Seasonal changes in biochemical composition of body walls,gonads and pyloric caeca in two populations ofSclerasterias mollis (Echinodermata: Asteroidea) during the annual reproductive cycle

Seasonal changes in biochemical composition of tissues are compared in a shallow inshore and deeper offshore population of the seastarSclerasterias mollis (Hutton, 1872) on the Otago shelf, New Zealand, from 1985 to 1986. The biochemical composition of the body wall, gonads and pyloric caeca of the two populations did not differ greatly, even though reproductive output and nutrient storage of reserves were consistently higher in the offshore population. The biochemical composition of the ovaries and testes differed. High levels of carbohydrate and lipid were maintained in the ovaries from the commencement of vitellogenesis until spawning occurred. The testes had higher ash levels during spermatogenesis. During the annual reproductive cycle, an inverse relationship in the carbohydrate levels between the ovary and the pyloric caeca suggests nutrient transfer from the pyloric caeca to the gonads during gametogenesis. Annual changes in biochemical composition indicate that the body wall is also used for nutrient storage.     

Population genetics and systematics of theLeptasterias hexactis (Echinodermata: Asteroidea) species complex

Morphological characters and 14 enzyme loci were examined for 1 040 sea stars, currently recognized as forms ofLeptasterias hexactis, from Lynn Canal, Alaska, and Puget Sound, Washington, USA, between March 1988 and April 1989. Three morphologically and two genetically distinctLeptasterias forms were identified. The most common form found at both localities wasL. epichlora (Brandt)sensu Verrill.L. hexactis (Stimpson)sensu Verrill co-occurred withL. epichlora at all study sites and apparently hybridizes extensively withL. epichlora in the Puget Sound region, but rarely, if at all, in Alaska. The presumptive product of this hybridization morphologically resembledL. aequalis (Stimpson)sensu Fisher, and was conspicuously absent from Alaskan samples. Considerable genetic distance existed betweenL. epichlora andL. hexactis (Nei'sD=0.19±0.01) and moderate genetic differentiation occurred between populations of each species from Alaska and Washington (Weir and Cockerham'sF _RT =0.29±0.04 forL. epichlora and 0.21±0.15 forL. hexactis). A significant (p<0.05) deficiency in the proportion of heterozygous individuals was found compared to Hardy-Weinberg expectations (Wright's fixation index,F _ID=0.12±0.04 and 0.31±0.08 forL. epichlora andL. hexactis, respectively). However, mean observed heterozygosity for each species (0.09±0.03, 0.14±0.04 and 0.14±0.04 forL. epichlora, L. hexactis andL. aequalis, respectively) fell within the range of reported values for other asteroid species (ca. 0.04 to 0.37). The results of this study indicate that considerable genetic integrity is maintained betweenL. epichlora andL. hexactis, which warrants their recognition as distinct species despite their apparent hybridization in the Puget Sound region.     

Responses of Leptasterias hexactis (Echinodermata: Asteroidea) to low salinity

The six-rayed starfish Leptasterias hexactis (Stimpson, 1862) is seasonally exposed to low salinities in southeastern Alaska. Individuals that were gradually exposed to reduced salinities in the laboratory had a 28-d TLm of 12.9 S. The activity of L. hexactis, as measured by its activity coefficient, varied directly with salinity. Individual feeding rates of the starfish on similarly exposed Mytilus edulis, measured daily for 21 d at salinities of 30, 20 and 15 S, also varied directly with salinity. The dry weight of mussel tissue consumed was 8.84, 8.49 and 0.58 mg ·starfish^-1·d^-1 at 20, 20 and 15 S. Expressed as percent of dry starfish weight, the daily feeding rate was 1.35, 0.76 and 0.10% at 30, 20 and 15 S. Absorption efficiency decreased from 64% at 30 S to 49% at 20 S, further reducing the energy available for metabolism. Growth, measured in terms of changes in total dry weight or dry weight of soft tissues, also varied directly with salinity. Although exposure to hyposmotic conditions did induce stress responses, as indicated by reductions in activity, feeding, absorption efficiency and growth rates, L. hexactis maintained positive growth for at least a 3-wk period in the laboratory at 20 S and 13°CC. The population of L. hexactis investigated must be considered euryhaline and brief periods of exposure to hyposmotic conditions should not limit its distribution.     

