The reproductive biology of Hymenaster membranaceus from the Rockall Trough,North-East Atlantic Ocean,with notes on H. gennaeus

Samples of the deep-sea spinulosan asteroid Hymenaster membranaceus Wyville Thomson were collected in a timeseries of 19 bottom trawls spanning the period April 1978 to October 1981 from a 2200 m-deep station in the northern Rockall Trough. The reproductive biology of this species was studied from histological sections of the gonad, and compared with that of H. gennaeus H. L. Clark which was collected in the same hauls. At first sexual development, oogonia develop in nests surrounded by small accessory cells. Previtellogenic oocytes remain in the periphery but, at maturity, oocytes ranging up to 1 100 m fill the ovary. A variety of accessory cells pack the lumen and may be nutritive or degenerative. It appears that a small number of oocytes are spawned intermittently, but there is no evidence of overproduction and break-down of superfluous oocytes. A few large oocytes become senescent and undergo internal break-down, releasing periodic acid Schiff-positive material into the lumen. Size-frequencies of oocytes indicate that eggs may be spawned as a continuous slow release, and there is no evidence of reproductive synchrony between or within samples. On reaching maturity, males appear always to be ready to release spermatozoa. Spawning is probably stimulated by egg release during chance encounters with mature females. There is no evidence for brooding, and from the large size and yolky nature of the egg direct lecithotrophic development at or near the seabed is inferred. A limited histological study of H. gennaeus indicates that egg production is very similar, but the two species differ in the nature of the accessory cells and amorphous material filling the lumen.     

Observations on the reproduction and postlarval morphology of pourtalesiid sea urchins in the Rockall Trough area (N.E. Atlantic Ocean)

Gametogenesis was studied histologically in the deep-sea species Pourtalesia jeffreysi (Wyville Thomson), P. miranda (A. Agassiz) and Echinosigra phiale (Wyville Thomson) collected during 1973–1983 from various positions at depths of 1 040 to 2 921 m in the Rockall Trough. The gonads are small and no synchrony in terms of the stage of gametogenic development was evident in females of the latter two species collected at different times of the year, indicating nonseasonal breeding. Males appear to be ripe at all times of the year. Follicle cells were well developed around the developing oocytes. Both the size range of the yolky ripe eggs (173 to 357 m diam) and potential fecundity within the range of 1 000 to 4 000 oocytes per individual, are indicative of an abbreviated, indirect, lecithotrophic mode of development occurring in all three species. Pourtalesiid postlarvae occurred also in fine-meshed trawl hauls, and their morphology is described. It was not possible to distinguish between P. miranda and E. phiale at sizes <3 mm in length.     

Growth of the deep-sea irregular sea urchins Echinosigra phiale and Hemiaster expergitus in the Rockall Trough (N.E. Atlantic Ocean)

Growth in the deep-sea irregular sea urchins Echinosigra phiale (family Pourtalesiidae) and Hemiaster expergitus (family Hemiasteridae) was studied from deep-sea samples taken during the years 1973 to 1985 from two stations at 2900 and 2200 m depth in the Rockall Trough (N.E. Atlantic Ocean). Growth zones, similar to those described from sea urchins in shallow water, are present as a series of wide white bands separated by narrow, dark rings in the calcite stereom of the test plates after heating to 350°C. In shallow water, such growth zones seem to result from seasonally varying growth rates. In the supposedly constant conditions in the deep sea, a seasonal growth pattern is unexpected but may occur in response to recently discovered annual pulses in downward flux of detritus from the euphotic zone, providing a seasonally varying food supply for such deposit-feeding species living in the bottom sediment. On this assumption, growth curves were fitted to counts of growth zones (as representing age in years), in the larger lateral and ventral test plates of E. phiale and H. expergitus. The opportunity was also taken to fit growth curves derived from counts of growth zones in samples of the inshore spatangoids Spatangus purpureus and Echinocardium pennatifidum. Plots of counts against test length of Echinosigra phiale and H. expergitus, although scattered and not clearly asymptotic, indicate, growth to be slower than in the two inshore spatangoids, and than in the coastal species Echinocardium cordatum, for which there are good recent growth data, available.     

