Observations on mesopelagic fish from the Red Sea

Mesopelagic fish were collected from a cruise with the R.V. “Dr. Fridtjof Nansen” to the northern Red Sea and the interior Gulf of Aden off the Republic of Djibouti in March 1981. Off the Republic of Djibouti five species of mesopelagic fishes were caught and in the Red Sea six species. Benthosema pterotum (Alcock) were dominant in both areas. In the Red Sea Maurolicus muelleri (Gmelin) were also abundant. The mesopelagic fish caught in the Red Sea showed a pattern of vertical migration similar to that observed in other areas, in spite of the special hydrographical regime. The number of gill rakers of B. pterotum caught in the Red Sea differed significantly from that of all other populations studied. The number of photophores of M. muelleri apparently do not differ much from other populations. B. pterotum from the Red Sea mature at a larger size than other populations, while M. muelleri mature at a smaller size. The relative fecundity of M. muelleri is higher in the Red Sea than in other regions studied, while no significant difference was observed in B. pterotum. Both species appear to be opportunistic feeders. Their feeding chronology in the Red Sea seems to be similar to those observed in other areas.     

Observations on the embryonic development of some deep-living decapod crustaceans,with particular reference to species of Acanthephyra

Eggs of meso- and bathypelagic decapod crustaceans were removed from gravid females and allowed to develop in vitro. The time course of embryonic development, as indicated by the appearance of certain morphological and physiological features, was determined for most species at a constant temperature of 12°C. The embryonic period, and the time between the appearance of certain key features such as naupliar and compound eyes and heartbeats, showed little variation in any one species. Significant differences were observed in the development times of different species. In general, species with large eggs have a longer embryonic period than those with small eggs, but there also variations between species with eggs of similar size. During the development of Acanthephyra eggs there is an approximate doubling of the egg volume, correlated with a decrease in egg density and an increase in the water content. The increase in the water content and egg volume is similar to that of other decapod eggs. It is concluded that those species with small eggs could have several broods in 1 year, whereas those species with large eggs are likely to be restricted to one, or in some cases possibly two, broods.     

Patterns and selectivity in the feeding of certain mesopelagic fishes

Stomach contents were analysed from the 7 most numerous species of mesopelagic fish caught in a series of 11 hauls over a 24 h period at 230 to 266 m depth in the eastern North Atlantic Ocean. The numerical abundance of organisms per filled stomach and the frequency of occurrence of empty stomachs were used to indicate feeding periodicity. The ecological significance of the feeding periodicity was considered by examining it in connection with an investigation of the day-night vertical distribution of zooplankton and micronekton to 2000 m at the same station. Additional dietary evidence on the 7 species considered was also obtained from the vertical series. Feeding selectivity was examined by comparing the composition of the zooplankton population, sampled separately but simultaneously with the micronekton, with that from the overall stomach contents of the species examined. Feeding periodicity was demonstrated for 6 species, of which 3 were found to be feeding selectively: Valenciennellus tripunctulatus on calanoid copepods, Argyropelecus aculeatus on ostracods, and Lampanyctus cuprarius on amphipods and possibly euphausiids. The limited data available on the other 3 species suggested that they were either random feeders (A. hemigymnus and Lobianchia dofleini) or perhaps selecting against a particular group (Notolychnus valdiviae). No indication of feeding periodicity or selectivity was found for Chauliodus danae. The overall pattern of results confirmed the supposed close correlation between vertical migration and feeding in mesopelagic fish.     

Concentrations of Mn,Fe, Cu,Zn and Cd in the mesopelagic decapod Systellaspis debilis from the East Atlantic Ocean

Specimens of the oceanic decapod Systellaspis debilis were collected from six sites in the East Atlantic Ocean between 1970 and 1984, and were analysed for Mn, Fe, Cu, Zn and Cd. The data confirm that there are small but significant differences in mean metal concentrations from some sites which showed no obvious pattern in relation to geographic location of the samples. As a result, ranges of site means are quoted as baseline levels for each metal (g g^-1 dry wt): 2.3 to 2.9 g Mn g^-1, 31.2 to 77.8 g Fe g^-1, 25.9 to 83.4 g Cu g^-1, 41.9 to 92.9 gZn g^-1, 11.1 to 31.8 g Cd g^-1. The concentration of cadmium in S. debilis from all sites was raised relative to cadmium concentrations reported for coastal decapods, perhaps as a result of dietary enrichment. Metal accumulation may provide useful information for understanding the complex feeding behaviour of many oceanic animals.     

