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.     

Relationships between the movements,growth, age structures,and reproductive biology of the teleosts Sillago burrus and S. vittata in temperate marine waters

Sillago burrus and S. vittata both use sheltered, nearshore shallow waters ( 1.5 m) as nursery areas. However, the juveniles of the former species remain there for only a few months, before migrating into deeper waters (5 to 15 m) as they increase in size, whereas some juvenile S. vittata do not undergo a similar migration until considerably later. S. burrus rarely exceeded 2 yr of age and was never found beyond 4 yr of age. Although only a small number of S. vittata exceeded 2 yr of age, a few individuals of this species were caught between 4 and 7 yr old. The maximum and asymptotic lengths of S. burrus (251 and 180 mm, respectively) were far lower than those of S. vittata (325 and 320 mm, respectively), whereas the growth coefficients (K) were much higher for the former species, i.e. 2.4 vs 0.4. Virtually all S. burrus, and also those S. vittata that moved into deeper waters early in life, spawned at the end of their first year of life. Since relatively few S. burrus reached 2 yr of age, the attainment of almost full size by the end of their first year of life enables a relatively large number of eggs to be produced by fish at the end of their first year-for many, their only spawning period. Those individuals of S. vittata that remained in their shallow nursery areas until the end of their first year of life, did not reach maturity until the end of their second year of life. The proportions of mature gonads and the numbers of yolk-vesicle and yolk-granule oocytes and post-ovulatory follicles in ovaries were far higher in both S. burrus and S. vittata during December to February than in any other month, demonstrating that these two species spawn largely in these summer months. During this period, the ovaries of individual S. burrus and S. vittata often contained post-ovulatory follicles, as well as yolkvesicle and yolk-granule oocytes that ranged widely in size, strongly suggesting that both species are multiple spawners. During the spawning period of S. burrus, the ovaries possessed large numbers of hydrated oocytes and no post-ovulatory follicles or the reverse situation, and the oocytes tended to form several relatively discrete size groups. This indicates that S. burrus produces eggs in batches and that the spawning of the members of this species is synchronised. The presence of large numbers of yolk-granule oocytes with migrating nuclei in the ovaries of many S. vittata at certain times suggests that this species is also a batch- and synchronised spawner. Comparisons between the results of the present study and past work emphasise that the relationships between the timing of offshore movements and the sizes and ages at first maturity vary considerably amongst whiting species.     

Diets of some deep-sea brittle stars in the Rockall Trough

The stomachs of 1 165 individuals of the species Ophiacantha bidentata (Retzius), Ophiactis abyssicola (M. Sars), Ophiocten gracilis (O. Sars), Ophiura irrorata (Lyman), O. ljungmani (Lyman) and Ophiomusium lymani Wyville Thomson, collected from the general area of the Rockall Trough from 1973 to 1983, were examined; 47% contained material. This varied in nature and in volume between individuals, but overall, the diets of the 6 species showed quite high similarity. The few differences evident are interpreted as reflecting different lifestyles. Except for Ophiocten gracilis, the probably motile, epifaunal species (Ophiura irrorata, O. ljungmani and Ophiomusium lymani) showed the greatest variety in items, and the lowest similarity with the remaining species; they are probably unselective omnivores, eating whatever small prey or organic detritus they are able to find. Ophiacantha bidentata and Ophiactis abyssicola showed both the least dietary variety, and the highest similarity to each other. This probably reflects their more sedentary lifestyle, feeding on current-borne particles and small prey from perches on sessile fauna. The stomach contents of Ophiocten gracilis consisted mainly of amorphous organic material and mineral particles, indicating a microphagous feeding mechanism. With O. gracilis, the proportion of stomachs with food decreased markedly in summer, this coinciding with seasonal gonadal development when stomachs frequently appeared ruptured or compressed. The possible importance of seasonal sedimentation of fast-sinking particulates from the surface is discussed in relation to finds of flocculent organic material, often containing diatom frustules, in stomachs of all 6 species. Some doubts remain as to whether our results are biased by specimens egesting stomach contents on capture. As in shallow water, deep-sea ophiuroids seem to be trophic generalists lacking in dietary specialization. Brief notes on parasites encountered are also given.     