Diets of clupeoid,stomiatoid and salmonoid fish of the Rockall Trough,northeastern Atlantic Ocean

The trophic interactions of species of fish of the continental slopes have not been investigated previously in detail. The present study examines the diets of the clupeoid, stomiatoid and salmonoid species occurring demersally and pelagically in the Rockall Trough to the west of Scotland and Ireland. Pelagic fish were collected between the surface and about 2 500-m depth between 1973 and 1978. Demersal fish were sampled at 250-m intervals of depths between 500- and 2 900-m depth during the years 1975 and 1981. Of the 28 species caught, 18 were strictly pelagic and dominated by the stomiatoids while five were strictly demersal. The remaining five species occurred in both the pelagic and demersal environment. The commonest clupeoid was Alepocephalus bairdii, which is a dominant species within the demersal fish associations at depths of 750 to 1 250m. This species, along with the deeper-living A. agassizi, and 3 other alepocephalid species feed primarily on benthopelagic prey but also exploit the epibenthos. The stomiatoid species, such as Cyclothone microdon, C. braueri, Maurolicus muelleri and Argyropelecus hemigymnus, dominate the associations of pelagic fish in the Trough. They feed on different proportions of ostracods, copepods and amphipods. Of the salmonoids, only Argentina silus and Bathylagus euryops occurred commonly and both feed on benthopelagic fauna, the latter at much greater depths than the former. The principal factor acting to prevent direct competition between species is the modal centres of bathymetric distribution of the species. Other contributing factors are briefly discussed.     

Diets and bathymetric distributions of the macrourid fish of the Rockall Trough,northeastern Atlantic Ocean

The Macrouridae are the most common fish caught in demersal trawls on the continental slope and rise of the Rockall Trough. They represented 41% of all fish caught, with Coryphaenoides rupestris amounting to 28% of the catch. No previous study of the trophic interaction of these fish has been made over a wide bathymetric range. Samples were obtained at 250-m intervals of depth between 400 and 2 900 m in the period 1975 to 1981. The stomachs of 5 326 fish belonging to 12 species were examined to define their diets. Eight species are primarily benthopelagic feeders while four are primarily epibenthic feeders. The bathymetric centres of distribution of the populations of the benthopelagic feeding Trachyrhynchus murrayi, Malacocephalus laevis, Coryphaenoides rupestis, C. guentheri, Nematonurus armatus, Chalinura brevibarbis, C. leptolepis and C. mediterranea are different from each other. Similar differences in bathymetric distribution occurred among the epibenthic feeding Nezumia aequalis, Coelorhynchus coelorhynchus, C. occa and Lionurus carapinus. These differences decrease competition among species that exploit similar resources.     

Growth and breeding of meso- and bathypelagic organisms of the Rockall Trough,northeastern Atlantic Ocean and evidence of seasonality

Seasonality of growth and reproduction of a variety of epimeso-and bathypelagic organisms living in the Rockall Trough is examined in samples collected between 1973 and 1978. Rates of growth change seasonally. Epipelagic and shallow-living mesopelagic migrators tend to grow most rapidly in the spring and summer while deeper-living mesopelagic migrators tend to accelerate their growth in the summer and autumn when seasonal water temperatures at these depths are at their seasonal maxima. Mesopelagic migrators all breed seasonally. Rates of growth of bathypelagic non-migrating species could not be determined through analyses of body length/frequency histograms because of the continuous (aseasonal) breeding and recruitment of juveniles to the adult populations. An exception was the mysid crustacean Boreomysis microps that had a seasonal period of spring recruitment of juveniles to the population. Several of the mesopelagic crustaceans and fish have extended periods of approximately linear growth in body length. There is no evidence that organisms living in the mesopelagic or bathypelagic environments have rates of growth that are markedly slower than those of congeners in shallower environments.     

