Feeding ecology and interannual variations in diet of larval jack mackerel,Trachurus declivis (Pisces: Carangidae), from coastal waters of eastern Tasmania

Changes in the biochemical composition of the digestive gland and in the proteins of the mantle muscle of Sepia officinalis L, collected in September 1989 from the Ria de Vigo (northwest Spain), were measured during periods of 2, 4, 10 and >53 d starvation. The digestive gland lost weight faster than the rest of the body throughout the whole period of starvation. In the digestive gland, carbohydrate and protein contents did not change during starvation; however, lipid levels decreased significantly after 53 d. Phospholipid content increased during longterm starvation. The content of free fatty acids rose after 16 d. Sterols, diacylglycerylethers, triacylglycerols and carotenoids contents did not change significantly. Of the total fatty acids, 18:0, 20:2n6, 20:4n6 and the monounsaturated moieties were preferentially consumed; others, such as 22:5n3, 22:6n3 and 16:4n1, were selectively retained. In the mantle muscle, water content increased and total protein content decreased. The myofibrillar proteins decreased after 53 d starvation, whereas the sarcoplasmic fraction did not change and the stromatic proteins increased. No changes were observed in the electrophoretic patterns of sarcoplasmic and myofibrillar proteins. The digestive gland of S. officinalis does not seem to be an important reserve organ during long-term starvation, but does seem to be important during shortterm starvation.     

Feeding ecology of five fishes from the mid-slope micronekton community off southern Tasmania, Australia

We examine the feeding ecology of characteristic micronekton species inhabiting the mid-slope region (~700 to 1,500 m depths) off southern Tasmania. Five fishes, Diaphus danae, Hygophum hanseni, Lampanyctus australis (Myctophidae), Phosichthys argenteus (Phosichthyidae) and Chauliodus sloani (Chauliodontidae), were sampled by depth-stratified midwater trawling on a diel and seasonal basis. Overall, 74 prey taxa were identified from 2,132 stomachs. Euphausiids (mostly Euphausia spinifera and E. similis) and calanoid copepods (mostly Pleuromamma species) were the main prey of the three myctophids; P. argenteus ate fishes and decapods in addition to the euphausiids, while C. sloani ate only fishes. Copepods were less important in the diets of larger D. danae, L. australis and P. argenteus and were replaced by euphausiids in the myctophids and by fishes in P. argenteus. In autumn, when euphausiid biomass increased six-fold, all three myctophids and P. argenteus fed most intensively and consumed a high proportion of euphausiids. The three myctophids appeared to feed nocturnally. Differences in the timing and duration of feeding corresponded to differences in their spatio-temporal overlap with key prey. Daily rations of H. hanseni (1.93% dry-weight biomass) and L. australis (1.43%), estimated from data on stomach fullness, were typical for temperate myctophids and higher than that of the non-migratory P. argenteus (0.43%). The vertical flux of near-surface plankton production to the mesopelagic food web is based primarily on diel feeding in the upper water column (<500 m) rather than consumption of species that migrate seasonally into the deeper mesopelagic zone. Because species such as P. argenteus and C. sloani feed above the third trophic level, their predators, including the commercially important orange roughy, are feeding between levels four and five.     

Feeding ecology of three species of midwater fishes associated with the continental slope of eastern Tasmania,Australia

The feeding ecology of Maurolicus muelleri, Lampanyctodes hectoris and Diaphus danae was examined from samples collected from continental slope waters of eastern Tasmania between April 1984 and April 1985. A total of 2 232 stomachs was analysed. M. muelleri, L. hectoris and D. danae fed primarily on euphausiids and secondarily on copepods, although larger D. danae (> 60 mm standard length) fed on other lanternfish (chiefly L. hectoris). The diets of M. muelleri and L. hectoris overlapped substantially. Diet overlaps between D. danae and the former species was low, however, due to the large biomass of fish present in D. danae. The prey taxa consumed changed with time of year and predator size. Stomach fullness (feeding intensity) varied seasonally in all three species, but only M. muelleri showed significant diel differences in fullness. The synchronization of the size structure of the predator populations and their feeding intensity, with seasonal variations in preferred prey, is proposed as a mechanism whereby each species maximizes its share of the available food resources.     