Reproductive strategies of two deep-sea gastropod species from the Porcupine Seabight (Northeast Atlantic)

Two species of small gastropods (<6 mm in length), Amphissa acutecostata (Philippi, 1844) and Gymnobela subaraneosa (Dautzenberg and Fischer, 1896), widely distributed in the northeast Atlantic, were found in large numbers in the Porcupine Seabight (Northeast Atlantic). Except for some aspects of taxonomy and distribution, as well as some data on larval development, the biology of these species is unknown. This study describes basic aspects of the life-history strategies of both species. Histological studies showed that oocyte and sperm development in both species was similar to the gametogenetic patterns observed in other deep-sea gastropods. In females, oogonia proliferated in the germinal epithelium and developed into previtellogenic oocytes (30–40 m), which grew into vitellogenic primary oocytes. Vitellogenic oocytes were covered by a thin layer of follicle cells involved in the vitellogenic processes. The maximum size for mature oocytes was 99.06 m for A. acutecostata and 114.82 m for G. subaraneosa. In A. acutecostata most of the volume of the ovary was occupied by previtellogenic and early vitellogenic oocytes, whereas in G. subaraneosa most of the volume was filled by large vitellogenic oocytes. Both species showed quasi-continuous production of oocytes. The oocyte size-frequency diagrams suggested a continuous release of a small number of oocytes throughout the year for A. acutecostata, and asynchronous periodic spawning events for G. subaraneosa. Gonad development and gametogenesis could be strongly affected by presence of parasites in one of the populations of G. subaraneosa.Communicated by J.P. Thorpe, Port Erin     

Resource use by five sympatric parrotfishes in the San Blas Archipelago,Panama

Resource use by five sympatric species of parrotfish was quantified in the San Blas Archipelago of the Republic of Panama from March to August 1987. Detailed observations of parrotfishes on patch reefs and surrounding seagrass beds showed that they partition resources with respect to habitat, food and size, but not time. Although parrotfishes shared resources, the proportions of each resource used differed significantly among species. Scarus iserti (Bloch) scraped filamentous microalgae that grew from eroded coral pavement on lower slopes of patch reefs and in halos, the area of sparse vegetation surrounding reefs. Sparisoma viride (Bonnaterre) foraged on upper slopes of patch reefs where they mostly took bites from dead coral and associated algae. S. aurofrenatum (Cuvier and Valenciennes) had the broadest diet, which consisted mostly of seagrasses and macro- and microalgae that were attached to dead coral on lower reef slopes and in halos. Although S. chrysopterum (Bloch and Schneider) commonly occurred on patch reefs, it primarily foraged in seagrass beds that surround them. S. rubripinne (Cuvier and Valenciennes) was distributed most widely, ranging from seagrass beds to reef crests, where it took bites from seagrasses, dead coral and macroalgae. Juveniles of all species occurred on lower slopes or in halos where they scraped filamentous microalgae from coral pavement. As they matured, parrotfishes moved into other habitats changing access to different types of food. All of these parrotfishes fed throughout the daytime, and resource use did not differ between morning and afternoon.     

Silicon and the ecology of marine plankton diatoms. II. Silicate-uptake kinetics in five diatom species

The variation of the rate of silicate uptake with varying silicate concentration in the medium was investigated in short-term experiments with the following marine diatom species:Skeletonema costatum, Thalassiosira pseudonana, T. decipiens, Ditylum brightwellii, andLicmophora sp. The uptake conformed to Michaelis-Menten kinetics only after a correction had been made for reactive silicate that apparently could not be utilized by the diatoms. The magnitude of this correction was in the range of 0.3 to 1.3 g-at Si/l. Mean values of the half-saturation constant of silicate uptake were calculated for the different species. The lowest value was found inS. costatum (0.80 g-at Si/l) and the highest inT. decipiens (3.37 g-at Si/l). Growth limitation by low silicate concentrations could be a cause of species succession in marine plankton-diatom blooms.     

