Brooding behaviour of the northern sea star Leptasterias polaris

The northern sea star Leptasterias polaris (Müller and Troschel) has a unique mode of brooding its young. The mother curves her arms along the plane of the disc, the whole individual assuming a flattened plate-like shape. In this form she covers the larvae, which are attached to the substratum beneath her. The central disc is not raised to form a brood chamber, as in related species, and there is no tendency to hide while brooding. In the St. Lawrence Estuary, brooding L. polaris are common from February through May and decline in numbers during June and early July. The genus Leptasterias originates in the Arctic and probably all species brood their young.This paper is dedicated to François Dorion who lost his life during benthic studies in the St. Lawrence Estuary. His enthusiasm in studying marine organisms will always be an inspiration to usContribution to the program of GIROQ (Groupe Interuniversitaire de Recherches Océanographiques du Québec)     

Genetic structure of populations of two closely related brittle stars with contrasting sexual and asexual life histories,with observations on the genetic structure of a second asexual species

Ophiocoma pumila Lütken andOphiocomella ophiactoides (H. L. Clark) are morphologically similar brittle stars with contrasting life histories, the former obligately sexual, the latter fissiparous (capable of both sexual reproduction and asexual proliferation by binary fission). Electrophoretic analysis of five polymorphic enzymes was used to assess the genetic consequences of these differing life histories and provide a genetic perspective on the taxonomic relationship between the two species. Genotypic diversity ofOphiocoma pumila collected at Discovery Bay, Jamaica, in 1985 conformed to expectations for a sexually reproducing population. In contrast, genotypic diversity ofOphiocomella ophiactoides at this site was significantly lower than expected for a sexually reproducing population, due largely to the predominance of clonal proliferation over larval recruitment. Large variation in clonal composition over a short (50 m) distance emphasized the very localized scale of clonal mixing in this species. Allozymic data are indicative of a close sibling species relationship betweenOphiocoma pumila andOphiocomella ophiactoides which suggests that the present generic separation of the two species should be re-examined. Electrophoretic analysis was also used to examine the genetic structure of sponge- and alga-dwelling populations of a second fissiparous brittle star,Ophiactis savignyi (Müller & Troschel), which was also collected at Jamaica in 1985. Striking differences in the allelic composition of sponge- and alga-dwellingO. savignyi were observed. Genotypic diversity ofO. savignyi in sponges was very low, each sponge being dominated by a single genotype. Genotypic diversity ofO. savignyi in algae was higher, although still significantly lower than expectations for a sexually reproducing population. In the light of the highly clonal composition of fissiparous brittle-star populations, the adaptive significance of clonal growth may be related to an increase in the overall fitness of dispersed clones (genets), compared to individuals of strictly sexual counterparts, through greater genotypespecific biomass and, hence, fecundity.Contribution No. 476 of the Discovery Bay Marine Laboratory of the University of the West Indies     

Reproduction of asteroids from Balsfjorden,Northern Norway: Analyses of lipids in the gonads of Ctenodiscus crispatus,Asterias lincki and Pteraster militaris

Seasonal lipid analyses were performed on the gonads and residual bodies of Ctenodiscus crispatus (Retzius) and Asterias lincki (Müller & Troschel), collected from Balsfjorden, northern Norway during March 1978–February 1980. The ovaries of both species had high percentages of total lipid (in the region of 60% of the dry weight) that were relatively constant throughout the season. Testes from both species had in the region of 20% of their dry weight as lipid, again with little seasonal variation. The majority of the total lipid in the ovaries of C. crispatus was accounted for by alkyldiacylglycerols and there was little seasonal variation in lipid class composition. Phospholipid accounted for the bulk of the lipid in the testes of C. crispatus, where alkyldiacylglycerols were present but in minor amounts. A slight degree of seasonal variation occurred in the composition of testis lipid in that maximal percentages of triacylglycerols and alkyldiacylglycerols occurred in January–April. The ovaries of A. lincki contained both triacylglycerols and alkyldiacylglycerols as major neutral lipids. Significant seasonal variation occurred in lipid class composition in the ovaries in that neutral lipids showed transient minima in February–April. The testes of A. lincki had substantial amounts of both triacylglycerols and alkyldiacylglycerols: maximal percentages of these lipids occurred in January–March. Analyses of gonads of near-mature Pteraster militaris (O. F. Müller) revealed that the ovaries were rich in lipid that was predominantly alkyldiacylglycerols, whereas the testes had much lower levels of lipid accounted for mainly by phospholipids. The residual bodies of all three asteroids had significant but small amounts of alkyldiacylglycerols in their lipids. A. lincki had triacylglycerols as a major lipid class in its body lipids, whereas in C. crispatus the major body lipid was phospholipid. Little seasonal variation was seen in the composition of the residual body lipid of either C. crispatus or A. lincki. The results are consistent with an aseasonal reproductive pattern in C. crispatus, although a main breeding activity occurred in mid-winter. A. lincki is a more seasonal breeder, but also had maximal activity in winter. Irrespective of their breeding habits, all three asteroids produced eggs rich in alkyldiacylglycerols. A major role for this lipid in the development of the large lecithotrophic eggs of asteroids is indicated.     