Testing the abundant center model using range-wide demographic surveys of two coastal dune plants

It is widely accepted that species are most abundant at the center of their geographic ranges and become progressively rarer toward range limits. Although the abundant center model (ACM) has rarely been tested with range-wide surveys, it influences much thinking about the ecology and evolution of species' distributions. We tested ACM predictions using two unrelated but ecologically similar plants, Camissonia cheiranthifolia and Abronia umbellata. We intensively sampled both throughout their one-dimensional distributions within the Pacific coastal dunes of North America, from northern Baja California, Mexico, to southern Oregon, USA. Data from > 1100 herbarium specimens indicated that these limits have been stable for at least the last 100 years. Range-wide field surveys detected C. cheiranthifolia at 87% of 124 sites and A. umbellata at 54% of 113 sites, but site occupancy did not decline significantly toward range limits for either species. Permutation analysis did not detect a significant fit of geographical variation in local density to the ACM. Mean density did not correlate negatively with mean individual performance (plant size or number of seeds/plant), probably because both species occur at low densities. Although size and seeds per plant varied widely, central populations tended to have the highest values for size only. For C. cheiranthifolia, we observed asymmetry in the pattern of variation between the northern and southern halves of the range consistent with the long-standing prediction that range limits are imposed by different ecological factors in different parts of the geographical distribution. However, these asymmetries were difficult to interpret and likely reflect evolutionary differentiation as well as plastic responses to ecological variation. Both density and seeds per plant contributed to variation in seed production per unit area. In C. cheiranthifolia only, sites with highest seed production tended to occur at the range center, as predicted by the ACM and assumed by theory proposing that range limits evolve via antagonism between natural selection and gene flow.     

Horizontal and vertical trends in the distributions of larval fishes in coastal waters off central New South Wales,Australia

Larval fishes were sampled across six transects perpendicular to a 50 km section of the coast off Sydney, Australia, in April/May and August/September 1990. Samples were collected at the surface and at depth (20 to 30 m) at three locations across each transect; over the 30, 70 and 100 m depth contours. There was a large level of heterogeneity in the horizontal and vertical distributions of most taxa examined, and no general pattern of distribution was evident for the whole assemblage. Classification analyses revealed that the major differences between assemblages were related to depth. Horizontal trends in the distributions of the abundant taxa were evident in the inshore-offshore direction, but not longshore. Seven taxa belonging to the families Gobiidae, Labridae, Sillaginidae, Sparidae, Ambassidae, Clupeidae and Clinidae/Tripterygiidae were most abundant inshore, whereas 4 taxa of the families Cepolidae, Percichthyidae, Cheilodactylidae and Gonorynchidae were generally more abundant offshore and 24 taxa showed no discrete horizontal trends across transects. More taxa and individuals were generally caught at depth than at the surface and this was evident across all transects. Twenty taxa were more numerous at depth, whereas 4 taxa, the Cheilodactylidae, Gonorynchidae, Mullidae and Scorpididae, were most abundant at the surface and 11 taxa showed no difference in densities between depths. Ontogenetic differences in the distributions of some larvae were evident. The mean size of larval Liza argentea (Mulgilidae) caught was greater offshore than inshore, and greater at the surface than at depth. In contrast, larger Pseudocaranx dentex (Carangidae) occurred in greater numbers at depth than at the surface. The data emphasise that the assemblages of larval fishes in coastal waters off central New South Wales cannot be modelled as a single unit, which concurs with the findings in other temperate and tropical vaters. Furthermore, the data denote the need to spatially stratify sampling in these waters in order to assess seasonal changes in these assemblages.     

Stable isotopic evidence for trophic groupings and bio-regionalization of predators and their prey in oceanic waters off eastern Australia