Deep-sea bottom-living fishes at two repeat stations at 2 200 and 2 900 m in the Rockall Trough,northeastern Atlantic Ocean

Repeated sampling of deep-sea bottom-living fishes was conducted at two stations in the Rockall Trough at depths of 2 200 and 2 900 m. The 2 200 m station (M) was sampled 16 times by an Agassiz trawl between April 1978 and April 1985 and yielded 473 fish belonging to 17 species. The 2 900 m permanent station (PS) yielded 781 fish belonging to 11 species from 9 Agassiz trawls, 2 small box otter trawls, 4 semi-balloon otter trawls and 7 epibenthic sledges between March 1975 and April 1985. Macrourid fishes were numerically dominant at both stations with Coryphaenoides guentheri and Coryphaenoides (Nematonurus) armatus being the most abundant species at M and PS, respectively. In terms of biomass the morid, Antimora rostrata, was dominant at M but C. (N.) armatus was dominant at PS. The only species that showed clear evidence of a seasonal reproductive cycle was C. guentheri. Many species showed no indication of reproducing in the Rockall Trough. The diets of almost all the species are described and considered in relation to the food resources exploited. Investigations on the pelagic and benthic invertebrate faunas at these stations should reveal the extent to which the bottom-living fishes exploit the available resources.     

Growth and production of Euphausiacea (Crustacea) in the Rockall Trough

The euphausiid fauna of the Rockall Trough has been sampled in time series at approximately two-monthly intervals between 1973 and 1976 between surface and depths greater than 2000 m. Length/frequency histograms have modes that are coherent for periods of at least 4 to 6 mo, indicating that populations are being sampled. Rates of growth were determined for six species. Epipelagic species have winter checks to the growth rates, but such checks are absent in the growth of Thysanopoda acutifrons, Nematobranchion boöpis and Stylocheiron maximum. Ratios of annual production: biomass were determined and are all within the previously determined range of values for epipelagic and coastal populations. Lack of a winter check to the growth rate in deep-living species enables these potentially to have overall rates of growth comparable to those of temperate, epipelagic, shelf and coastal species.     

Multi-marker estimate of genetic connectivity of sole (Solea solea) in the North-East Atlantic Ocean

A thorough knowledge on the genetic connectivity of marine populations is important for fisheries management and conservation. Using a dense population sampling design and two types of neutral molecular markers (10 nuclear microsatellite loci and a mtDNA cytochrome b fragment), we inferred the genetic connectivity among the main known spawning grounds of sole (Solea solea L.) in the North-East Atlantic Ocean. The results revealed a clear genetic structure for sole in the North-East Atlantic Ocean with at least three different populations, namely the Kattegat/Skagerrak region, the North Sea and the Bay of Biscay, and with indications for a fourth population, namely the Irish/Celtic Sea. The lack of genetically meaningful differences between biological populations within the southern North Sea is likely due to a large effective population size and sufficient connection (gene flow) between populations. Nevertheless, an isolation-by-distance pattern was found based on microsatellite genotyping, while no such pattern was observed with the cytochrome b marker, indicating an historical pattern prevailing in the latter marker. Our results demonstrate the importance of a combined multi-marker approach to understand the connectivity among marine populations at region scales.     

Reproductive strategies and diet in deep-sea nuculanid protobranchs (Bivalvia: Nuculoidea) from the Rockall Trough

Although the taxonomy of deep-sea protobranch bivalves is becoming better known, relatively little information is available on their reproductive biology and whether or not populations show reproductive periodicities. We have examined the reproductive biology of three common sympatric species as part of a long-term time-series of samples taken from 2900 m in the Rockall Trough from 1973 to 1983. Malletia cuneata Jeffreys, 1876 produces a maximum of 30 oocytes at any one time and these grow to a maximum size of 240 m. Maximum fecundity of Ledella pustulosa (Jeffreys, 1876) and Yoldiella jeffreysi (Hidalgo, 1877) is 174 and 360, respectively, and both species produce an egg of 120 m in diameter. These data indicate lecithotrophic early development in L. pustulosa and Y. jeffreysi, but direct development in M. cuneata; however, evidence from the prodissoconch length of M. cuneata suggests lecithotrophic development. L. pustulosa and Y. jeffreysi also differ from M. cuneata in having a distinct reproductive cycle with spawnout in the winter months. Both the continuously breeding M. cuneata and the seasonally breeding L. pustulosa ingest diatoms, coccoliths and foraminiferans, but whereas the diet of M. cuneata appears to be constant throughout the year there is an apparent reduction in the feeding activity of L. pustulosa concomitant with the deposition of phytodetritus on the deep-sea bed.     