Reproductive cycle of two populations of Ophionereis schayeri (Ophiuroidea) in New South Wales

Reproduction of two populations of Ophionereis schayeri (Müller and Troschel) in New South Wales was investigated from November 1991 through January 1993. The reproductive cycle was documented by histological examination of the gonads and by image analysis to determine the oocyte size-frequency distributions. Both populations of O. schayeri were mature in December and January and had a major summer spawning period between January and February. Thereafter, the condition of the gonads exhibited interindividual variability, with asynchronous low-intensity gamete release through August. O. schayeri has the potential to spawn for up to 8 mo of the year. This breeding pattern with synchronous spawning in summer and asynchronous gamete release through winter was similar in both populations. During autumn and winter, the gonads contained developing gametes and advanced gametes ready for spawning. The eggs spawned by O. schayeri during these seasons started their growth during the previous spring vitellogenic period, and continued to grow through summer. Upon reaching full size, they were stored for spawning outside the main breeding period. Spring is marked by increased spermatogenic and vitellogenic activity followed by maturation and spawning. Throughout their development, the oocytes of O. schayeri are surrounded by a follicle consisting of periodic acid-Schiff-positive (PAS+) haemal fluid and PAS+ yolk by the oocytes. In the testes, the haemal sinus projects into the centre of the spermatocyte columns, thus appearing to play a nutritive role in support of both oogenesis and spermatogenesis. Late vitellogenic oocytes of O. schayeri are firmly anchored to the germinal epithelium by an attachment complex consisting of specialised attachment cells and basophilic strands that radiate between the oolemma and the nucleus. Spawning is associated with rupture of the follicles, which remain as prominent, empty U-shaped profiles in the ovaries. O. schayeri produces copious numbers of 200 m-diam oocytes, suggesting that this species is a broadcast spawner and that it has a modified ophiopluteus or vitellaria larva.     

Elemental distribution in relation to skin necroses of marine flatfishes from the english channel

It has been suggested that industrial waste discharges are associated with the occurrence of fin damage and skin lesions. A mixed population of diseased and healthy blatfish consisting ofLimanda limanda, Pleuronectes platessa, Platichthys flesus andSolea solea were collected from an industrialised area (Calais, France) and were compared with diseased and healthyl. limanda caught in a non-industrialised area (Brixham, England). Analysis of skin samples from necrosed and healthy fish by X-ray microanalysis and atomic absorption spectroscopy has shown that no elemental accumulation occurs either in necrosed fish, or fish from an industrialised area. The fish from Calais were heavily infested with the ectoparasitic copepodLepeophtheirus pectoralis (O.F. Müller) and, although few were found on fish from Brixham, it is unlikely that this infestation was related to the discharge of industrial effluents.     

Modification of sediment permeability and shear strength by two burrowing invertebrates

Laboratory experiments have been conducted on the effects of bioturbation by the amphipod Corophium volutator (Pallas) and the polychaete Nereis diversicolor (O.F. Müller) on the physical properties of estuarine muddy sand. Invertebrates and muddy sediment were collected from Langbank, Firth of Clyde, Scotland. Sediment columns were seeded with low, medium and high densities of invertebrates in single-and mixed-species experiments. Permeability decreased with increasing density of C. volutator, but increased with increasing density of N. diversicolor. Water content decreased with increasing density of both species. Shear strength increased with increasing density of both species, but more so with C. volutator than with N. diversicolor. Mixed-species effects on shear strength were additive, but some of the permeabilities were lower than predicted. Bivariate and multivariate correlation, and partial correlation analyses were applied to the data. Shear strength was negatively correlated with water content. The correlations between shear strength and permeability and between permeability and water content were affected differently by the two species. There were few partial correlations between the variables.     