Corophium multisetosum (Amphipoda: Corophiidae) in Canal de Mira,Portugal: Some factors that affect its distribution

The population ofCorophium multisetosum (L.) from Canal de Mira, Ria de Aveiro, Portugal, was sampled seasonally from December 1985 to September 1986, as part of a larger survey of the benthic invertebrate macrofauna. Its distribution along the channel exposes the species to a range of salinities from freshwater to above 30. A principal components analysis, using the physical and chemical parameters of the sediment as variables and sampling stations as operational taxonomic units, indicated that abundance is negatively correlated with salinity, depth, and the occurrence of sediments rich in particles below 125µm and rich in organic matter. Abundance is positively correlated with temperature. The distribution of the species does not seem to be affected by the occurrence of sediment grades between 125 and 1 000µm. Paired-choice salinity experiments indicated thatC. multisetosum prefers salinities within the range 2.5 to 10. In multi-choice experiments concerning sediment grade, amphipods did not show any significant preference within the 125 to 500µm range, although the 125µm grade was chosen less frequently. The influence of temperature on the overall distribution ofC. multisetosum is discussed.     

Zooplankton of the Bristol Channel and Severn Estuary. The distribution of four copepods in relation to salinity

The seasonal variations in distribution and abundance of the common zooplankton species in the Bristol Channel and Severn Estuary were related to the salinity regimes observed over the period November 1973 to February 1975. The dominant constituents in all regions were the calanoid copepods, which reached maximum densities in July: approximately 100 times their winter levels. Four zooplankton assemblages were recognised using an objective classification program which computed similarity coefficients and used group-average sorting. The assemblages existed along the salinity gradient observed from the Severn Estuary to the Celtic Sea. The assemblages were classified as true estuarine, estuarine and marine, euryhaline marine and stenohaline marine and were characterized by the copepods Eurytemora affinis (Poppe) (<30S), Acartia bifilosa var. inermis (rose) (27 to 33.5S), Centropages hamatus (Lilljeborg) (31 to 35S) and Calanus helgolandicus (Claus) (>33S), respectively.     

Macrozoobenthic community structure in the Ría de Foz,an intertidal estuary (Galicia,Northwest Spain)

The community structure of the macrobenthic (>1 mm) fauna of an intertidal soft-bottom estuary, the Ría de Foz (Galicia, Northwest Spain) was investigated on 99 occasions during June, September and December 1984 and March 1985. The infaunal data were subjected to classification and ordination techniques. The analyses indicated that sediment characteristics (grain size, percentage silt-clay and organic-matter content) and tidal height are the most important factors governing the distribution and abundance of the intertidal communities. Three faunal assemblages were identified and examined in terms of species dominance, constancy and fidelity. In the clean sands of the outer estuary two main assemblages were distinguished; one, corresponding to the boreal LusitanianTellina community, inhabited the middle and lower tidal levels; the other, typical of a biocoenosis of sands and slightly muddy mediolittoral sands, inhabited the upper tidal level. The third benthic assemblage occupied the muddy sands and mud of the inner estuary and displayed some features in common with theCardium edule-Scrobicularia community described by Thorson.     

Long-term changes in reproductive patterns of the holothurian<Emphasis Type="Italic"> Oneirophanta mutabilis</Emphasis> from the Porcupine Abyssal Plain