Muscle tissue was collected for stable isotope analysis (SIA) from the main fish predators and their fish and cephalopod prey from oceanic waters off eastern Australia between 2004 and 2006. SIA of δ¹⁵N and δ¹³C revealed that the species examined could be divided into three main trophic groups. A “top predator” group consisted mainly of large billfish (Xiphias gladius and Tetrapturus audax), yellowfin (Thunnus albacares), bigeye (T. obesus) and southern bluefin (T. maccoyii) tunas and sharks; with mako (Isurus oxyrinchus) the highest. Below this tier was a second group composed of mid-trophic level fishes including albacore tuna (Thunnus alalunga), lancet fish (Alepisaurus ferox), mahi mahi (Coryphaena hippuris) and ommastrephid squid. Underlying both groups was a grouping of small fishes including myctophids, small scombrids and nomeids as well as surface fishes including macrorhamphosids. These groupings were based largely on mean animal size which showed a positive linear relation to δ¹⁵N (r ² = 0.58). Some species showed significant ontogenetic variation in either δ¹⁵N (swordfish, lancet fish, yellowfin and albacore tuna) or δ¹³C (mako shark). We also noted a consistent latitudinal change in δ¹⁵N and δ¹³C at ~28°S for the top predator species, particularly albacore and yellowfin tuna. The differences were consistent with a change from oligotrophic Coral Sea to nutrient rich Tasman Sea waters. These differences suggest that predatory fishes may have extended residence time in distinct regions off eastern Australia.     

Metal concentration changes in growing Pacific oysters,Crassostrea gigas,cultivated in Tasmania,Australia

Pacific oyster spat (Crassostrea gigas Thunberg) transferred from the Tamar Estuary to two growing areas in southern Tasmania were monitored for their metal contents over one growing season (1974–1975). Oysters at Pipeclay Lagoon were grown with stick and tray culture while those at Dart Island were cultured with the longline technique. Metal content of the oysters increased with time and the trend was similar to the weight growth curves. Mean dry weights of oysters increased from 0.07 to 1.19 g at Pipeclay Lagoon and from 0.25 to 1.47 g at Dart Island. Metal contents (g) increased at each site, respectively: Fe 57 to 326; 91 to 446; Zn 259 to 6 555; 755 to 5 335; Cd 1.5 to 13.3; 1.9 to 16.3; Cu 26 to 142; 9 to 116; Pb 1.9 to 11.9; 0.6 to 3.8. Concentration curves generally showed a downward trend with time. The relationships of metal concentrations with weight did not differ from sample to sample at a site nor did they differ at one site compared with the other. The only exception was lead, which showed no relationship of concentration with weight at Pipeclay Lagoon and a negative one at Dart Island. It is postulated that higher winter concentrations of metals in the oysters were linked with greater solubility of metal ions in lower salinity water.     

Resource partitioning by four species of elasmobranchs (Batoidea: Urolophidae) in coastal waters of temperate Australia

Four species of stingarees (Urolophus lobatus, U. paucimaculatus, Trygonoptera personata and T. mucosa) were abundant in samples collected from over sandy substrates in four regions along ∼200 km of the lower west coast of Australia in each season between summer 1990/1991 and winter 1992. The main aim of this study was to determine whether these four species were segregated by habitat, size, sex and/or diet, thereby reducing the potential for intra- and interspecific competition. The densities of U. lobatus and T. personata were almost invariably greater in deep, offshore waters (20 to 35 m) than in shallow, inshore waters (5 to 15 m), while those of U. paucimaculatus were greatest in the southernmost region, irrespective of depth, and those of T. mucosa were highest at the single inshore site at which the water was deep. None of these species was segregated by either size or sex. The volumetric dietary compositions of the two Urolophus species, which consisted predominantly of various crustaceans, and those of the two species of Trygonoptera, which consisted mainly of different polychaetes, were significantly different from each other. The two Urolophus species initially fed largely on amphipods, mysids and carid decapods but, with increasing size, U. lobatus started to ingest teleosts and U. paucimaculatus commenced feeding on errant polychaetes and penaeid decapods. The contribution of errant polychaetes to the diets of T. personata rose progressively from 0% in the smallest fish to 90% in the largest fish, thereby accounting for a concomitant progressive decline in dietary breadth and for intraspecific dietary overlap typically to occur only between successive size classes. In contrast, the volumetric contribution of sedentary polychaetes to the diets of T. mucosa remained high (65 to 80%) throughout life, which accounts for the relatively low and constant dietary breadth of most size classes and a significant overlap in the diets of all but one pair of size classes. The diets of each of the four stingaree species occasionally differed between sites and sometimes underwent seasonal changes, presumably reflecting spatial and temporal differences in the types of prey in the environment. Since the ratios of benthic to epibenthic prey in the diets of the two Trygonoptera species were greater than in those of the two Urolophus species, the first two species apparently forage deeper into the substrate. The combination of partial segregation of the four stingaree species among habitats and the differences in the dietary compositions and feeding strategies of these species, would presumably facilitate the co-existence of these batoid elasmobranchs when they are abundant in the same general region. Received: 29 October 1997 / Accepted: 25 March 1998     

