Factors controlling spatial distribution of the sea cucumber Psolus chitonoides: Settling and post-settling behavior

Substratum selection experiments were carried out in situ with larvae of the holothurian Psolus chitonoides Clark in the rocky subtidal of the San Juan Islands, Washington. The experiments indicated, in agreement with laboratory findings, that pentactula larvae settle gregariously, either on the adult, or (when the bivium is obscured by epizooites) by clustering around the margin of the adult sole. Late larvae and early juveniles are strongly photonegative. Following metamorphosis, juveniles migrate from the adults into nearby shaded habitats, where they may take up residence indefinitely. Although adults retain the capacity to move, they relocate only when their living conditions become unsuitable; in most circumstances they are effectively sessile. Small-scale spatial pattern of the adults, which was documented quantitatively on subtidal rock walls, boulders and cobbles, correlates well with microtopographical features. The aggregation observed in the field is primarily due to post-settling migration rather than larval substratum choice at settlement.     

Spawning behaviors of northeast Pacific sea cucumbers (Holothuroidea: Echinodermata)

Spawning behaviors observed from May 1981 through August 1983 in the laboratory and field are described and compared for twelve species of northeast Pacific holothuroids collected from waters of the San Juan Archipelago, Washington. For many species, elevation of the anterior end, stretching of the introvert, tentacle waving, and swaying are behaviors commonly associated with gamete release. Less common activities include waving of introvert podia, waves of body wall contractions, errant males, aggregations, and pseudocopulation. Positively buoyant eggs of four species [Psolus chitonoides Clark, Cucumaria miniata (Brandt), C. piperata (Stimpson), and C. fallax (Ludwig)] were packaged in pellets, ropes, or strands and released passively; a female Molpadia intermedia (Ludwig) discharged negatively buoyant eggs in a single explosive burst. Two externally brooding sea cucumbers [C. lubrica (Clark) and C. pseudocurata Deichmann] produced strands of dilution-resistant sperm that were carried by currents to nearby females. Three species commonly sympatric [P. chitonoides, C. miniata, and Eupentacta quinquesemita (Selenka)] in high-current rocky habitats have overlapping spring breeding seasons, but spawn at different hours of the day.Please address all requests for reprints to Dr. McEuen at the Harbor Branch Oceanographic Institution     

Development,settling behaviour,metamorphosis and pentacrinoid feeding and growth of the feather star Florometra serratissima

Embryonic development of the northeastern Pacific feather star Florometra serratissima takes place within a ridged fertilization membrane. Cleavage is radial, resulting in a coeloblastula, and gastrulations is by invagination. Cilia are swollen terminally during ciliogenesis whereas fully grown cilia possess several swellings along the length of their shafts. Young doliolaria larvae begin to hatch from the fertilization membranes 35 h after fertilization (9.5° to 11.5°C); by 4 d the doliolaria has acquired ciliated bands, a vestibular invagination and an antero-ventral adhesive pit. The surface of the larva is covered with a delicate glycocalyx supported by microvilli. Larvae swim along a vertical sinusoidal path just below the water surface; they begin to explore the substratum at 4.5 d and settlement begins as early as 4.6 d, but can be delayed for up to 9 more days. Larvae settle gregariously in culture and it is suggested that gregarious settlement plays a role in the formation and maintenance of adult aggregations of F. serratissima. Metamorphosis into a stalked cystidean following settlement is rapid. Major changes at this period include: loss of cilia; withdrawal of ectoderm from the glycocalyx; covering over of the vestibular invagination; and a 90 degree rotation of the vestibule to the former posterior end of the doliolaria. Transformation from cystidean to pentacrinoid includes the opening of the 5 oral plates, the extension of the 15 papillate tube feet and further elongation of the stalk. The pentacrinoid is able to feed on small food particles. Rudiments of all 10 adult arms are present by 4 months; at 6 months the pentacrinoid has an arm span of 6.5 mm but cirri and pinnules are not yet present.     

Observations on the function of the glands of the foot of the pediveliger of Ostrea edulis during settlement

The use during settlement of secretions of the 9 different types of gland in the foot of the pediveliger of Ostrea edulis L. was examined experimentally. During settlement, there is a progressive decrease in the crawling speed of the larva and a strengthening of its bond with the substratum. Both changes are related to (a) a change from ciliary to muscular pedal locomotion; (b) the increasing acidity and viscosity of the mucoid secretions used; (c) the secretion of a byssus thread by the glands of the byssus duet; (d) changes in the form and composition of this byssus. At the termination of settlement, the entire contents of 2 types of gland are discharged. Their secretions give rise to the cement which permanently fixes the larva to the substratum. The 9 types of gland are utilised sequentially in a manner closely related to the 6 phases recognised in settlement behaviour.     