Using fatty acid analysis to elucidate the feeding habits of Southern Ocean mesopelagic fish

Fatty acid biomarkers were used to investigate the feeding ecology of 17 mesopelagic fish species occurring in the Southern Ocean. Fatty acid signatures of species where little or no dietary information exists were compared to fatty acid signatures of species of known diets in order to elucidate their trophic position. Principal component analysis grouped species of known diets into two clusters with amphipods and copepods comprising the main prey species, respectively. Although the majority of species of unknown diet were grouped with either of these feeding guilds, a third cluster comprising only Gymnoscopelus bolini was identified suggesting a significantly different diet for this species. Electrona antarctica also exhibited significant changes in fatty acid signatures with size. Furthermore, discriminant analysis of the four most abundant species classified species with a 90% success rate thus validating the usefulness of fatty acid signatures when trying to resolve the trophic position of species where no or little dietary information exists.     

Observations on the burrows and burrowing behaviour of two mud-dwelling decapod crustaceans,Nephrops norvegicus and Goneplax rhomboides

The burrows of the Norway lobster Nephrops norvegicus (L.) and of the crab Goneplax rhomboides (L.) were studied in Loch Torridon, Scotland. Polyester resin casts of burrows in the sea were made by divers to reveal their subsurface form. Tunnels made by N. norvegicus were usually simple, with two or more openings on the mud surface, and penetrated to a depth of about 30 cm. G. rhomboides burrows did not descend more than about 15 cm beneath the surface, but were usually more complex than the lobster burrows and had several openings. The methods of burrow construction used by the two crustaceans are described from aquarium observations. Neither N. norvegicus nor G. rhomboides show obvious morphological adaptations for burrowing, and it is suggested that the fossorial habit was adopted very early by decapods. The burrows of N. norvegicus do not seem to have assumed any functions in addition to the original one of providing refuge from predators. There is not sufficient known of the biology of the crab to indicate whether the same is true in its case.     

Molt cycle alterations in behavior,feeding and diel rhythms of a decapod crustacean,the spiny lobster Panulirus argus

The crustacean molt cycle manifests extensive behavioral changes in addition to physiological and integumentary modifications. The paucity of quantitative studies led us to characterize molt stage dependent alterations in rhythmic locomotor, feeding, and agonistic behavior of subadult spiny lobster Panulirus argus held grouped and solitarily in simulated natural conditions. Non-disruptively determined molt stages were defined as proportions of intermolt duration. Significant nocturnal rhythmicity persisted through the full molt cycle, though daily form varied as a function of activity level and molt stage. A distinct early evening peak signaled initiation of foraging and walking behavior. Thereafter, rhythm amplitude either declined progressively (low activity: unimodal pattern), remained high (high activity: nocturnal plateau), or exhibited a secondary early morning peak (intermediate activity: bimodal and polymodal patterns). Activity ceased at or immediately prior to sunrise. Ecdysis was predominantly nocturnal, probably reflecting species specific spatiotemporal movement patterns and social behavior. A selective advantage of nocturnal ecdysis was postulated concerning avoidance of cannibalism and diurnal predators. Locomotor activity and feeding rates were not equivalent through the molt cycle, though both peaked in stages B₂-C₁. Locomotor activity remained high in metecdysis, decreased sharply at proecdysis initiation (D₀), reaching lowest levels in D₁-D₃, then increased in D₃ through B₁. Activity dropped steeply at ecdysis, though lobsters were capable of intense and coordinated activity. Feeding decreased slowly through metecdysis after the B₂-C₁ maximum, then increased temporarily in C₄ and D₀, indicating heightened feeding motivation. This contrasts with the locomotor activity decrease at proecdysis. Food consumption declined rapidly in D₁ and D₂ and ceased at the D₂-D₃ transition. Feeding resumed in B₁ or B₂, intensifying to maximum in late B₂. Feeding remained relatively constant within stages, whereas locomotor activity varied greatly, though both correlated with metabolic needs. Grouped and solitary lobsters displayed similar patterns of foraging and walking, equivalent to those of locomotor activity and feeding of solitary individuals. Frequency of agonistic interactions (not aggression per se) remained relatively constant through the cycle, peaking in metecdysis, though the highest relative proportion occurred near ecdysis. Lobsters then were submissive and avoided physical contact with conspecifics. Clearly, locomotor activity, feeding, and social behavior of P. argus are not simply determined. Indeed, behavior is distinctly phase coordinated with varying metabolic requirements dependent on the proximity to ecdysis and ecological pressures.     

