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     

Mating behavior of Centropages typicus (Copepoda: Calanoida)

Observations were made on approximately 300 mating couples of Centropages typicus Krøyer. Mating behavior consists of 4 distinct stages: pre-copulation, spermatophore extrusion, copulation, and post-copulation. Successful copulation requires proper positioning of the spermatophore complex on the female urosome and is dependent upon precise movements and orientation of the male during the mating encounter. Sub-optimal mating encounters, resulting in the improper placement of the spermatophore complex, were observed and adaptations tending to reduce spermatophore wastage are discussed.Contribution No. 62, Harbor Branch Foundation, Inc.     

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.     

Annual reproductive cycle, spatial distribution, abundance, and size structure of Oreaster reticulatus (Echinodermata: Asteroidea) in Bocas del Toro, Panama

The structure, distribution, and population abundance of Oreaster reticulatus (Linnaeus, 1758) in 47,157 ha of shallow-water habitat in the archipelago of Bocas del Toro, Panama, were assessed from May to October 2000. The reproductive cycle of the sea star was studied in Isla Solarte, from February 2000 to February 2001. In total, 4,818 sea stars were recorded with a mean density of 149.7 ind. ha^-1, and a population of over 7 million was estimated for the archipelago. O. reticulatus was absent in ca. 50% of the evaluated areas, possibly due to high runoff and sedimentation; highest density was observed in an intermediate-runoff regime (255 ind. ha^-1). About 45% of the population was found in substrata dominated by seagrass (Thalassia testudinum) and coarse, calcareous sand, 51% occurred in habitats where coral reef patches were mixed with seagrass, and 4% exclusively on coral reefs. The average size, based on the major radius, was 9.5 cm (3-21 cm), with a population structure composed of ca. 83% juveniles and 17% adults. The average reproductive size, measured as major radius, was 15 cm, and the minimum was 7 cm. Both males and females with a maximum stage (IV) of gonad development were observed throughout the year. The gonad index showed three peaks of maximum reproductive activity, which is not comparable to studies from other localities. The reproductive cycle did not seem to be related to water temperature, which ranged from 27°C to 30°C, but may respond more closely to changes in local rainfall. This relationship was not statistically significant based on this 1-year study. These data provide a useful baseline for management of local populations in the face of an increasing harvest for the aquarium trade and as souvenirs.     

The annual reproductive cycle of the holothurian Aslia lefevrei (Dendrochirota: Echinodermata)

The dendrochirote holothurian Aslia lefevrei (Barrois) is locally and densely aggregated (>70 individuals m^-2) in Galway Bay, on the west coast of Ireland. It has a discrete, relatively short breeding period from the end of February to the beginning of April, with peak activity in March. The sexes are separate and the single gonad, consisting of numerous tubules (100 to 800), lies in the anterior coelom. Both gonad index values and histological observations indicate that gametogenesis resumes in June/July after a brief post-spawned resting period. Little oocyte growth and sperm production occurs over the warm summer months when the holothurian is actively feeding. Gonad index values, however, increase rapidly during this period and reach high values by early winter without concomitant increases in maturity indices. This is subsequently followed by slight reductions in the gonad indices, before reaching peak values in February. It is suggested that this is probably due to the initial build-up and subsequent assimilation of storage products in the gonad, the development of gametogenic products and the frequent assimilation thereafter of unwanted ova. Spawning generally takes between 5 and 60 min, with the entire contents of the gonad rarely being shed in a single release. During this process, the holothurian raises its anterior end and, with the extended tentacles close together, emits a stream of gametes from the genital pore in the mid-dorsal line. Laboratory experiments and general observations indicate that increased temperature and low light levels may influence spawning behaviour.Contribution No. 248 to the School of Marine Sciences, University College Galway, Ireland     

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.     

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.     

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.     

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.     

Reproductive cycle,larval development,juvenile growth and population dynamics of Patiriella pseudoexigua (Echinodermata: Asteroidea) in Taiwan

A field study was conducted at Wanlitung, southern Taiwan, in 1986–1089, to determine the reproductive cycle, development mode, growth rate and population dynamics of the small seastar Patiriella pseudoexigua (Dartnall), which occurs in highly stressful and disturbed intertidal pools in this area. An inverse relationship between gonad index and pyloric-caccum index was only recorded immediately prior to spawning. A short, well-synchronized seasonal spawning occurs in October. When reared at 25 °C, lecithotrophic larvae develop directly, lack a bipinnaria stage, and metamorphose completely on the seventh day after fertilization. The growth curves of field juveniles are linear, those of laboratory-reared juveniles are sigmoid. Juveniles appear in tide pools in spring-early summer of each year. Adults spawn mainly in late fall, enabling spawning to occur in time for the larvae to benefit from the environmentally favorable winter season. Populations in high-tidal pools decrease in later summer, but remain more stable in lowtidal pools and lagoons.     