Dispersion of gastrotricha in a marine beach of the San Juan Archipelago,Washington

Gastrotricha were quantitatively sampled at low tide on several dates from two transeets at Eagle Cove, San Juan Island, Washington, USA, during the summers of 1963/1964. They showed low species diversity and very high whole-beach densities. Chaetonotus testiculophorus Hummon, 1966 and Turbanella cornuta Remane, 1925 were found abundantly, Paraturbanella intermedia Wieser, 1957 only rarely. Abundant C. testiculophorus were found above 100 cm elevation; T. cornuta occurred abundantly in a rip-channel trough and in the lower beach area. T. cornuta, but not C. testiculophorus, showed significantly greater clumping at high than at low density. Curves of LC₉₀ temperature-salinity tolerance for both species show that they tolerate widely fluctuating temperatures and salinities. Causes of whole-beach distribution patterns are discussed.Research was supported by an NSF College-University Association Grant (G-17537), administered by the Graduate School, University of Washington, Seattle, and was conducted at the Friday Harbor Laboratories, San Juan Island, Washington, during the summers of 1963 and 1964.     

Microhabitat-associated variability in survival and growth of subtidal solitary ascidians during the first 21 days after settlement

Newly settled juvenile ascidians were transplanted into shaded and unshaded sites at two subtidal depths to test the hypothesis that photonegative settlement behavior enhances juvenile survival. Silt, filamentous algae and grazing gastropods (Margarites pupillus) were identified as sources of mortality associated primarily with non-cryptic microhabitats. Silt inflicted heavy mortality on all six species tested. Algal overgrowth was important in shallow water, where it increased mortality and depressed growth rates over a 14-d period. Laboratory experiments demonstrated that gastropods can remove juvenile ascidians from the substratum. The sources of post-settlement mortality investigated may help determine distributional patterns of ascidians in the San Juan Islands, Washington, USA, and may also represent selective pressures that maintain negative phototaxis in the behavioral repertoires of ascidian tadpoles.     

Reproduction,development and population biology of the Caribbean ophiuroidOphionereis olivacea,a protandric hermaphrodite that broods its young

Reproduction and development of the ophiuroidOphionereis olivacea H. L. Clark, a species that broods its young, were investigated in Florida from January through December 1985 and in Belize in June 1985 and April 1986.O. olivacea has a maximum disc diameter of 5.2 mm and is a protandric hermaphrodite. Sex reversal occurs at disc diameters ranging between 2.2 and 4.0 mm; it has 400 µm diameter oocytes and is ovoviviparous. As expected from an obligate out-crosser, this ophiuroid has a similar allocation to male and female reproduction, in terms of gonad volume. Reproduction of the Florida population was monitored for one year.O. olivacea is a sequential brooder and incubates its young during the coldest months of the year. The females produce a single clutch of juveniles each year. Spawning starts in November and peaks in January and February. The ova are spawned synchronously from all the ovaries of each female into the bursae and are retained there. Fertilization is accomplished by intake of sperm into the female bursae.O. olivacea develops through a ciliated embryonic stage that appears to be a modified vitellaria larva lacking ciliary bands. These modified larvae swim in seawater in the bursae. Direct development in this species has resulted from heterochrony in the loss of ophiopluteus features. The number of brooding females in the population increases in November and reaches a maximum in March. JuvenileO. olivacea emerge from the bursae at a disc diameter of 480 µm and each arm has three segments. Recruitment was first detected in April, and juveniles were particularly abundant in June. Small size inO. olivacea is associated with paedomorphic heterochrony with precocious maturation of the males at a disc diameter of 1.5 mm, ~2 mo after taking up their free existence. Brooding as a life history mode inO. olivacea is examined and the adaptations of this species to its warm-water habitats are discussed.     

