Guide to the coastal and surface zooplankton of the south-western Indian Ocean. Occasional Publication of the Marine Biological Association 15

A guide compiled as an aid to researchers in the identification of the coastal and shallow water, south-western Indian Ocean pelagic zooplankton, as much of the identification literature covering this area of amazing biodiversity is currently spread through the scientific literature and not accessible without extensive library resources. Most zooplankton groups, except fish larvae and eggs, have been covered, but some specialist groups have not yet been dealt with in great detail. However, a selection of representative members of most groups have been given, so that organisms can at least be assigned to perhaps a particular genus within the main group. The species list is based on zooplankton sampling carried out round the coastal areas of the islands of Mahé and Aldabra (Seychelles), Rodrigues (Mauritius), Madagascar and from a sampling transect between Seychelles and Rodrigues. The guide therefore includes a high proportion of the island-coastal and surface water zooplankton of the whole Indian Ocean. The location where a particular species has been sampled has been noted and some species that have not been sampled, but are known to occur in the region, have been included. Comprehensive taxonomic information has not been presented, but sufficient information should be given to identify each species. Keys have not yet been included for genera, as further species will be added. A bibliography of relevant plankton references has also been included.

Identification of the copepodite developmental stages of twenty-six North Atlantic copepods. Occasional Publication of the Marine Biological Association 21.

In zooplankton copepod studies there is often the requirement to be able to identify the six copepodite developmental stages of different species, or to know their body dimensions. However, this information is not available for many species, or is dispersed through the literature. This guide gathers together both original and previously published information on morphology and measurements for the stages of twenty-six common North Atlantic copepod species and tabulates them in a standard format. For each species additional notes useful in their identification are also given.

A new species of Tortanus (Atortus) (Copepoda: Calanoida) from the Seychelles, Mauritius and Madagascar

A new species of the planktonic copepod Tortanus (Atortus) (Calanoida: Tortanidae), T. (A.) magnonyx is described from Seychelles, Mauritius and Madagascar. This is the sixth species of the Indian Ocean recticauda species group, of the Indo-West Pacific recticauda species complex, that has been described from the western Indian Ocean. The inshore areas where these copepods are found have been poorly surveyed, so the number of species found implies a high diversity.

A new species of Tortanus (Atortus)(Copepoda: Calanoida: Tortanidae) from Rodrigues Island, Mauritius

A new species of the copepod genus Tortanus (Atortus) (Copepoda: Calanoida), T. (A.) insularis, is described from the coastal region of Rodrigues Island, Mauritius, an isolated island in the south-western Indian Ocean. This new species can be assigned to an Indian element, the recticauda species group, in the Indo-West Pacific recticauda species complex. It exhibits the closest relationship to T. (A.) recticauda recorded from the southern Red Sea and Arabian Gulf, rather than the more closely distributed T. (A.) capensis, recorded from off South Africa.

Epipelagic mesozooplankton distribution and abundance over the Mascarene Plateau and Basin, south-western Indian Ocean

The crescent shaped Mascarene Plateau (southwestern Indian Ocean), some 2200 km in length, forms a partial barrier to the (predominantly westward) flow of the South Equatorial Current. Shallow areas of the Mascarene Plateau effectively form a large shelf sea without an associated coastline. Zooplankton sampling transects were made across the plateau and also the basin to the west, to investigate the role the partial interruption of flow has on zooplankton biomass and community structure over the region. Biomass data from Optical Plankton Counter (OPC) analysis, and variability in community structure from taxonomic analysis, appear to indicate that the obstruction by the plateau causes upwelling, nutrient enrichment and enhanced chlorophyll and secondary production levels downstream. The Mascarene Basin is clearly distinguishable from the ridge itself, and from the waters to the south and north, both in terms of size-distributed zooplankton biomass and community structure. Satellite remote sensing data, particularly remotely-sensed ocean colour imagery and the sea surface height anomaly (SSHA), indicate support for this hypothesis. A correlation was found between OPC biovolume and SSHA and sea surface temperature (SST), which may indicate the physical processes driving mesozooplankton variability in this area. Biomass values away from the influence of the ridge averaged 24 mg m-3, but downstream if the ridge biomass averaged 263 mg m-3. Copepods comprised 60% of the mean total organisms. Calanoid copepods varied considerably between regions, being lowest away from the influence of the plateau, where higher numbers of the cyclopoid copepods Oithona spp., Corycaeus spp. and Oncaea spp., and the harpacticoid Microsetella spp. were found.