The trophic status of two Northern Irish sea loughs

Studies were conducted on nutrient cycling in two coastal embayments; Strangford Lough and Belfast Lough, in order to classify these coastal waters in compliance with the European Union Urban Waste Water Treatment and Nitrates Directives. Analysis of the data suggests that Strangford Lough is nitrogen-limited throughout much of the growing season with chlorophyll-a levels comparable to the open Irish Sea. In contrast, much of Belfast Lough is never nitrogen-limited and, with chlorophyll-a levels at time exceeding 50 μg/l, is exhibiting symptoms of eutrophication.     

The trophic status of two northern Irish Sea loughs

Studies were conducted on nutrient cycling in two coastal embayments; Strangford Lough and Belfast Lough, in order to classify these coastal waters in compliance with the European Union Urban Waste Water Treatment and Nitrates Directives. Analysis of the data suggests that Strangford Lough is nitrogen-limited throughout much of the growing season with chlorophyll-a levels comparable to the open Irish Sea. In contrast, much of Belfast Lough is never nitrogen-limited and, with chlorophyll-a levels at times exceeding 50 μg/l, is exhibiting symptoms of eutrophication.     

Temporal changes in the trophic ecology of the asymbiotic gorgonian Leptogorgia virgulata

Gorgonians are widespread sub-littoral benthic suspension feeders in the world oceans. However, data on their trophic ecology and role in benthic–pelagic coupling and biogeochemical cycles remain limited. This study assesses the trophic ecology of Leptogorgia virgulata in the Skidaway River estuary, USA (31.9896N, 81.0242W) by analysing carbon and nitrogen isotopic signatures of its soft tissue and different-sized fractions of particulate organic matter (POM) in its environment. Samples were taken in 5 April, 2 August, and 15 October 2012 and 11 January 2013. Results support a distinct temporal shift in the diet of L. virgulata from the POM fraction <10 µm (i.e. pico- and nanoplankton) to the 10–63-µm fraction (i.e. microplankton). This trophic regime is likely associated with the natural abundance of prey items within these size classes, thus suggesting that L. virgulata may be an opportunistic feeder, and seasonal shifts in food availability in the water column affect its diet. As such small prey items affect the bioenergetics of L. virgulata, it is important to understand the implications of changes in food availability associated with environmental drivers on the physiology and population dynamics of this dominant species in the western Atlantic Ocean.     

Reproductive biology of commercially exploited deep-water royal red prawns (Haliporoides sibogae,Solenoceridae) in south-east Australia

Deep-water royal red prawns (Haliporoides sibogae de Man, 1907; Solenoceridae: Penaeidea) were sampled from commercial catches off south-east Australia between November 1988 and May 1990. Four stages of sexual development were distinguished macroscopically for each sex. For the females, histology and analysis of oocyte size were used to validate the macroscopic staging of ovaries into undeveloped, developing, early ripe and ripe. Females were larger than males, reaching a maximum size of 46 mm carapace length (CL) compared to 32 mm CL for males. Average size at sexual maturity was 30.8 mm CL for females and 25.8 mm CL for males. Females apparently breed several times in their life, whereas males probably breed only once. Individual fecundity (ranging from about 58 000 to 140 000 oocytes) was relatively low compared with other penaeids, but oocytes were larger (mean diameter=0.41 mm). Two short breeding seasons occurred per year (February to April and July to August). There was a clear size-related latitudinal distribution of the prawns, with largr prawns (spawners) being more abundant in the northern region of the fishing area, and juveniles (CL20 mm) being more abundant in the south. Most of the spawning stock probably lies outside the main fishing ground, while the fishery targets mainly prawns below the size at maturity.     