Time-series studies (1989–2002) in the NE Atlantic have shown large-scale changes in the composition and structure of the benthic community on the Porcupine Abyssal Plain. Radical changes in the abundance of some species in 1996 led to a significant shift in the way in which organic matter was reprocessed at the seabed. This article examines the reproductive processes of the holothurian Oneirophanta mutabilis collected during the time series. The reproductive biology of O. mutabilis is reviewed. No males were evident in any of the samples. The sex ratio (females: no gender) was biased significantly towards no-gender individuals. The maximum egg size was 650 m and there was no evidence of synchrony in reproduction. Significant changes in the oocyte-size distribution and the fecundity of O. mutabilis were noted through time, coinciding with the time of greatest faunal change in the benthic community. There was an increase in the proportion of previtellogenic oocytes and a decrease in the proportion of mature vitellogenic oocytes in 1997 and 1998, in parallel with a significant decrease in fecundity of the post-1996 samples. Samples from 2002 showed a reversal in the reproductive trends, with an increase in the proportion of mature vitellogenic oocytes and fecundity. The results are discussed in relation to large changes in abundance of the epibenthos on the Porcupine Abyssal Plain. It is suggested that the superabundance of certain megafaunal species on the abyssal seafloor affected the availability of trophic resources for O. mutabilis, leading to the changes in its reproductive effort.     

Reproductive biology of the common antarctic crinoid Promachocrinus kerguelensis (Echinodermata: Crinoidea)

Histological examinations of the gonads of Promachocrinus kerguelensis Carpenter, 1888 collected in November 1984 from McMurdo Sound, Antarctica, revealed ripe ovaries containing three distinct size classes of oocytes: 20 to 90, 91 to 150 and 151 to 180 m diameter as well as ova. Testes were ripe, and filled with mature sperms. There was marked inter-pinnule and inter-individual synchrony of gametogenesis and spawning is probably synchronous, at least within the sub-population sampled. The number of eggs per pinnule is allometrically related to body size. Body weight and arm length are positively correlated with total egg number. Annual fecundity was estimated to be 29 000 eggs (full-grown oocytes and ova) for a crinoid weighing 53.6 g wet weight with an arm length of 16.3 cm. The eggs float. Female genital pinnules contain more lipid and energy than male pinnules. Reproductive output and effort (23.4 to 65.2 kJ and 26 to 38%, respectively) increase with body size and are similar to values reported for other echinoderms with lecithotrophic larvae. The greatest overall energetic contribution is represented by the arms and genital pinnules. Spawning in P. kerguelensis probably occurs in November and December within McMurdo Sound. If development is slow, as in all other antarctic echinoderms studied to date, then settlement most likely occurs 2 to 3 mo later, after the short summer period of high phytoplankton productivity. High fecundity, a pelagic mode of reproduction, and the high probability of a slow rate of development may contribute to the wide geographic distribution of this common antarctic crinoid.     

Potential effects of a resuspension event on the vertical distribution of copepod eggs in the sea bed: a laboratory simulation

Adult female Acartia tonsa and Labidocera aestiva were obtained from inshore marine waters near Turkey Point, Florida (29°54.5N;84°31W) during April and May 1991. For each species, eggs spawned overnight in the laboratory were collected, mixed with sediment, and added to a plastic tube (26 to 29 cm longx5 cm diam) to yield a 2 cm-thick layer. This tube was connected to a 245 cm-long tube, seawater was added and the contents were thoroughly mixed by turning the assembly end-over-end several times. The vertical distribution of eggs in the sediments following resuspension varied according to the grain size of the sediments. For sediments of 125 m particle-diameter, 68 to 72%, and 70 to 73% of the eggs of A. tonsa and L. aestiva, respectively, occurred in the upper 4 mm.For sediments 63 to <125 m particle-diameter, 34 to 36% of the eggs of A. tonsa and L. aestiva, respectively, occurred in the upper 4 mm. For sediments of <63 m particle-diameter, egg distribution was uniform for both species throughout the 2 cm layer. The results indicate that the vertical distribution of eggs in the sea bed following a resuspension event should be highly dependent upon the grain-size composition of the sediments.     