Flexible diel vertical migration behaviour of zooplankton in the Irish Sea

The diel vertical migration (DVM) of the whole plankton community was investigated in the central and coastal Irish Sea. Generally, more than 60% of the plankton did not perform significant DVM. A correlation analysis of the weighted mean depths of different organisms and their potential predators suggested relationships between two groups, Oithona spp., copepod nauplii and fish larvae, and between Pseudocalanus elongatus, Calanus spp. and chaetognaths. The organisms showing significant DVM were chaetognaths (Sagitta spp.), Calanus spp. and P. elongatus. Calanus spp. showed clear ontogenic variations in DVM, and along with P. elongatus demonstrated great flexibility both in the amplitude and direction of migration. P. elongatus did not migrate in the coastal area and Calanus spp. showed a clear reverse migration. The direction of migration appeared to be related to the vertical position of the chaetognaths in the water column during the day.

Use of satellite data for modelling food availability and survival of marine fish larvae

A modelling scheme is described which uses satellite retrieved sea-surface temperature and chlorophyll-a to derive monthly zooplankton biomass estimates in the eastern North Atlantic; this forms part of a bio-physical model of inter-annual variations in the growth and survival of larvae and post-larvae of mackerel (Scomber scombrus). The temperature and chlorophyll data are incorporated first to model copepod (Calanus) egg production rates. Egg production is then converted to available food using distribution data from the Continuous Plankton Recorder (CPR) Survey, observed population biomass per unit daily egg production and the proportion of the larval mackerel diet comprising Calanus. Results are validated in comparison with field observations of zooplankton biomass. The principal benefit of the modelling scheme is the ability to use the combination of broad scale coverage and fine scale temporal and spatial variability of satellite data as driving forces in the model; weaknesses are the simplicity of the egg production model and the broad-scale generalizations assumed in the raising factors to convert egg production to biomass.

Wind mixing, food availability and mortality of anchovy larvae Engraulis encrasicolus in the northern Adriatic Sea

A study was carried out in June/July 1996 in the River Po outflow in the northern Adriatic to investigate spawning of anchovy Engraulis encrasicolus and survival of larvae in relation to food availability and wind mixing. Hydrographic- and bongo net sampling was carried out on 2 grid surveys; one after a period of low winds and settled weather, and the other after an intervening period of strong winds, which resulted in a decrease in water column stratification. The spawning areas of anchovy and the larval distributions were associated with the river outflow plume (most clearly on the second survey grid, after the period of higher winds). Potential food particles for anchovy larvae, primarily copepod nauplii and copepodite stages, were also concentrated in the area influenced by the river outflow. Although there was a nearly 50% reduction in the mean water column abundance of potential food particles between the 2 survey grids, mostly due to a decline in abundance outside the immediate river plume area, there was no significant change in mortality of anchovy larvae between the 2 grids; the exponential decline in numbers of eggs and larvae to 10 mm in length being equivalent to overall mortality rates of 43.2%/d on the first survey and 44.7%/d on the second. The resilience of larval survival under potentially less favourable feeding conditions, following the period of wind mixing, was ascribed, in part, to the maintenance of local water column stratification by the superficial low salinity input from the River Po. This stratification in the immediate outflow area was associated with the presence of concentrated layers of potential food particles (typically >50 particles/L and 1.5 to 2.8 times the mean water column abundance) in the upper 10 m of the water column, coincident with peak numbers of anchovy larvae. However, since there was no evidence for lower larval survival in areas, less influenced by the immediate river outflow plume, a simple direct relationship between enhanced water column stability, improved feeding conditions and larval survival was not supported.

Buoyancy measurements and vertical distribution of eggs of sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus)

Measurements were made of the density and settling velocity of eggs of sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus), using a density-gradient column. These results were related to observed vertical distributions of eggs obtained from stratified vertical distribution sampling in the Bay of Biscay. Eggs of both species had slightly positive buoyancy in local seawater throughout most of their development until near hatching, when there was a marked increase in density and they became negatively buoyant. The settling velocity of anchovy eggs, which are shaped as prolate ellipsoids, was close to predictions for spherical particles of equivalent volume. An improved model was developed for prediction of the settling velocity of sardine eggs, which are spherical with a relatively large perivitelline volume; this incorporated permeability of the chorion and adjustment of the density of the perivitelline fluid to ambient seawater. Eggs of both species were located mostly in the top 20 m of the water column, in increasing abundance towards the surface. A sub-surface peak of egg abundance was sometimes observed at the pycnocline, particularly where this was pronounced and associated with a low-salinity surface layer. There was a progressive deepening of the depth distributions for successive stages of egg development. Results from this study can be applied for improved plankton sampling of sardine and anchovy eggs and in modelling studies of their vertical distribution.