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.     

Prey capture by Pisaster brevispinus (Asteroidea: Echinodermata) on soft substrate

Observations on the subtidal seastar Pisaster brevispinus (Stimpson) indicate that this predator can extend the central tube feet into sand-mud substrate for a distance roughly equal to the radius of the seastar. Field and laboratory evidence demonstrates the use of the elongated tube feet by the asteroid for the capture of burrowed prey items. A relationship between predator size and the size of juveniles of the deep-burrowing clam Tresus nuttallii taken as prey is a product of the ability of larger seastars to extend the tube feet further into the substrate and capture deeper prey. The role of P. brevispinus as an important subtidal predator along the Pacific Coast of North America is discussed.     

Reproduction in Amphiura filiformis (Echinodermata: Ophiuroidea): Seasonality in gonad development

The burrowing ophiuroid Amphiura filiformis (O. F. Müller) (Echinodermata: Ophiuroidea) colonises an extensive area and is a numerically dominant member of the macrofauna in Galway Bay on the west coast of Ireland. There, it has a discrete, relatively short annual breeding period, from June to September, with peak activity in the month of August. The breeding period approximates in timing those of other European populations. The oocytes of a particular year are laid down in the autumn of the previous year; they begin to develop in spring, with the period of fastest growth in May/June. Gonad growth corresponds quite closely to the annual rise in water temperature, with spawning taking place during the warmest months of the year. The species is polytelic, and it is suggested that the long-lived Galway Bay individuals may breed every year for several years. The relationship between spawning and population dynamics is discussed in the light of current, available literature concerning A. filiformis from other parts of Europe. The role of so-called ultimate factors in the process of ophiuroid maturation and spawning is discussed.     

Terrestrial runoff as a cause of outbreaks of Acanthaster planci (Echinodermata: Asteroidea)

Outbreaks of adult Acanthaster planci (Linnaeus) have appeared at irregular intervals, arriving 3 yr after heavy rains (>100 cm in 3 months) following droughts (<25 cm in 4 months) or 3 yr after rains exceeding intensities of 30 cm in 24 h. Outbreaks of A. planci follow typhoons that bring heavy rains, but do not follow dry typhoons of equivalent wind force. Outbreaks occur around the high islands in Micronesia and Polynesia, but not around the atolls at intermediate locations. Phytoplankton blooms appear off high islands at the beginning of the rainy season in bays with large watersheds and with sufficient residence time of the waters; these are the initial sites of A. planci abundance on Guam. The spawning seasons of A. planci occur at the beginning of the rainy season on both sides of the equator. I hypothesize that, on rare occasions, terrestrial runoff from heavy rains (following the dry season or a record drought) may provide enough nutrients to stimulate phytoplankton blooms of sufficient size to produce enough food for the larvae of A. planci. The increased survival of larvae results in an outbreak of adults 3 yr later. This hypothesis can be tested by predicting future outbreaks. An outbreak of A. planci on Saipan in the summer of 1981 was predicted on the basis of heavy rains in August 1978.Contribution no. 169, University of Guam Marine Laboratory     

Population structure, growth and recruitment of the euryalinid brittle-star Astrobrachion constrictum (Echinodermata: Ophiuroidea) in Doubtful Sound, Fiordland, New Zealand