Embryogenesis of decapod crustaceans with different life history traits,feeding ecologies and habitats: a fatty acid approach

Variations in embryo size and fatty acid (FA) dynamics during embryogenesis were evaluated in deep-sea pandalids and portunid swimming crabs from the Portuguese continental margin and Madeira Island slope and compared with previous data on neritic and deep-sea lobsters and shrimps (collected between February 2001 and March 2004). Inter-specific variations in embryo size seem to be dictated primarily by phylogeny rather than by differences in reproductive or early life history traits. FA reserves were significantly correlated with embryo size (P < 0.001). Principal component analysis revealed differences among three groups (1—neritic caridean shrimps, 2—deep-sea pandalids of the genus Plesionika, and lobsters, 3—portunid crabs and the deep-sea pandalid Chlorotocus crassicornis, Costa 1871). Group 1 was clearly separated by PC1 mainly due to the higher percentage of essential C₁₈ (linoleic and linolenic acids) and C₂₀ (namely eicosapentaenoic) polyunsaturated FA (specific markers of primary producers). PC2 separated Group 2 from Group 3 due to differences in the percentage of several saturated FA (including odd-numbered FA—bacterial markers) and C₁₈ monounsaturated FA (namely 18:1n − 9, a general marker of carnivory). Therefore, these differences among groups seem to result from distinctions in diet and ecological niche. Intra-specific differences in FA composition between western and southern Plesionika martia martia (A. Milne-Edwards, 1883) populations may reflect higher water temperatures on the south sub-tropical coast. Lobster embryonic development was more demanding of lipid energy than that of the other decapod species, which may reflect an evolutionary trend in decapod taxa related to an increasing degree of lecithotrophy. However, a lower FA catabolism can be interpreted as an enhanced independence of the newly hatched larvae from external energy sources. Higher FA content at hatching and, as a consequence, a greater independence from the external environment should increase the chances of larval survival. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Presence of coelenterazine in mesopelagic fishes from the Strait of Messina

Coelenterazine, a luciferine, and luciferase activity specific to coelenterazine were detected and assayed in various tissues of mid-water fishes, Argyropelecus hemigymnus, Chauliodus sloani, Myctophum punctatum, Vinciguerria attenuata and Cyclothone braueri. Coelenterazine was found mainly in digestive systems and photophores of these fishes. A large species to species variation was found: extremely high levels of coelenterazine and luciferase were found in the M. punctatum digestive system whereas only a very low level of coelenterazine was detected in C. braueri. Coelenterazine was detected in A. hemigymnus eggs supporting the hypothesis of a maternal transfer of luminous capabilities. Luciferase activity specific to coelenterazine was found in photophores as well as in various other tissues suggesting another (besides light emission) biological function for this enzymatic activity. Distribution of coelenterazine in all tissues of the individuals supports the hypothesis of the dietary acquisition of coelenterazine by these fishes.     

Observations of copepod feeding and vertical distribution under natural turbulent conditions in the North Sea

We present results of simultaneous measurements of turbulent-dissipation rate, zooplankton vertical distribution and copepod gut pigments in the northern North Sea. Analysis shows that some, but not all, copepods (by species, sex and stage) exhibit significant dependence on turbulence in respect to vertical distribution and feeding rate. Oithona similis (female and copepodite stages) exhibits an avoidance of the surface layer when turbulence is strong there. For the range of turbulence (10⁻⁷ to 10⁻³m² s⁻³) and ambient chlorophyll concentration (0.5–0.8 μg l⁻¹) encountered, Calanus spp. and Metridia lucens exhibited a significant negative response in feeding-rate index with increasing turbulence. Centropages typicus and Pseudocalanus spp. also exhibited a negative response but of less significance. Received: 12 October 2000 / Accepted: 11 December 2000     

The permeability to oxygen of the swimbladder of the mesopelagic fish Ceratoscopelus maderensis

The permeability to oxygen of the swimbladder of physoclistous fish is important because of the usually high oxygen content of the bladder. The oxygen permeability and guanine content of the swimbladder wall of the mesopelagic myctophid Ceratoscopelus maderensis (Lowe) were measured. The oxygen permeability was 0.0672 cm³ m/atm min cm² and the guanine content was 73 g/cm². Attention is drawn to the high rates of gas loss which would occur from a swimbladder with these characteristics and the problems of maintaining the gas content, especially at depth.     