Studies of primary productivity in coastal waters of southern Long Island,New York

Phytoplankton productivity of the tidal estuaries and coastal waters of southern Nassau Country, Long Island, New York, USA was determined monthly at 28 stations during 1966. Diatoms alternated with dinoflagellates in dominating the standing crop in the coastal area. The estuaries were characterized by sustained blooms of green flagellates and dinoflagellates during the spring/summer period, 1966. Chlorophyll a ranged from 1.0 to 27.6 mg/m³ in the estuarine area, and 1.45 to 10.15 mg/m³ in adjacent coastal waters. Rate of phytosynthesis per unit weight chlorophyll a for surface samples in the region under study a veraged from 3.1 to 3.5 mgC/mg chlorophyll a/h. At light saturation, however, the ratio varied according to water temperature and species' composition. Primary productivity decreased seawards, with mean values for 1966 of 0.35, 0.22, and 0.16 gC/m³/d for the estuarine, nearshore and offshore areas, respectively.This study was carried out at the Lamont Geological Observatory of Columbia University Palisades; New York, USA.The study was conducted with financial assistance from Nassau County, New York, and the G. Unger Vetlesen Foundation.     

Age and growth of the lanternfish Lampanyctodes hectoris (Myctophidae) from eastern Tasmania,Australia

The age, growth and mortality of the lanternfish Lampanyctodes hectoris (Gunther) were investigated by examining the otoliths, length-frequency data, and seasonal abundance of fish collected from continental slope waters of eastern Tasmania between April 1984 and June 1985. Although L. hectoris can live to 3 yr old, it has a high annual mortality (79%) and few live past their first year. Growth, described here by a variation of the Von Bertalanffy model, takes place mainly in the first six months of life and only during late spring and summer. The hypothesis of daily deposition of primary growth increments in myctophids was supported by comparing the back-calculated birth dates of daily-aged individuals of L. hectoris with the spawning season of this species.     

Use of seaweeds for monitoring trace elements in coastal waters

Concentrations of a wide range of trace elements: arsenic, cadmium, cobalt, chromium, hafnium, nickel, thorium, uranium, zinc and the rare earth elements, cerium, europium, samarium, terbium and ytterbium were determined by instrumental neutron activation analysis in the brown alga,Fucus vesiculosus from Eckwarder Hörne, North Sea and from Rügen, Baltic Sea. Another brown alga,Sargassum filipendula from Sri Lanka, Indian ocean (representing an unpolluted control station) was similarly investigated. Cobalt, chromium and nickel concentrations were highest inF. vesiculosus from the North Sea while zinc was highest in samples from the Baltic Sea, reflecting high levels of these elements in coastal waters of the North and the Baltic sea. Cadmium, cobalt, nickel and zinc levels were lowest inS. filipendula from Sri Lanka, probably demonstrating lower levels of those elements in coastal waters. Concentration levels of hafnium, thorium, uranium, and the rare earth elements were highest inS. filipendula. Two years later in 1994,S. filipendula along withUlva sp. (green alga) was resampled from the same sampling site, and in addition to the above elements, six other trace elements (Ag, Ba, Br, Rb, Se and Sr) were determined.Sargassium filipendula showed a particular affinity for Ag, As, Br and Sr. For the other elements, marginal concentration differences were observed betweenS. filipendula andUlva sp., probably reflecting the regional background levels. Substantially higher concentrations of Hf, Th, U, and the rare earths were found again in the 1994Sargassum andUlva samples, reflecting the effect of a substrate rich in rare earth elements. The brown algae used in this study may be used to monitor trace elements in coastal waters.     

Variability of dissolved organic matter in northern adriatic coastal waters

The results obtained in the four seasonal cruises planned in the PRISMA II project are reported. These concern dissolved and colloidal organic carbon, free amino acids and total dissolved carbohydrates and heterotrophic activity. Main factors controlling organic matter degradation, resulting from laboratory tests not planned in the above project, are also discussed. Dissolved organic matter shows seasonal accumulation, which may be markedly different from year to year, and large contributions by colloidal and saccharide components. Heterotrophic activities play an important role in the carbon cycle, although laboratory runs highlight limitations caused by aging of organic matter and phosphorus deficiency.     