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     

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.     

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.     

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     

Mating behaviour of Macrophthalmus hirtipes (Brachyura: Ocypodidae)

 We examined the mating behaviour of the New Zealand ocypodid crab Macrophthalmus hirtipes in the laboratory between February and June 1998. This species has a discrete breeding season. Mating and moulting were not linked and only intermoult females with mobile gonopore opercula were attractive to males. Allometry and compatibility of gonopods and gonopores of different-sized crabs was investigated. Under laboratory conditions, the opercula of intermoult females remained mobile on average for 11.4 d, but the duration of receptivity did not appear to be under female control. The operational sex ratio in the laboratory fluctuated greatly, but was always male-dominated. During the period of opercular mobility, females mated many times with several different males. Matings in the absence of burrows were relatively short (mean duration = 23 min, max. = 122 min) and the mating behaviour of M. hirtipes lacked courtship and mate-guarding. Males used a search-intercept method to acquire mates, with very low levels of intrasexual competition. There was no evidence of mate preference in M. hirtipes, and males spent just as long mating with ovigerous females as with non-ovigerous ones. Although M. hirtipes has ventral-type spermathecae, as do several other ocypodid crabs, it is unclear whether this promotes last-male sperm precedence. The role of burrows in modifying the mating behaviour of M. hirtipes in the field remains to be established. Received: 7 January 2000 / Accepted: 5 June 2000     

Population differences in gonad and pyloric caeca cycles of the New Zealand seastarSclerasterias mollis (Echinodermata: Asteroidea)

Inshore and offshore populations ofSclerasterias mollis (Hutton, 1872) were sampled on the outer continental shelf off the Otago Peninsula, New Zealand, and their nutritional and reproductive cycles are described from 1985–1986. Histological changes in the gonads are generally typical of other asteroids. The gametogenic cycle takes 12 mo. The gonad and pyloric caeca indices in both females and males had an inverse relationship.S. mollis accumulates nutrients in the pyloric caeca during summer and early autumn. The gonads develop in the autumn and winter. Offshore seastars were much larger and had significantly higher gonad and pyloric caeca indices than inshore individuals. These differences in body sizes and organ indices arise from differences in food availability and/or population density.     

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.     

Ontogenetic habitat shift,population growth,and burrowing behavior of the Indo-Pacific beach star, <Emphasis Type="Italic">Archaster typicus</Emphasis> (Echinodermata; Asteroidea)

Archaster typicus, a common sea star in Indo-Pacific regions, has been a target for the ornamental trade, even though little is known about its population biology. Spatial and temporal patterns of abundance and size structure of A. typicus were studied in the Davao Gulf, the Philippines (125°42.7′E, 7°0.6′N), from February 2008 to December 2009. Specimens of A. typicus were associated with intertidal mangrove prop roots, seagrass meadows, sandy beaches, and shoals. Among prop roots, specimens were significantly smaller and had highest densities (131 ind. m⁻²) between November and March. High organic matter in sediment and a relatively low predation rate seemed to support juvenile life among mangroves. Size and density analyses provided evidence that individuals gradually move to seagrass, sandy habitats, and shoals as they age. Specimens were significantly larger at a shoal (maximum radius R = 81 mm). New recruits were found between August and November in both 2008 and 2009. Timing of recruitment and population size frequencies confirmed a seasonal reproductive cycle. Juveniles had relatively high growth rates (2–7 mm month⁻¹) and may reach an R of 20–25 mm after 1 year. Growth rates of larger specimens (R > 30 mm) were generally <2 mm month⁻¹. The activity pattern of A. typicus was related to the tidal phase and not to time of day: Specimens moved over the sediment surface during low tides and were burrowed during high tides possibly avoiding predation. This is one of the first studies to document an ontogenetic habitat shift for sea stars and provides new biological information as a basis for management of harvested A. typicus populations.