Gender- and oxygen-related irrigation behaviour of the decapod Nephrops norvegicus

The Norway lobster Nephrops norvegicus (L.) inhabits burrows in muddy clay sediments (e.g. on the Swedish west coast), where an autumnal oxygen deficiency in the bottom water can occur. Our experiments investigated whether the irrigation of the burrows would reflect a behavioural adaptation to hypoxia, and whether any gender differences of such behaviour exist. Irrigation is performed by the pleopods which may compensate for a decreasing oxygen tension. Pleopod activity (total number of strokes per sampling time), associated with oxygen concentration and gender, was studied in N. norvegicus kept in artificial burrows resembling their natural habitat. Male and female lobsters were separately exposed to either normoxia (70% oxygen saturation) or hypoxia (30% oxygen saturation). A sexual difference in behaviour was found, where females irrigated the burrow less than males during normoxia. Females showed a significant increase of pleopod activity in hypoxia compared with normoxic conditions, which was not displayed by the males probably due to the degree of individual variation found. However, when only males were studied during progressive hypoxia (from 60 to 5% oxygen saturation), following any changes of irrigational behaviour, a significant increase of accumulated pleopod activity occurred. A major increase of pleopod activity appeared between 60 and 50% oxygen saturation, below which the activity remained high until a critical point (<10% saturation, 11 °C, 33 psu) where irrigation dropped to a level close to that of normoxic values. Activity sessions during hypoxia were longer and had a higher stroke rate than during normoxia. Received: 22 October 1997 / Accepted: 26 February 1998     

Vertical distribution and migration of fishes of the lower mesopelagic zone off Oregon

An exceptionally large midwater trawl (50 m² mouth area) with 5 opening and closing codends was towed horizontally in the lower mesopelagic zone at depths of 500, 650, 800 and 1000 m off Oregon (USA) from 1–6 September, 1978. In comparison to more conventional trawls, ours collected more fish, including rare species and large individuals of common species. Comparison of collections made by day and by night revealed that 12 of the 15 most common species probably migrated vertically. Bathylagus milleri evidently migrates from 650 m during the day to 500 m at night. Cyclothone acclinidens and C. atraria were more abundant by night than by day at 800 m, possibly due to an upward migration from deeper depths at night. C. pseudopallida, C. signata, Chauliodus macouni, Tactostoma macropus and Stenobrachius leucopsarus were more abundant by day than by night at 500 m, suggesting that they migrated out of this depth horizon at night. Lampanyctus regalis, and large individuals of B. pacificus were more abundant by night than by day at 500 m, possibly because they migrated upward from near 650 m. Many species exhibited trends of increasing or decreasing size with depth, and several species showed changes in migratory behavior with size. For example, only small (<240 mm) T. macropus migrated vertically, whereas only large (>110 mm) B. pacificus appeared to migrate. Depths of maximum abundance of congeneric species were usually separated. B. milleri and B. pacificus had similar distributions by day, but the former was shallower at night. S. leucopsarus tended to live shallower than S. nannochir both day and night. Congeners always occurring at the same depth were Cyclothone pseudopallida and C. signata (both most abundant at 500 m) and C. acclinidens and C. atraria (both most abundant at 800 m).     

Distribution and diel vertical movements of mesopelagic scattering layers in the Red Sea

The mesopelagic zone of the Red Sea represents an extreme environment due to low food concentrations, high temperatures and low oxygen waters. Nevertheless, a 38 kHz echosounder identified at least four distinct scattering layers during the daytime, of which the 2 deepest layers resided entirely within the mesopelagic zone. Two of the acoustic layers were found above a mesopelagic oxygen minimum zone (OMZ), one layer overlapped with the OMZ, and one layer was found below the OMZ. Almost all organisms in the deep layers migrated to the near-surface waters during the night. Backscatter from a 300 kHz lowered Acoustic Doppler Current Profiler indicated a layer of zooplankton within the OMZ. They carried out DVM, yet a portion remained at mesopelagic depths during the night. Our acoustic measurements showed that the bulk of the acoustic backscatter was restricted to waters shallower than 800 m, suggesting that most of the biomass in the Red Sea resides above this depth.     

