with the three marine cladoceran genera Penilia, Podon and Evadne, with Penilia avirostris feeding on plankton communities from Blanes Bay (NW Mediterranean, Spain), covering a wide range of food concentrations (0.02–8.8 mm³ l^–1, plankton assemblages grown in mesocosms at different nutrient levels), and with Podon intermedius and Evadne nordmanni feeding on the plankton community found in summer in Hopavågen Fjord (NE Atlantic, Norway, 0.4 mm³ l^–1). P. avirostris and P. intermedius showed bell-shaped grazing spectra. Both species reached highest grazing coefficients at similar food sizes, i.e. when the food organisms ranged between 15 and 70 µm and between 7.5 and 70 µm at their longest linear extensions, respectively. E. nordmanni preferred organisms of around 125 µm, but also showed high grazing coefficients for particles of around 10 µm, while grazing coefficients for intermediate food sizes were low. Lower size limits were >2.5 µm, for all cladocerans. P. avirostris showed upper food size limits of 100 µm length (longest linear extension) and of 37.5 µm particle width. Upper size limits for P. intermedius were 135 µm long and 60 µm wide; those for E. nordmanni were 210 µm long and 60 µm wide. Effective food concentration (EFC) followed a domed curve with increasing nutrient enrichment for P. avirostris; maximum values were at intermediate enrichment levels. The EFC was significantly higher for P. intermedius than for E. nordmanni. With increasing food concentrations, the clearance rates of P. avirostris showed a curvilinear response, with a narrow modal range; ingestion rates indicated a rectilinear functional response. Mean clearance rates of P. avirostris, P. intermedius and E. nordmanni were 25.5, 18.0 and 19.3 ml ind.^–1 day^–1, respectively. Ingestion rates at similar food concentrations (0.4 mm³ l^–1) were 0.6, 0.8 and 0.9 g C ind.^–1 day^–1.Communicated by O. Kinne, Oldendorf/Luhe     

Trophic impact, metabolism, and biogeochemical role of the marine cladoceran Penilia avirostris and the co-dominant copepod Oithona nana in NW Mediterranean coastal waters

In this work we studied the trophic ecology and feeding impact of the cladoceran Penilia avirostris and the cyclopoid copepod Oithona nana, the two dominant zooplankters in the summer communities of the coastal NW Mediterranean, on the naturally occurring microbial communities. In order to ascertain carbon surplus for growth and reproduction and the contribution to carbon and nitrogen recycling of these two species, we also determined their basal metabolism and excretion rates. The experiments conducted during summers 2002, 2003, and 2004 indicate that P. avirostris grazed mostly upon small flagellates, dinoflagellates, and diatoms, whereas O. nana had a narrower prey range, selecting motile organisms such as ciliates and occasionally dinoflagellates. The grazing impact of both species accounted, on average, for <10% of the standing stock of the microbial groups considered. In spite of the oligotrophic conditions, the feeding activity of P. avirostris is in general sufficient to compensate basal metabolism and allows a surplus for growth and reproduction. This was not the case for O. nana, its daily rations being often lower than the carbon basal demands. Regarding excretion rates, both species presented different N:P excretion ratios, the ones of O. nana falling within values typical for copepods, whereas the absence of detectable phosphorus excretion by P. avirostris implied an unbalance recycling with respect to typical Redfield ratio composition of marine seston.     

The effect of temperature on the development and hatching of resting eggs of non-indigenous predatory cladoceran Cercopagis pengoi in the Gulf of Finland, Baltic Sea

The effect of temperature on the development and hatching of resting eggs of the Ponto-Caspian Cercopagis pengoi was studied experimentally in the Gulf of Finland, Baltic Sea. Morphological changes were monitored as the development of the resting eggs proceeded. Sexual reproduction of the C. pengoi population in the Gulf of Finland was evaluated by combining the data from hatching experiments and resting egg abundances in the sediment. Development time of resting eggs was dependent on temperature: increase in the temperature shortened the time needed until hatching. Hatching success was also dependent on incubation temperature. Almost sixfold increase in hatching success was detected when temperature increased two degrees above the storage temperature. Average resting egg abundances varied between 0.16 and 0.49 eggs cm⁻³ in the 0–6 cm sediment layer.