The euryalinid brittle-star (snake star) Astrobrachion constrictum (Farquhar) lives coiled around the branches of black coral (Antipathes fiordensis) colonies. Twenty-two vertical transects, 10 m wide by 30 m deep, were swum in Doubtful Sound over a 2.5 yr period from 1993 to 1995. Numbers, disc diameters and colour morphotypes of brittle-stars inhabiting coral colonies were recorded. 36.3% of the coral colonies >200 mm tall (n = 292) hosted ≥1 Astrobrachion constrictum (range 0 to 12). Overall, the population was patchily distributed on the available coral habitat. The dark red colour morphotype of A. constrictum was most common (87%, n = 279) followed by the yellow, striped, and then spotted varieties. The population was comprised mainly of large (≥10 mm disc diam) individuals, and juveniles were rarely encountered, indicating low rates of recruitment or a high mortality of recruits. Disc-diameter data gathered from this and previous studies indicated that growth in A. constrictum is initially rapid, with individuals reaching a disc diameter of 15 mm in ≃2.5 yr; growth decreases with age, as in other deep-sea ophiuroids. Growth rate within years, however, was not constant, with faster growth in the spring/summer. Maximum size for A. constrictum is reached in ≃8 yr at ˜23 mm disc diam. Anecdotal evidence indicates that A. constrictum may not be confined solely to black coral colonies. Received: 25 September 1996 / Accepted: 16 October 1996     

Costs of extended brood protection in the Antarctic sea star, Neosmilaster georgianus (Echinodermata: Asteroidea)

Changes in size, morphology, and biochemical composition in adults and embryos of a brooding sea star, Neosmilaster georgianus (Sladen), were studied in a population adjacent to Anvers Island, Antarctic Peninsula (64°46′S; 64°04′W) during the austral spring, 1991. Five morphological stages of development were designated in 24 broods, and for each the weight and biochemical composition of the brooding adults and their embryos were determined. Between Stage 1 and 2, the dry weight (dw) and organic weight (ow) of the embryo did not change. From Stage 2 to 3, the dw and ow increased significantly by 10%. Stage 2 and 3 embryos were in clusters of a few (2 to 10) to as many as 40 individuals. In the smaller clusters, individual embryos were attached by tissue cords to another, sometimes atrophied, brood member. In the larger clusters, they were attached to a central mass of tissue containing remnants of embryos. We interpret these interactions as a form of cannibalism which may account for the weight gains between Stage 2 and 3. During Stages 4 and 5, as juvenile form was approached, the dw and ash weight of the young increased significantly and the ow decreased significantly. The calculated energy content for the juvenile (Stage 5) was not significantly different from the energy content of the earliest undifferentiated stage (Stage 1), an indication that most of the organic matter in the egg is the primary contribution to the large juvenile. In brooding females, pyloric caeca indices declined by 52% from Stage 1 to Stage 5 and pyloric energy stores declined by 63% due to proportionately equivalent declines in protein and lipid. The ovary index was low and increased only slightly during brood protection, while the size of the largest oocytes remained approximately 23% that of ova. Energy stores in the pyloric caeca of brooding N. georgianus thus become depleted over a long period of incubation during which the adults apparently starve. This may delay oocyte development and ultimately limit the frequency of individual reproduction. Received: 15 September 1998 / Accepted: 9 March 1999     

Acclimation of gametes to reduced salinity prior to spawning inLuidia clathrata (Echinodermata: Asteroidea)

AdultLuidia clathrata (Say) were held at salinities lower and higher (25 to 35) than environmental salinity (29) during gametogenesis and after sperm and primary oocytes were produced. The sperm and primary oocytes acclimated to the experimental salinities in both cases. Fertilization and development through gastrulation were not affected at the salinities at which the parents were held, but were affected at altered salinities. This indicates that intracellular osmoregulation can occur both during gametogenesis and after the stage of arrested activity has been reached. This is an important capacity forL. clathrata, which lives in estuaries with variable salinities.     

On the distribution and variability of the Indo-Pacific sea star Asterina wega (Echinodermata: Asteroidea) in the Mediterranean sea

The distribution of the Mediterranean populations of Asterina wega Perrier is described. Four populations were found to differ in size of animals and regeneration rate. The possible migratory pattern is duscussed.     