Allozyme electrophoretic analysis of the Hediste limnicola — H. diversicolor — H. japonica species complex (Polychaeta: Nereididae)

Three morphologically similar species of the polychaete genus Hediste [H. limnicola (Johnson, 1901), H. diversicolor (O. F. Müller, 1776), and H. japonica (Izuka, 1908)] have been described, respectively, from the west coast of North America, Europe, and Japan. Although these taxa are geographically isolated by oceans, the morphological criteria by which they can be separated are sufficiently variable as to make identification difficult. We have compared these three taxa biochemically using ten allozyme loci and have found that they are genetically distinct and constitute valid species. H. japonica (collected in June 1990 from Kyushu, Japan) appears more closely related to H. limnicola (collected in July 1989 from two sites in California and two in Oregon, USA) than it is to H. diversicolor (collected in October-November 1989 from the Weser estuary and the Jadebusen in Germany). H. limnicola has been recognized previously as a self-fertilizing hermaphrodite, but it is quite polymorphic in the four populations examined and we suggest that some cross-fertilization must occur in the field. Marked allele frequency differences between the two H. diversicolor populations examined, which were separated by only about 30 km, suggested that there was restricted gene flow between populations, or that there had been strong short-term selection.     

Reproductive pattern in the brooding and broadcasting sea star Pteraster militaris

Pteraster militaris (O. F. Müller, 1776) broods young in an aboral chamber and broadcasts gametes and offspring as well. The reproductive cycle of a population of P. militaris from the northwest Atlantic Ocean was examined over a 23 mo period from September 1985 to July 1987. The species is gonochoric (sex ratio 1:1), with only one incidence of hermaphroditism observed (n=261). Gonad indices were lower in females than in males, possibly an adaptation to the limited space available in the brood chamber. Body component and maturity indices, gonad histology and response to 1-methyladenine indicate that females in the population have a continuous breeding season, with a slight decrease in intensity in the spring. Males have a more pronounced decrease in the intensity of reproduction in the spring. Gonad development in males in synchronous both within and between individuals. Gonad development in females is asynchronous.     

Mating behaviour and reproductive cycle of Archaster typicus (Echinodermata: Asteroidea)

This study examines the reproductive cycle, the mechanism of male-on-female pairing behavior and the spawning behavior of Archaster typicus Müller et Troschel. Field studies were conducted in the intertidal zone of the sand beaches at Penghu, Taiwan (23 32N; 119 33E) at ebb-tide in 1984 and 1985. The pairing behavior of A. typicus is a reproductive behavior which leads to simultaneous spawning, increasing the probability of fertilization. As the breeding season approaches, sea stars, especially males, display increased mobility. Because only males tend to mount another individual and because males can detect the sex of another individual by contact with the side of their arms, a male-on-female pair is formed when a male encounters a female. Eighty-five percent of the sea stars observed were paired during the pairing season. Spawning by a paired female is closely followed by spawning of its paired male; male spawning, however, does not induce spawning in its paired female. During spawning, the male turns slightly so that its arms overlap the arched arms of the female. The gonad volume of males is much less than that of females. This may result from the high efficiency of fertilization in this species, which does not require a large amount of gametes to be released, or from the higher energetic demands made on the males.     

Population dynamics and reproduction of Mysella bidentata (Bivalvia: Galeommatacea) in Galway Bay,Irish west coast

Mysella bidentata (Montagu) is found in high densities in an Amphiura filiformis (O. F. Müller) community in Inner Galway Bay, west coast of Ireland. As part of an ongoing investigation at one station within this community, the population dynamics and reproductive pattern of this bivalve were investigated during the period November 1978-December 1981. Monthly sampling over the 3 yr period revealed a mean population density of 745±672 individuals m^-2. Recruitment occurred during August-October and three year-classes were identified. Growth rates were assessed by analysis of length-frequency histograms and winter growth checks. Individuals from the Y1 and Y2 year-classes functioned as hermaphrodites during the 1980 and 1981 reproductive seasons. The Y0 year-class contained both males and hermaphrodites. Results are compared with available information on the same species from other locations.Contribution No. 225 from the School of Marine Sciences, University College, Galway Ireland     