Proximate composition and energy content of mesopelagic fishes from the eastern Gulf of Mexico

The proximate composition of 33 species of mesopelagic fishes collected from the eastern Gulf of Mexico during May–June 1984, July–November 1985 and January 1986 was determined. Water level increased and ash-free dry weight (% dry weight) and protein level (% wet weight) decreased with increasing species' minimum depth of occurrence (that depth below which 90% of a species' population lives). Lipid level (% wet weight or % ash-free dry weight), did not change with species' minimum depth of occurrence. Skeletal ash level (% wet weight) generally decreased with minimum depth of occurrence, whereas carbohydrate level did not change with depth. The variable water level, low lipid level, and high protein level in eastern Gulf of Mexico fishes resulted in a low energy content. These results are generally characteristic of fishes from warm, stable regions of low productivity, such as the eastern North Pacific Gyre. The constant food supply provided by a stable environment may obviate the need for large lipid reserves, in contrast to colder waters at higher latitudes where food availability is seasonal. In addition, the large energy requirements for diel excursions into high-temperature surface waters by the many vertically migrating fishes of this region may influence lipid deposition. The relatively high protein level found in migrators compared to that in non-migrators or weak migrators indicates that locomotory demands influence the percentage of protein found in Gulf fishes. The lack of a positive correlation between protein level and the food availability of a zoogeographic region, suggested in previous studies, is supported here.     

Persistent near-bottom aggregations of mesopelagic animals along the North Carolina and Virginia continental slopes

Submersible observations during four missions over the North Carolina and Virginia continental slopes (184–900 m) documented the occurrence of large aggregations of mesopelagic fishes and macronektonic invertebrates near or on the bottom. Aggregated mesopelagics formed a layer up to tens of meters deep positioned from a few centimeters to 20 m, usually <10 m, above the substrate. Aggregations were numerically dominated by microvores, notably the myctophid fish Ceratoscopelus maderensis and the penaeid shrimp Sergestes arcticus. Consistently present but in relatively lower numbers, were mesopelagic predators, including the paralepidids Notolepis rissoi and Lestidium atlanticum, the eel Nemichthys scolopaceus, the stomiid fishes Chauliodus sloani and Stomias boa ferox, and squids Illex spp. Near-bottom aggregations do not appear to be an artifact due to attraction to the submersible. Based on submersible observations in three areas in 4 years spanning a decade, near-bottom aggregations of midwater organisms appear to be a geographically widespread and persistent phenomenon along the continental slope of the southeastern US Aggregations may exploit areas of enhanced food resources at the bottom.     

Rhythms of larval release of decapod crustaceans in the Mira Estuary,Portugal

Rhythms of larval release of the most abundant decapod crustaceans in the Mira Estuary, Portugal, were observed based on plankton samples made over consecutive ebbing tides during one lunar cycle in June 1986. Most species showed a semi-lunar rhythm of larval release, centered on crepuscular high tides around the quarter moons. The study suggests that the larval-releasing activity is connected to the hour of the high tide rather than to tidal amplitude.     

Species-specific predation on amphipod crustacea by the pinfish Lagodon rhomboides: Mediation by macrophyte standing crop

The amphipod species consumed by Lagodon rhomboides represented only a small subset of the amphipod assemblage available at three seagrass habitats in Apalachee Bay, Florida (USA). Predatory preferences were related most closely to the microhabitat of prey species and were unrelated to amphipod abundances. Important prey species were all epifaunal types. Consumption of preferred amphipod species was non-selective at a site with sparse macrophyte cover, but selectivity increased with macrophyte biomass. The amphipod species that were preferentially selected as prey by pinfish correspond with those that have been suggested as being limited by fish predators. It was suggested that mediation of predator behavior by physical structure in seagrass meadows may play an important role in the regulation of species richness and abundances. Species-specific identification of prey is recommended for food-habit studies.