Mass spawning by the sea urchin Evechinus chloroticus (Echinodermata: Echinoidea) in a New Zealand fiord

A mass synchronous spawning of the sea urchin Evechinus chloroticus (Valenciennes) was observed in situ in Doubtful Sound, a large New Zealand fiord. Spawning occurred between 17:30 hrs and 18:30 hrs on 27 January 1994 and coincided with a full moon, spring tides and a period of decreasing sea temperatures. During spawning, the sea urchins formed a dense spawning aggregation of both sexes, with >90% of the urchins observed spawning at the time. Spawned gametes clouded the water column, and some were eaten by small labrid fish species. The spawning, which may have been as widespread as 40 km, marked a 42 to 50% decrease in gonad indices and resulted in a widespread, dense cohort of  E. chloroticus larvae within the fiord. Received: 25 September 1997 / Accepted: 6 March 1998     

The effect of temperature-salinity combinations on survival and growth of juvenile Patiriella pseudoexigua (Echinodermata: Asteroidea)

Adult Patiriella pseudoexigua were collected in October 1989 from Wanlitung, Taiwan and then induced to spawn in the laboratory. Post-metamorphosed juvenile P. pseudoexigua were reared on a diet of benthic algae Navicula sp. at 25°C and salinity (34). Six weeks after metamorphosis, juvenile P. pseudoexigua at ca. 400 m in radius were reared on a diet of benthic algae Navicula sp. at different combinations of temperatures (20, 25, 30°C) and salinities (26, 30, 34) for 40 d. Both temperature and salinity had a significant effect on juvenile survival and growth. Juveniles survived best (>90%) at 25°C and 34 and grew best (to ca. 750 m in radius) at 30°C and 34. Variation in juvenile size was small immediately after metamorphosis and increased with time.     

Differences in reproductive strategies of morphs of the genus Echinaster (Echinodermata: Asteroidea) from the Eastern Gulf of Mexico

Two morphs of the asteroid genus Echinaster, E. Type 1 and E. Type 2, have reciprocal annual cycles of reproduction and nutrient storage. The pyloric caeca (storage organs) reach maximal size over winter then decrease with increasing gametogenic activity in the spring. E. Type 1 broadcasts buoyant eggs in late May to early June. E. Type 2 deposits benthic eggs in late April. Lecithotrophic larval development is similar in both morphs except that larvae of E. Type 1 undergo a brief planktonic phase. E. Type 1 reaches sexual maturity at a larger size, expends a lower reproductive effort but has a greater absolute reproductive output than E. Type 2. E. Type 2 produces fewer, larger eggs and has a greater parental investment per egg. In both morphs, females have much higher gonadal lipid levels than males and expend a higher reproductive effort in terms organic matter and energy. Reproductive effort, reproductive output, nutrient storage in the pyloric caeca and body size varied between populations and between years in each morph. Egg size and parental investment per egg were constant. Within-morph variability is attributed to differences in nutritional state. Differences in reproductive strategy support the hypothesis that the morphs are separate species.     

Mitochondrial DNA polymorphisms reveal additional genetic heterogeneity within the Leptasterias hexactis (Echinodermata: Asteroidea) species complex

Sea stars of the Leptasterias hexactis complex are important members of the rocky intertidal community of the temperate and subarctic North American Pacific coasts. Recent attempts to determine the genetic structure and taxonomic relationships of these sea stars have not been completely successful, due to the combination of extreme morphological variability and relative scarcity of taxonomically-informative genetic (allozyme) polymorphisms. The present study used mitochondrial DNA restriction-site polymorphisms for PCR (polymerase chain-reaction)-amplified products to provide additional information about this complex. Five restriction-site haplotypes were identified in southern Alaska and in the Puget Sound region that were allozymically distinct. Because the allozymic differences (Nei's unbiased genetic differences >0.08) are maintained in sympatry, the haplotypes behave evolutionarily as conventional species. Although most haplotypes show deficiencies of heterozygotes for polymorphic allozyme loci, there is no evidence for the existence of cryptic species within any of the haplotypes. Rather than consisting of a few polytypic and cosmopolitan species, the L. hexactis complex is apparently composed of numerous species with more restricted ranges. Determining the exact ranges and taxonomic relationships of the various species will require additional study.     