Use of pelagic prey subsidies by demersal predators in rocky reefs: insight from movement patterns of lingcod

Allochthonous subsidies of energy and nutrients can affect community structure in patchy marine habitats, including rocky reefs, and their ecological consequences may depend on the mechanism of energy transfer. Lingcod (Ophiodon elongatus) are demersal predators that trophically link nearshore rocky reefs with offshore pelagic habitats through consumption of pelagic fishes. We quantified lingcod habitat use and movement patterns to make inferences about the temporal and spatial conditions under which lingcod may acquire pelagic prey. Lingcod maintained small home ranges (21,272 ± 13,630 m²) within a rocky reef in the San Juan Archipelago, Washington; eight of nine individuals used rocky habitat exclusively. Depths occupied by lingcod (0–50 m) coincided with pelagic fish distribution on the rocky reef; however, diel patterns in lingcod activity varied inversely with occurrence of pelagic fishes on the reef. Our findings suggest that the pelagic subsidy to lingcod is not strongly mediated through directed off-reef foraging by lingcod.     

Utilization of a fecal mass as food by the pelagic mysis larva of the penaeid shrimp Solenocera atlantidis

The undetached peritrophic membrane of the mysis shrimp larva (Solenocera atlantidis) provides a substratum on which microbes, organic debris and planktonic organisms accumulate. Behavioral, metabolic and biochemical data indicate that this fecal mass serves as a concentrated food resource for the larva.     

Ingestion of a medusa (Aegina citrea) by the nematocyst-containing ctenophore Haeckelia rubra (formerly Euchlora rubra): phylogenetic implications

The rare ctenophore Haeckelia rubra (formerly Euchlora rubra) has long been known to have nematocysts rather than colloblasts in its tentacles. Five specimens were collected in the San Juan Archipelago, Washington State, USA in 1980 and 1981, and their feeding behavior was observed in the laboratory. We found that H. rubra readily eats the tentacles of a medusa, Aegina citrea, whose nematocysts (apotrichous isorhizas) are nearly identical in morphology to the nematocysts of the ctenophore. When H. rubra was offered 16 other species of hydromedusae and 1 siphonophore in the laboratory, the ctenophores showed little or no tendency to ingest these potential prey items. In addition to its routinely positive response to A. citrea, the ctenophore could be induced by manipulation and starvation to accept and ingest bits of the bodies of 4 additional species of hydromedusae and 1 siphonophore. These results, combined with the histological and rearing experiments of other investigators, leave little doubt that the nematocysts in H. rubra are not endogenous, but are kleptocnidae similar to those nematocysts retained and subsequently used by some species of nudibranchs that feed on Cnidaria. A close phylogenetic link between the Cnidaria and the Ctenophora is most unlikely.     

Reproductive biology and early life history of the hermaphroditic feather star Dorometra sesokonis (Echinodermata: Crinoidea)

Reproduction and larval/post-larval development of the one of the smallest known comatulid Dorometra sesokonis were studied on the coral reefs of Sesoko-jima Island, Okinawa, Japan. Breeding individuals were found every month from May 2003 to April 2004 indicating continuous reproduction. Individuals possessed both mature ovaries and testes, and were therefore characterized as simultaneous hermaphrodites. This species was observed to be an external brooder in which the embryos developed on the surface of parental genital pinnules until the doliolaria larval stage, and subsequently released doliolariae settled on substrata within 4 days after hatching. Larvae then metamorphosed into the post-larval cystidean stage, which lasted for more than a month. The reproductive features of this species are unique among crinoids, and, together with juvenile morphological features observed in adults, can be interpreted as adaptations to their cryptic habitat, which is subject to frequent natural disturbances.     

Settlement of barnacle larvae: surface structure of the antennular attachment disc by scanning electron microscopy

The surface morphology of the antennular attachment disc of the cypris larva of Balanus balanoides has been investigated by scanning electron microscopy, and its function is considered in the context of settlent behaviour. It is concluded that the attachment organ acts as an adhesive pad for temporary attachment while the larva explores the substratum. The surface of the disc is covered with villi, which could retain an adhesive substance thought to be produced by the antennular glands. The open-ended hair of the axial sense organ on the disc is shown.     