Dietary effects on oocyte yolk-composition inCapitella sp. I (Annelida: Polychaeta): Fatty acids and sterols

In 1988, immatureCapitella sp. I (initial biomass 80µg dry wt) were raised on four diets: Gerber cereal, TetraMin fish food,Ulva sp., and benthic diatoms. After 2 wk of culturing different populations on these diets, eggs were dissected from gravid females, frozen and analyzed for fatty acid and sterol composition. Eggs produced by worms on different food types were discriminated by fatty acid composition in a principal component analysis (PCA), with 18:1⁹, 18:2, 20:5 and 20:4 fatty acids showing the greatest differences between experimental groups. The sterol profiles of all eggs were dominated by cholesterol (cholest-5-en-3-ol) and cholest-5,24-dien-3-ol (>60%). A PCA of egg sterols discriminated between adult diets with cholesterol, 23,24-dimethyl-cholesta-5-en-3-ol, cholest-5,24-dien-3-ol and a C-29 stenol showing the greatest differences. In field populations ofCapitella sp. I, oocytes produced at different times of the year may have different levels of lipids depending on their dietary availability during vitellogenesis. Variations in oocyte composition may influence larval growth and development and thus have an impact on population dynamics. Alternatively, variations in fatty acid and sterol composition of oocytes may allow us to identify the food resources utilized by this species.     

Brood reduction in the American white pelican (Pelecanus erythrohynchos)

Summary American white pelicans (Pelecanus erythrorhynchos) breeding in colonies at East Shoal Lake, Manitoba, Canada exhibited a mean hatching asynchrony of 2.5 days in 2-egg clutches. This resulted in a size difference between chicks which facilitated sibling dominance, harassment and lack of food for the subordinate chick. Only one young survived per nest. In marked broods, the secondhatched chick survived in 20% of successful nests. Manipulated clutch sizes (1, 2 and 3 eggs or chicks per nest) revealed that the presence of a second chick contributes significantly to the reproductive success of the parents. Results support the hypothesis that the second egg functions as a form of insurance against early loss of the first egg or chick. The parents, by establishing hatching asynchrony, by nonintervention in sibling aggression, and by selectively feeding the dominant chick, maximize their chance of rearing the most viable young.     

Does displacement of spawning time occur in the sibling species Cerastoderma edule and C. lamarcki?

The spawning cycles of the sibling species Cerastoderma edule (L.) and C. lamarcki (Reeve) were studied in allopatric and sympatric Danish populations from 1978–1980. The results do not support earlier findings of reproductive character displacement in sympatric populations of the two species. New records of hybrid formation and intermediate forms are presented and discussed.     

Allozyme and colour differences between two sibling species of the heteronemertean Lineus torquatus from the Sea of Japan

Brown and cherry colour morphs of the heteronemertean Lineus torquatus Coe, 1901 from the Vostok Bay of the Sea of Japan were compared using allele frequencies at 18 isozyme loci. These morphs shared no common alleles at 9 of the 18 loci. The observed genetic differences are strong evidence that the brown and cherry colour morphs, which occur sympatrically, are different species. Nei's genetic distance between these species (D=1.135) is considerably higher than typical D values for congeneric animal species. As with much previous work on nemerteans, intraspecific allozymic variability in both species was high, with mean expected heterozygosity values 0.286±0.049 and 0.159±0.051 for L. torquatus (brown) and L. torquatus (cherry) species, respectively.     

Skeletal growth markers in the deep-sea brittle starsOphiura ljungmani andOphiomusium lymani