Spawning season and temperature relationships for sardine (Sardina pilchardus) in the eastern North Atlantic

Spawning temperature preferences for sardine (Sardina pilchardus) in the eastern North Atlantic were determined from field data. These were compared with climatological temperature cycles (1986-2002) derived from satellite data by geographical region, to predict spawning seasons. Optimum spawning temperatures were determined as 14.0-15.0oC from the English Channel to Portugal and 16.0–18.0oC for all north-west African regions. Spawning seasons were closely related to the general latitudinal trend of the annual temperature cycle, with modification by upwelling in the western Iberian and north-west African regions. Some differences between temperature-based spawning season predictions and field observations were related to variations in seasonal plankton production. Correlations in the annual time-series of favourable spawning temperatures suggested relatively strong linkages between the southern areas from Portugal to Senegal. There was no consistent relationship between annual variations in duration of temperature-predicted spawning seasons and observed field abundance of eggs.

Feeding of anchovy Engraulis encrasicolus larvae in the northwestern Adriatic Sea in response to changing hydrobiological conditions

Results from depth integrated and vertically stratified plankton sampling in the northwestern Adriatic Sea were used for comparison of gut contents of larvae of European anchovy Engraulis encrasicolus with composition and concentration of potential prey in the plankton. Sampling was carried out over a grid of stations both before and after a period of increased wind mixing to investigate changes in food availability and larval feeding success. All larvae had empty guts soon after dusk, indicating daytime feeding and rapid gut clearance. With increasing larval length there was a greater percentage of specimens with empty guts, despite suitable food being available in the plankton for these larger larvae; this suggests differential gut evacuation during sampling-possibly related to the degree of gut development. Larval diet was principally the various developmental stages of copepods, especially calanoid and cyclopoid nauplii, which were preferentially selected by larvae, whereas selection was against harpacticoid nauplii. Lamellibranch larvae and Peridinium were generally abundant in the plankton, but were only present in the gut contents in any number when the preferred dietary organisms were present in the plankton at low concentrations. The number of food organisms in the gut contents increased with concentration of the preferred food organisms in the plankton up to a limit of similar to 50 organisms/l. Within the upper 18 m of the water column, there was a reduction in the proportion of larvae with food in their guts with increasing depth, irrespective of the vertical profile of food concentration. Following a period of wind mixing the composition of the plankton changed. This was reflected in the diet of anchovy larvae, which altered in parallel. There was also an overall 41% decrease in concentration of the preferred food particles of larvae in the plankton following the period of wind mixing, but larvae were still able to maintain their food intake. These results show that anchovy larvae can successfully adapt their diet to a changing prey field and suggest that in the conditions observed in the northern Adriatic, quite radical changes in the feeding environment were probably insufficient to affect overall larval mortality.

Digestion of natural food by larval and post-larval turbot Scophthalmus maximus

The digestion of natural, mainly crustacean zooplankton, by different age groups of turbot Scophthalmus maximus larvae was evaluated by comparisons of visual appearance, dry weight and carbon and nitrogen content of fresh food organisms with material recovered from faeces. Visually, the degree of digestion of food particles ranged from no discernible change of lamellibranch larvae, copepod eggs, intact copepod faecal pellets and some phytoplankton species, to varying degrees of removal of body constituents in copepods, cladocerans and decapod zoea. For crustaceans, the proportion of body constituents removed was related to the size and construction of their apparently indigestible exoskeleton. Uppon defaecation larger organisms showed the greatest percentage loss in dry weight and carbon. A high percentage of nitrogen was extracted from all organisms.

Digestion of copepod eggs by larval turbot Scophthalmus maximus and egg viability following gut passage

Between 20.5 and 93.6 % of the subitaneous eggs of 6 species of egg-carrying copepods passed undigested through the digestive tracts of larval and early postlarval turbot Scophthalmus maximus. Viability of the eggs of Eurytemora affinis, E. velox and Euterpina acutifrons remained high on egestion (67.0 to 91.7 %), Pseudocalanus elongatus and Oncaea venusta eggs had low viability (1.1 to 1.5 %), while all Corycaeus anglicus eggs were rendered inviable. The indigestibility of the eggs denies the turbot larvae a potentially valuable food resource, while retention of high egg viability in certain species reduces the effect of predation.