Reproductive energetics of two species of dorid nudibranchs with planktotrophic and lecithotrophic larval strategies

Onchidoris muricata (Müller) and Adalaria proxima (Alder and Hancock) are sympatric, potentially competing species of dorid nudibranchs, which preferentially graze the cheilostome polyzoan Electra pilosa (L.). O. muricata is small and lays small eggs which hatch as poorly-developed planktotrophic veliger larvae. A. proxima is larger and reproduces by means of larger eggs which hatch, as well-developed lecithotrophic larvae, that can metamorphose within approximately 24 h of release. A. proxima larvae can feed in the plankton, but do not require extrinsic nutrition to undergo complete development. Both species spawn in February–april, and have a strictly annual life-cycle. Comparisons of the calorific content of spawn have shown that A. proxima apportions a greater number of calories to reproduction, but that O. muricata makes a greater relative effort. A. proxima shown considerable individual variability in reproductive effort, which fails to correlate with, body size or rate of spawning. A more deterministic situation applies to O. muricata, because body size and fecundity follow an allometric relationship. It appears that there is a threshold of absolute energy required to support the lecithotrophic larval strategy in nudibranchs, and that this is not attained by the smaller species, O. muricata. A. proxima thus appears to have both reproductive strategies open to it, and to have adopted lecithotrophy in order to offset the unpredictability of energy available for reproduction.Communicated by J. M. Mauchline, Oban     

Reproductive behaviour ofNereis diversicolor (Annelida: Polychaeta)

Nereis diversicolor O. F. Müller, collected on the tidal flats of the Jadebusen (North Sea, FRG) in autumn 1987 and 1988, was exposed to different temperatures in the laboratory. Results indicated that maturation was induced by temperatures above 6°C; spawning in early spring was synchronized by raising temperatures after a period of low temperatures in winter, and occurred mainly at new and full moon. During reproduction the female stays inside the burrow; the male releases sperm in front of it; the resultant larvae remain in the tube for 10 to 14 d protected by the female.     

Joint breeding in female burying beetles

Summary Burying beetles (Nicrophorus) exhibit advanced parental care, by feeding and guarding their offspring on buried vertebrate carrion. Till now, interactions between two conspecific females on a carcass have been thought to be mostly competitive, and parental care was thought to be provided by single females or male-female pairs exclusively. Here we demonstrate that cooperative brood care occurs in this species, and that its incidence is contingent on carcass size. Small carcasses are usually monopolized by one female; typically the larger of two female combatants secures the carcass for her offspring (Figs. 1 and 2). On large carcasses fights still occur, but in most cases both females stay on the carcass long enough to provide care for the brood. The use of genetic markers revealed that the maternity of offspring is shared evenly among joint breeders (Figs. 3, 4). We hypothesize that cooperative breeding is an adaptive response to a situation that arises partly as a consequence of a physical constraint. Correspondence to: J.K. Müller     

Influence of cultivation and food concentration on body length of calanoid copepods

The cephalothorax length of the marine pelagic Copepoda Acartia clausi Giesbrecht, Temora longicornis (Müller), Centropages hamatus (Lilljeborg) and Pseudocalanus sp. was monitored at 15°C during prolonged cultivation through up to 55 filial generations and at different concentrations of food. The length of T. longicornis decreased considerably during the first 15 generations and remained rather constant thereafter. In the other species, body length increased slightly or remained almost constant. Genetic changes are probably involved. Food concentration influenced body size of all species, particularly C. hamatus, in which 80% of the natural size range may be explained by differences in food concentration. The idea that temperature is a dominant factor in determining the length of copepods should be reconsidered.     

Reproductive cycle of the subarctic brooding asteroid Leptasterias polaris

The reproductive cycle of the large brooding seastar Leptasterias polaris Müller and Troschel was examined over an 18-month period in the St. Lawrence Estuary, Québec, Canada. There is a distinct annual cycle with spawning over several months in the autumn. The female has an unusual cycle in that the ovary only slightly decreases in size during spawning, and the size structure of the oocyte population is remarkably stable throughout the year. The major annual change observed in the oocyte population is the development of a small number of 600-to 800- oocytes prior to spawning and their loss during spawning. This stability, combined with the lack of evidence of phagocytosis, suggests that oocyte development takes place over many years. By contrast, the developmental cycle of the testis is similar to that of most echinoderms. The large reservoir of oocytes probably guarantees a steady annual recruitment, and brooding during the winter probably minimizes metabolic costs for the female and ensures the release of the juveniles when conditions are favourable in the spring and summer.     