Life-history biology of Peniagone azorica and P. diaphana (Echinodermata: Holothurioidea) from the north-east Atlantic Ocean

Aspects of the reproduction and population biology of two elasipodid holothurians collected during the period 1973–1984 from the north-east Atlantic Ocean were examined. The reproductive biology of both the benthic species Peniagone azorica and the benthopelagic P. diaphana are similar. A primary oocyte forms from an oogonium and grows to about 100m diameter, whereupon it undergoes vitellogenesis and increases to 300m before being spawned. A high proportion of primary oocytes are not spawned and undergo a complicated breakdown process resulting in the formation of an amorphous sac in the ovary wall. It is possible that when an ovarian tubule is full of these sacs it atrophies and drops off, allowing other tubules to develop. The maximum egg size for both species suggests abbreviated larval development. In P. azorica, juveniles reach sexual maturity for the first time at about 30 mm length. The population structure suggests that the adults grow slowly, although it is possible that the observed unimodal distribution in size-frequencies results from one or several recruitments. Recruitment to the population is probably infrequent and may occur irregularly. Comparison of the population structure of closely spaced samples suggests a patchy distribution on the bottom. The benthopelagic lifestyle of adult P. diaphana suggests that the larvais also planktonic.     

Reproductive cycle of Zidona dufresnei (Caenogastropoda: Volutidae) from the southwestern Atlantic Ocean

Gonadal development of Zidona dufresnei (Donovan, 1823) (Caenogastropoda: Volutidae) was studied over a period of two consecutive years, through analysis of gonadal tissue. Individuals were sampled monthly at the Mar del Plata area, Argentina. Gonadosomatic index was estimated for males and oocyte size was used to estimate the stage of gonadal development in females. The reproductive season in this species in the sampled locality extended from October to March (austral spring-summer). During summer, a stage of advanced gonadal development and spawning predominated in the adult population. In autumn, gonads were generally under atresia; during wintertime (June-August), they underwent a period of recovery that lasted until the spring months, when gametes were released again. Synchronism between both sexes was evident. Marked periods of spawning were followed by resorption periods and then a growing phase; it was very clear that reproductive seasonality was linked to changes in bottom water temperature. These results suggest that Z. dufresnei gonads have a yearly cycle of gamete production, with two major activity peaks in September-October and January-February.     

Food habits and intestinal parasites of deep demersal fishes from the upper continental slope east of Newfoundland,northwest Atlantic Ocean

Stomach contents and intestinal parasite faunas of 471 individuals of demersal fishes in 14 species were examined from the Carson Canyon region (Lat. 45°30N; Long. 48°40W) of the upper continental slope of the Grand Banks off Newfoundland, Canada. Individual species tended to feed either on benthic or on pelagic/benthopelagic organisms, but pelagic prey assumed the greatest importance overall. Data from stomach contents were supported by the parasite information. Prevalence of parasites was higher in benthic feeders (53.1%) than in pelagic feeders (28.9%), and relative abundance by major group was: Digenetic Trematoda 5.8% benthic vs 27.8% pelagic, Nematoda 53.1% vs 72.2%, and Acanthocephala 40.9% vs 0%. Of the dominant fishes, there were more species of benthic feeders (5) than pelagic feeders (3), but pelagic feeders were numerically more abundant (pelagic 70.9%, benthic 20.5%). Benthic feeders were on average larger (=270.6g) than pelagic ones (=130.6g), but pelagic feeders represented a larger proportion of the biomass (pelagic 43.3%, benthic 25.9%). The results of this study combined with those from other areas suggest that feeding from the pelagial by demersal fishes at upper continental slope depths is probably the general rule.     

Interactions among endocrinology,seasonal reproductive cycles and the nesting biology of the female green sea turtle