Growth bands have been found in the calcitic vertebral arm ossicles of the commonly occurring deep-sea brittle starsOphiura ljungmani Wyville Thomson andOphiomusium lymani (Lyman) (Echinodermata: Ophiuroidea) trawled from the Rockall Trough (N.E. Atlantic) at 2 200 and 2 900 m depth from 1973 to 1982. InOphiura ljungmani, the study of ossicle microstructure by SEM shows that growth bands reflect differences in stereom porosity and surface relief, similar to that previously found amongst shallow-water brittle stars. The pattern inOphiomusium lymani was much less clear from the microstructure, but could be revealed by heating the ossicle to 450 °C and clearing in xylene. The bands showed up as fine, translucent rings separated by more opaque, wider zones, perhaps reflecting differences in organic material incorporated within the calcite. Both growth-banding patterns probably reflect an annual cycle in skeletal growth rate. On this assumption, the number and spacing of the banding in the two species indicates contrasting growth strategies. WithOphiura ljungmani, a rather regular annual growth increment and perhaps shorter lifespan (up to ca. 10 yr) thanOphiomusium lymani is indicated. The latter shows a relatively wide spacing of early bands, followed by tight clustering of the outermost bands corresponding to adult sizes. This growth pattern is characteristic of species escaping from predation by rapid growth to relatively large adult size. AdultO. lymani probably grow slowly, some perhaps reaching 20 yr of age. Growth curves were fitted to size-at-age corresponding to measurements of the size and ordering of growth bands. These corroborate age structure previously estimated from analysis of size frequencies in time series from the stations sampled in the present study. Such skeletal growth markers should be of value in analysis of the demography of deep-sea populations.     

New morphological characteristic of Xyrichtys novacula (Pisces: Labridae) from the Adriatic Sea

In 1983 curred horns were detected in the rare Adriatic Sea fish Xyrichtys novacula (Linnaeus, 1758). Their dimensions and histological structures have been investigated. This observation, compared with previous occurrences, suggested the existence of heteromorphism within this species.     

Vertical distribution and feeding patterns of midwater fish in the central equatorial Atlantic II. Sternoptychidae

The vertical distribution of seven sternoptychid species was examined from RMT 1+8 samples collected aboard R. V. Meteor in March-April 1979 and from Royal Research Ship R.R.S. Discovery in July 1974 in the central equatorial Atlantic. During daytime sternoptychids occupied depths between 200 and 1250 m, with Sternoptyx pseudobscura living deepest, centering between 800 and 900 m, and Argyropelecus sladeni most shallow, aggregating predominantly at 300 and 400 m. They are all considered limited or partial migrants, ascending only some 100 and 200 m towards the surface at night. Only A. sladeni was observed to enter the epipelagic zone (0 and 200 m).-Feeding patterns were investigated from stomach content analyses of Sternoptyx diaphana, S. pseudobscura, Argyropelecus sladeni and A. affinis. Additional stomach contents were analysed from samples of S. diaphana, A. hemigymnus and A. olfersi collected in June 1985 from F.R.V. Walther Herwig in the temperate NE Atlantic at 46°N, 17°W by means of the Engel Trawl. The food spectrum of the six species is generally described, and additional dietary evidence regarding calanoid copepod prey is provided for four of these taxa. All sternoptychid species investigated were planktivorous, feeding predominantly on copepods and ostracods, except for the largest size class, which preyed heavily on euphausiids and amphipods. The relationship of predator size towards prey type and prey size is analysed for both Sternoptyx species. Of these, S. pseudobscura in particular exhibits taxonomic selectivity towards polychaete prey. The diet of both species of Sternoptyx included a number of epipelagic or even neustonic calanoid copepod species which contributed more than 50% of the total copepod population by numbers. So far it is not known how the predators find access to prey organisms of the upper 200 m, as netfeeding is considered unlikely. Cyclopoid copepods of the genus Sapphirina were observed as dietary component particular of S. diaphana.     

Effects of salinity fluctuation on the hemolymph composition of the blue crab Callinectes sapidus

The hemolymph of the blue crab Callinectes sapidus was hyperosmotic during 20-10-20 S and 30-10-30 S diurnal cycles. The hemolymph became isosmotic at 26 S and hyposmotic at 28 S in the 10-30-10 S diurnal cycle. Hemolymph Na⁺ was hyperionic to seawater throughout all cycles. Hemolymph Cl^- was hyperionic below 24 S and either isionic or hypoionic from 24 to 30 S. Hemolymph K⁺ concentrations were hyperionic below 26 S and either isionic or hypoionic from 26 to 30 S. Hemolymph Mg⁺⁺ values were hypoionic over the experimental salinity range (10 to 30). Hemolymph ninhydrin-positive substances (NPS) levels were directly related to ambient salinity.