Metamorphosis and developmental evolution in <Emphasis Type="Italic">Ophionereis</Emphasis> (Echinodermata: Ophiuroidea)

The brittle star Ophionereis schayeri has abbreviated non-feeding development through a reduced ophiopluteus and a vitellaria larva. The metamorphic changes involved with development from a bilateral larva to a radial juvenile were examined in detail. The reduced ophiopluteus has a continuous ciliated band that breaks up into discontinuous ciliary ridges at the vitellaria stage. As the vitellaria develops, the juvenile rudiment forms in the mid-ventral region. The rudiment then undergoes a morphogenetic movement to the left of the larval anterior/posterior axis. This results in a dramatic transformation from bilateral to radial symmetry and is accompanied by development of juvenile structures and settlement, 6–7 days after fertilisation. Ophiuroid development through one larval stage, the ophiopluteus, is termed Type I; whereas development through two larval stages, the ophiopluteus and vitellaria, is termed Type II. We examined the evolutionary changes in the expression of Type II development in Ophionereis by comparing the ontogeny of six species with a range of larval forms. O. fasciata has the ancestral-like planktotrophic ophiopluteus. Vestigial pluteal structures in the larvae of O. schayeri, provide a link between ophiopluteal and vitellarial forms during evolution of non-feeding development. The diversity of larval forms in Ophionereis indicates that evolution of non-feeding development through a vitellaria (Type II) may have involved an increase in egg size, reduction of pluteal structures and shortening of the time to metamorphosis. Assessment of the phylogenetic significance of Types I and II development awaits additional comparative data on the metamorphic stages of other ophiuroid genera.Communicated by G.F. Humphrey, Sydney     

Aspects of the reproduction and life cycle of Rhabdopleura compacta (Hemichordata)

Rhabdopleura compacta Hincks is shown to be monoecious. It is suggested that the ovum is ovulated and extruded into the lumen of the coenecium via a temporary opening at the base of the metasome, and that the oviduct itself may serve only as a way for sperm to fertilise the ova internally. The zooids degenerate when reproducing sexually and when conditions are unfavourable. The larva and initial stages of astogeny are described. Some observations on seasonal variation in the colonies indicate that the dormant buds serve for storage of nutrients. Reproduction seems to occur throughout the year, but most actively in spring. Relationships with Cephalodiscus and the Bryozoa are briefly discussed.     

Some observations on gametogenesis,larval development and substratum selection of the sea pen Ptilosarcus guerneyi

Ptilosarcus (Leioptilus) guerneyi (Gray) maintained in the laboratory, were observed to spawn in late March, 1972. Gametes, developed in the leaf proper, discharged through the mouths of feeding polyps and were fertilized externally in the sea water. The sea pen's eggs are 500 to 600 in diameter; a large female is capable of producing over 200,000 eggs in one season. A pear-shaped and free-swimming planula larva developed 4 days after fertilization, at a temperature of 12 °C. The larvae were ready to settle and metamorphose when 7 days old if favorable substratum was available, but would remain as planulae for at least 30 days if kept in glass dishes only. The 30-day-old larvae would metamorphose if a suitable substratum (coarse sand, for example) was presented. The larvae do not feed and, hence, development is lecithotrophic. Studies of histogenesis showed that metamorphosis greatly enhanced the rate of cellular differentiation. The high fecundity, lecithotrophic development, and the ability of substratum selection by the larvae explain the success of this species in maintaining a high-density population in many areas of sandy substratum in the shallow waters of Puget Sound (USA), despite the fact that it is preyed upon by 7 species of predators.     

Unusual lecithotrophic development of the caribbean brittle star Ophiothrix oerstedi

The ophiuroid Ophiothrix oerstedi Lütken spawned in the laboratory at Barbados, West Indies, from August, 1975 until early December. The embryo passes through a wrinkled blastula stage, and the larva is a reduced lecithotrophic ophiopluteus with a shortened pelagic existence. A larva of this type is unusual for brittle stars in general and unique for ophiothricids for which development has been described. Metamorphosis is completed 4.3 days (24° to 27°C) after fertilization with a single pair of ciliated larval arms, the postero-lateral arms, being retained as a swimming device for the late larva. Settlement, with subsequent separation of the postero-lateral arms from the young brittle-star, begins as early as 4.5 days, but can be delayed for at least one week, at the end of which time midwater separation can occur resulting in the pelagic dispersal of post-larvae. A comparison of gametic and larval characteristics of O. oerstedi with the literature suggests that the larva of this species is most closely allied to the abbreviated developers. The adaptive significance of this larval form is discussed.     