Species composition, biomass and vertical distribution of micronekton over the mid-slope region off southern Tasmania, Australia

Seasonal sampling was carried out based on day/night, vertically stratified tows (100 or 125 m strata) in the upper 900 m of the water column over the mid-slope commercial fishing grounds south of Tasmania. A large midwater trawl (105 m² mouth area) was used with an opening/closing cod-end. Subtropical convergence and subtropical species dominated the fauna, but many less abundant, more widely-distributed species were also present. Fishes, which contributed 89% of micronekton biomass and 135 of 178 species, were dominated by the Myctophidae (48% biomass and 48 species). Twenty micronekton species made up 80% of the total biomass. Overall, the micronekton fish biomass in this region was 2.2 g m⁻² wet weight. A pronounced day/night shift in the distribution of biomass was attributable to diel migratory species. During the day, <0.2% of the total micronekton biomass was found in 0 to 300 m; most biomass was below 400 m, with peaks at 400 to 525 m and 775 to 900 m. At night, 53% of the biomass was found in 0 to 300 m, with progressively less in each deeper stratum. The vertical ranges of individual species typically exceeded 400 to 500 m during the day and night and were non-coincident, although nyctoepipelagic migrators were concentrated in the surface 200 m at night. Distinct epipelagic, lower and upper mesopelagic assemblages were identified, and patterns of epipelagic migration, limited migration and non-migration were categorised for species from each of the lower and upper mesopelagic assemblages. The vertical distribution of these assemblages was coincident with the primary water masses: subantarctic mode water (∼250 to 600 m) and antarctic intermediate water (below ∼700 m). The flux of migrating micronekton, estimated at 0.94 to 3.36 g C m⁻² yr⁻¹ to the lower mesopelagic and 1.14 to 4.06 g C m⁻² yr⁻¹ to the upper mesopelagic, appeared to be considerably outweighed by the consumption needs of aggregated mid-slope benthopelagic predators. We suggest that advection of mesopelagic prey in antarctic intermediate water may sustain aggregated populations of orange roughy (Hoplostethus atlanticus) and other predators on the micronekton in mid-slope depths at this site. Received: 2 April 1997 / Accepted: 21 August 1997     

Diets of fishes of the upper continental slope of eastern Tasmania: content,calorific values,dietary overlap and trophic relationships

Diets of 15 species of demersal and pelagic fishes on the upper continental slope (420 to 550 m) were determined, based on samples taken every two months over 13 mo (April 1984 to April 1985) off eastern Tasmania. The calorific contribution of each prey item to the diets was determined. The fish could be divided into four trophic categories: pelagic piscivores, epibenthic piscivores, epibenthic invertebrate feeders and benthopelagic omnivores. Dietary overlap between the groups was low. The pelagic piscivores Apogonops anomalus, Trachurus declivis, Brama brama, Lepidopus caudatus and Macruronus novaezelandiae primarily consume the shelf-break myctophid Lampanyctodes hectoris; their diet is narrow, with a large overlap between species. The epibenthic piscivores Deania calcea and Genypterus blacodes both take a greater variety of prey, but have little dietary overlap. The fish feeding on epibenthic invertebrates, Coelorinchus sp. 2 and Centriscops humerosus, obtain most of their energy from benthic Crustacea and Ophiuroidea, supplemented with Lampanyctodes hectoris; the diet is broad, with little overlap. Among the benthopelagic omnivores (Cyttus traversi, Coelorinchus sp. 4, Lepidorhynchus denticulatus, Neocyttus rhomboidalis, Helicolenus percoides, Epigonus denticulatus and E. lenimen), most diets are broad and show slight overlap. All but E. denticulatus consume significant quantities of Lampanyctodes hectoris as well as Crustacea, particularly Polychaeta, Euphausiacea and Pyrosoma atlanticum. Seasonal changes in diet occurred in G. blacodes, T. declivis, Lepidopus caudatus, Coelorinchus sp. 4, Lepidorhynchus denticulatus, H. percoides, E. denticulatus and E. lenimen; these were related to changes in abundance of particular prey species, not to alterations in feeding habits. Only three species, Lepidopus caudatus, Coelorinchus sp. 2 and H. percoides, showed significant diel feeding periodicity. Ontogenetic dietary changes were evident in Cyttus traversi, Coelorinchus sp. 2, Lepidorhynchus denticulatus and H. percoides. Cyttus traversi and H. percoides progressively changed from crustaceans to fish as their size increased. The diets of size classes within species showed little overlap, except for Lepidorhynchus denticulatus, which eats chiefly euphausiids and Lampanyctodes hectoris at all sizes. In addition to describing the diets and trophic relationships of 90% of the fish biomass, the results emphasize the importance to the entire fish community of mesopelagic food resources, particularly Lampanyctodes hectoris. Many benthopelagic species undertake extensive vertical migrations in search of prey, thus playing a major role in the transport of energy from midwater regions to the benthos of the continental slope.