We collected data on plasma levels of testosterone+5a-dihydrotestosterone (T+DHT) and corticosterone (CORT) from adult female green sea turtles (Chelonia mydas) from southern Queensland during distinct stages of their reproductive cycle. Those females capable of breeding in a given year had elevated plasma steroid levels (T+DHT 0.91ǂ.08; CORT 1.05ǂ.29 ng/ml), associated with follicular development, until courtship began in October. At the beginning of the nesting season in November plasma levels of CORT were related to when the female first nested (r²=0.06; F=10.45; P=0.01). However, they were not correlated with the number of clutches a female laid in that season (F=3.65; P=0.08). We repeatedly sampled 23 turtles over the nesting season and profiled changes in steroids immediately following oviposition of each clutch. Levels of T+DHT (range 0.41-0.58 ng/ml) and CORT (range 2.13-2.81 ng/ml) were similar through the early stages of the nesting season and inter-nesting period, and declined to near basal levels (T+DHT 0.37ǂ.03 and CORT 1.85-ng/ml) following the last clutch for the season. Steroid hormone levels were also low (T+DHT 0.38ǂ.16; CORT 0.46ǂ.21 ng/ml) in four independent post-breeding (atretic) females; samples for these females were taken at a time when body condition was presumably at the lowest for the season. Subtle changes in the nesting environment, such as variation in nesting habitat or the time of night that nesting occurred, were associated with a small and slow CORT increase. We suggest CORT is increased in nesting females to assist in lipid transfer to prepare the ovarian follicles and/or the reproductive organs for ovulation.     

Genetic variance in Ophiomusium lymani (Ophiuroidea: Echinodermata) from lower bathyal depths in the Rocklall Trough (northeast Atlantic)

Specimens of the deep-sea brittle-star Ophiomusium lymani were collected from six sites in the Rockall Trough (northeast Atlantic_. Four monomorphic and four polymorphic loci were detected, with up to 75 individuals screened at any one locus. The results showed little difference in allele frequencies between sites and, consequently, estimates of genetic identity indicated no significant genetic differentiation between the populations sampled. Deficits of heterozygotes were observed at all polymorphic loci, but the deficiencies were only significant at the phosphoglucose isomerase locus from two locations. The deficit of heterozygotes was not the same across loci, suggesting that inbreeding is not the cause of the excess homozygosity. No relationship between heterozygosity and depth was observed.     

Interactions between sea urchins (Strongylocentrotus droebachiensis) and their predators in field and laboratory experiments

Field observations and manipulative experiments in a nearshore cobble bed (2 to 3 m below mean low water) at Eagle Head, Nova Scotia, Canada, between 1984 and 1986, showed that small juveniles ofStrongylocentrotus droebachiensis (3 to 6 mm diam) sheltering beneath cobbles had a refuge from predators such as rock crabs, small lobsters, and fish. Sea urchins gradually outgrew these refuges and small adults (25 to 30 mm) required larger rocks as shelter from predators, particularly large cancrid crabs. Small juveniles were usually solitary and well dispersed beneath cobbles, whereas small adults tended to aggregate on the undersides and in the interstices of boulders. These aggregations may develop passively as sea urchins accumulate in suitablysized refuges. Chemotaxis experiments indicate that juvenileS. droebachiensis are repelled by waterborne stimuli from conspecifics. In a factorial experiment, effects of the presence of potential predators (rock crabs and lobsters) and/or food (kelp) on the behaviour of large juvenile (10 to 15 mm) and small adult sea urchins were examined in flowing seawater tanks. Both size classes formed exposed feeding aggregations when kelp was provided as food, irrespective of the presence or absence of predators. In the absence of kelp, each size class responded differently to the presence of a predator: juveniles became more cryptic, whereas adults aggregated on the tank sides. Increased movement to the sides of a tank in the presence of a predator may reflect a flight response, since chemotaxis experiments indicated thatS. droebachiensis is repelled by waterborne chemical stimuli from predators. Observational and experimental data suggest that predation at the late juvenile and early adult stages may influence population structure, distribution and abundance ofS. droebachiensis.     

The occurrence of wax esters in crustaceans from the North-east Atlantic Ocean

The lipids of 2 species and 1 group of copepods, 4 species of euphausiids, 2 species of mysids, and 6 species of decapods, taken from different depths in the North-eastern Atlantic Ocean and Western Mediterranean Sea, have been analysed specifically for their wax ester content. The mid-water animals living below 500 m contained large amounts of wax ester (30 to 80% total lipid), while those animals living in the upper 600 to 700 m or on the sea floor had only small wax ester fractions (5 to 10% total lipid). It is suggested that the wax esters are laid down by those animals living in areas of low food density, where energy conservation is important, partly to achieve a more neutral buoyancy. In addition, it would appear that wax esters can function as energy reserves.