Form and feeding mechanism of a living Planctosphaera pelagica (phylum Hemichordata)

We describe aspects of the anatomy and suspension-feeding mechanism of a single Planctosphaera pelagica captured from the plankton in June 1992 off Bermuda in the western Atlantic. We also describe several unusual features of the larva, including its occurrence in surface waters, unusually large size, and limited swimming ability. Our account of the form and feeding behavior of P. pelagica is the first based on observations of a specimen captured and observed alive. Our limited observations suggest that the planctosphaera may use a suspension-feeding mechanism much like that of the other feeding deuterostome larvae (the pluteus and bipinnaria larvae of echinoderms and the tornaria larva of enteropneust hemichordates) known to capture food particles using a single ciliated band. Although we could not observe cilia directly, the movement of dye streams and food particles and the structure of the ciliated band suggest that some particles may be captured at the ciliated band by the reversal of ciliary beat. The planctosphaera possesses many prominent mucous glands near the food grooves. This suggests an important role of mucus in the biology of the larva, but we were not able to observe directly any role of mucus in particle capture.     

Comparative foraging ecology and ecological niche of a superabundant tropical seabird: the sooty tern Sterna fuscata in the southwest Indian Ocean

Over 6-million pairs of sooty terns Sterna fuscata breed once a year in the southwest Indian Ocean, mostly on three islands of the Mozambique Channel (Europa, Juan de Nova and Glorieuses) and in the Seychelles region. Seasonal reproduction in either winter or summer is the dominant strategy in the area, but non-seasonal reproduction also occurred in some places like at Glorieuses Archipelago. The feeding ecology of the sooty tern was investigated during the breeding seasons to determine whether terns showed significant differences in their trophic ecology between locations. Regurgitations were analyzed to describe the diet of individuals when breeding, and stable isotopes and mercury concentrations were used to temporally integrate over the medium-term of the trophic ecology of both adults and chicks. Overall, the diet was composed of fish, flying squid and fish larvae in different proportions. At Europa and Aride in the Seychelles, where winter reproduction occurs, large epipelagic prey like flying fish or squid dominated the diet. At Juan de Nova, sooty terns reproduce in summer and rely mostly on fish larvae. At Glorieuses (non-seasonal breeding), the diet was intermediate with fish larvae and flying squid being important prey items. The stable-carbon and nitrogen isotope values in blood confirm the differences observed in dietary analysis, and demonstrate different feeding strategies between colonies. δ¹³C values of feathers showed spatial segregation between birds from the Mozambique Channel and the Seychelles region. Terns from the Seychelles had also higher δ¹⁵N values. Feather δ¹³C values also suggest a significant shift from summer to wintering habitat for birds from Juan de Nova. This study emphasizes the high phenotypic plasticity of the species, which may explain its numerical dominance in all tropical waters of the World’s Ocean.     

Comparison of larval development and growth of the sea cucumberActinopyga echinites: Ovary-induced ova and DTT-induced ova

AdultActinopyga echinites (Jaeger) were collected from northern Taiwan in September 1989. Oocytes were induced to mature by bathing them in ovary juice (ovary-induced ova) or in 10^–2 M dithiothreitol (DTT-induced ova). The percentage of germinal vesicle breakdown (GVBD) increased from 0.4 to 6.4% in the former treatment and to 84% in the latter. After artificial fertilization, the embryos were cultured in seawater (35 S) at 25 to 28°C. Larvae were fed with the algaIsochysis aff.galbana at 10^4–5 cells/ml. Larvae from ovary-induced ova developed faster (18 d to the doliolaria stage) and grew to a larger size (1.13 mm length) than those from DTT-induced ova (20 d to the doliolaria stage and 0.62 mm in length). On the twelfth day, larvae from ovary-induced ova bear lipid spheres. The number of spheres is positively correlated with larval size. Lipid spheres may provide nutrient reserves for larvae during metamorphosis.     

The larval settling response of the lined chiton Tonicella lineata

The lined chiton Tonicella lineata (Wood, 1815) is found on enerusting coralline algae in the lower rocky intertidal zone of Oregon and San Juan Island, Washington, USA. An encrusting coralline alga is the major food item of this chiton. Experiments were performed to test the settling response of T. lineata larvae to various algae and other substrata. In these experiments, the larvae would settle only on pieces of encrusting coralline algae and piecesof ceramic roofing tile soaked in a coralline algal extract. The settling stimulus is probably chemical in nature, and is inactivated by boiling. In laboratory cultures, normal development stops at the trochophore stage (110 to 160 h, depending on the temperature). Metamorphosis and further development will take place only after stttlement on encrusting coralline algae.