Feeding rate and duration of daily activity of Astropecten aranciacus (Echinodermata: Asteroidea) in relation to prey density

In a feeding experiment, 16 individuals of Astropecten aranciacus were kept in two enclosures on the sea floor, and were provided with a surplus of food. Feeding at a daily rate of 1.3% of their dry tissue weight, their food intake was four times higher than that of free-living sea stars. The periods of daily activity were shorter than in control individuals kept in enclosures without additional food.     

Reproductive cycles of tropical sea cucumbers (Echinodermata: Holothuroidea) in southern Taiwan

Reproductive cycles of Afrocucumis africana (Dendrochirotida), Actinopyga echinites, Holothuria leucospilota, H. cinerascens, H. difficilis (Aspidochirotida), Synaptamaculata, Opheodesoma grisea, Patinapta taiwaniensis and Polycheira rufescens (Apodida), representing three orders of intertidal holothurians in southern Taiwan, were determined by gonad index and histological examination from March 1990 through September 1991. All nine species of holothurians have annual spawning periods lasting 2 to 4 mo in spring or summer. The early spawning of H. cinerascens and P. rufescens (suspension feeders) suggests that their feeding mode is related to food resources and may influence the reproductive period. In the direct-developing dendrochirotid (suspension feeder), gonad development occurs in cool months, and gametes are released at the beginning of warm months. In indirect-developing aspidochirotid and apodid (deposit feeder), gamete release occurs in late spring or summer and appears to be correlated with summer phytoplankton growth. The holothurians in the present study display spawning periodicity and duration similar to those of temperate species. Four species of holothurians lengthened spawning season with decreased latitude.     

Genetic consequences of long larval life in the starfish Linckia laevigata (Echinodermata: Asteroidea) on the Great Barrier Reef

Gene flow between populations of the asteroid Linckia laevigata (Linnaeus) was investigated by examining over 1000 individuals collected from ten reefs throughout the Great Barrier Reef (GBR), Australia, for genetic variation at seven polymorphic enzyme loci. Despite geographic separations in excess of 1000 km, Nei's unbiased genetic distance (0 to 0.003) and standardised genetic variation between populations (F _ST) values (mean 0.0011) were small and not significant. Genetic homogeneity among L. laevigata populations is consistent with the long-distance dispersal capability of its 28 d planktonic larval phase, and is greater than that observed for other asteroid species, including another high-dispersal species, Acanthaster planci, which has a 14 d larval phase. Variation within populations was also higher than previously recorded for asteroids (mean heterozygosity=0.384; number of alleles per locus ranged from 5.1 to 6.0 in each population). Among asteroids, dispersal ability is positively correlated with gene flow and levels of variation, and negatively correlated with levels of differentiation.     

The reproductive cycle of Asterina stellifera (Möbius) (Echinodermata: Asteroidea) in the Cabo Frio region, southeastern Brazil

Reproduction of Asterina stellifera was studied in two populations from the Cabo Frio region, southeastern Brazil. One was located in an intertidal seagrass flat at Japonês Island (1994–1995); the second was on a sublittoral rocky shore at Pontal beaches (1999–2000). Reproductive cycle was studied at monthly intervals by organ indices and histological examinations of gonads. Although these two populations were studied in different habitats and at different times, the periodicity of the annual reproductive cycle was very similar for the two populations. The highest gonad index (GI) values were found in August/September (late winter), followed by a drop in GI values, indicating spawning. The GI and pyloric caeca index (PCI) tended to have an inverse relationship in both populations, but it was only statistically significant in the rocky-shore population. Gametogenesis in the two populations was described and roughly confirmed the reproductive cycles determined by GIs. Published online: 9 August 2002