Transfer of radioactive carbon within the copepod Temora longicornis

Standard autoradiographic techniques have been used to describe the temporal pattern in the transfer of radioactive carbon from the chlorophyte Tetraselmis suesica (Kylin) to females of the neritic copepod Temora longicornis (Müller, 1792). After 4 and 6 h exposure to labelled phytoplankton, silver grains were deposited only in the cells of the gut-wall. After 24 h exposure, grains were deposited in most tissues, with highest concentrations in cells of the gut-wall and in oocytes within the ovary. Experiments with Centropages typicus (Krøyer), Centropages hamatus (Lilljeborg), and Labidocera aestiva Wheeler demonstrated that much of the assimilated label was not turning over rapidly and that the amount of label assimilated was correlated with known food preferences of these species.     

Population dynamics and reproductive cycle of Pholoë minuta (Polychaeta: Sigalionidae) in Galway Bay

Pholoë minuta (Fabricius) is a prominent member of the Amphiura filiformis (O.F. Müller) subcommunity in Galway Bay, Ireland. The aim of the present investigation was to study the population dynamics and reproductive cycle of P. minuta in this area. Monthly sampling over a 2 yr period (November 1978–October 1980), at a permanent station (18 m) with a fine sand substrate, revealed a mean density of 96 individuals per 0.1 m². The width of the fifth elytron was correlated with total body length and was used as a basis for analysing population structure. Four distinct year classes were evident and estimates of recruitment levels and growth rates were calculated. P. minuta is polytelic and reproduces for the first time when approaching 3 yr of age. Spawning occurred in March—April and appeared to be associated with a seasonal low in bottom-water temperature.     

An osteological study of the dugong Dugong dugon (Sirenia) from India

An osteological study of the dugong (sea cow) Dugong dugon (Müller) from India, based on complete skeletons of an adult male and a juvenile female indicated certain morphological variations with age. It has also revealed a close osteological similarity between the dugongs from India and the Red Sea. Based on a statistical analysis, regression equations for certain typical skull measurements have been calculated; these equations, characteristic of the dugong from India, could be used for comparing dugongs of different regions.     

Uptake of 55Fe from contaminated sediments by the polychaete Nereis diversicolor

The polychaete Nereis diversicolor O. F. Müller was exposed to marine sediments which had been contaminated with ⁵⁵Fe either through the testing of nuclear devices or by the release of liquid waste effluent from a nuclear fuel reprocessing plant. The amount of ⁵⁵Fe taken up reached a concentration in the worms about 6% of the concentration in both types of sediment. Bioavailability of ⁵⁵Fe was shown to depend on its concentration in sediment and not on sediment type, stable iron concentration, or the source of contamination. N. diversicolor may not extract from sediment the same fraction of ⁵⁵Fe as does 0.1N HCl and caution should be exercised in using such measurements to represent the bioavailable fraction.     

Predator avoidance behaviour of Baltic littoral mysids<Emphasis Type="Italic"> Neomysis integer</Emphasis> and<Emphasis Type="Italic"> Praunus flexuosus</Emphasis>

The predator avoidance behaviours of two littoral mysid species, Neomysis integer (Leach) and Praunus flexuosus (Müller), were studied experimentally. In ingestion experiments, mysids responded to a combination of chemical and visual signals of perch (Perca fluviatilis), but not to each stimulus alone. In the presence of the combined visual and chemical predator signal the swimming activity and choice of habitat (open vs. artificial vegetation, the Charophyte Chara tomentosa or the brown alga Fucus vesiculosus) were also influenced. The two mysid species behaved differently when perceiving predation risk: N. integer reduced swimming activity, whereas P. flexuosus increased their use of the vegetation. The different antipredator strategies of the two mysid species reflect their different lifestyles, N. integer being a swarm-forming species and P. flexuosus living in association with aquatic macrophyte vegetation.Communicated by L. Hagerman, Helsingør