Alarm signal response in the brittle star Amphiura filiformis

 Flat fishes, such as the dab Limanda limanda, commonly prey on arms of Amphiura filiformis. We demonstrate in flume experiments that A. filiformis showed a more or less simultaneous response to an up-stream predation by dab on conspecifics: the arms, which are stretched up into the water column when filter feeding, responded by bending down to the sediment surface, and some arms submerged, at least partly, into the sediment. We interpret this as an alarm response. A similar, but significantly weaker response in A. filiformis was also recorded when dab ate Amphiura chiajei. Homogenates of the two Amphiura species and arm pinching – to simulate partial predation – elicited a similar response in A. filiformis, the arms bent to the sediment surface. In contrast, however, the behavioural response of the arms to the homogenates was to protrude further out of the sediment and sweep the surface vigorously. We interpret this as a feeding behaviour. Response to arm pinching was less clear. The ecological consequences and origin of alarm signal response in A. filiformis are discussed. Received: 8 February 1999 / Accepted: 16 July 1999     

Reproduction and population dynamics of the fissiparous brittle star Ophiactis savignyi (Echinodermata: Ophiuroidea)

Reproductive cycle, asexual reproduction, and population dynamics of the fissiparous brittle star, Ophiactis savignyi, which inhabits the exhalant passages of the sponge Haliclona sp. were examined monthly from February 1991 to January 1992 at Wanlitung, southern Taiwan (22°N; 120°E). Mature gametes were found from March to December, but release was mainly in May and June. Sexual recruits were found from May to December, with the highest frequency (14.1% of population) in June. Sexual recruits composed 2.4% of the 1-yr sample. Sex ratio of male to female was 24:1. Fission occurred throughout the year, although the frequency of recently split individuals was lower from January to June (6 to 31%), and higher from July to December (42 to 52%). The occurrence of fission was highest after spawning. Regenerating individuals composed 48.2% of the 1-yr sample. Population density fluctuated greatly during summer due to recruitment by fission and mortality or dispersal due to the stressful environmental conditions. Both sexual and asexual reproduction of O. savignyi were successful at this site.     

Reproductive biology of a deep-sea brittle star Amphiura carchara (Echinodermata: Ophiuroidea)

 New information on the brooding reproduction of Amphiura carchara, and previous studies of several deep-sea congeners, contradict the prevailing notion that direct development is exceptional among deep-sea echinoderms. Over 500 specimens of A. carchara from 2,850 to 4,100-m depths off the coast of California, USA, were studied. The species was found to be gonochoric, although most brooding species of ophiuroids are hermaphroditic. Females each brooded up to 72 embryos at a time, with up to 10 in a single bursa. The embryos of individual adults were often at different stages of development, but those brooded in a single bursa tended to be at the same stage. Thus, the species has characteristics that breach the distinctions between sequential and simultaneous brooding exhibited by shallow-water ophiuroids. The embryos developed from yolky eggs that are large for an ophiuroid, with the mean largest oocytes 0.45 mm, and ranging to 1.28 mm in diameter. Almost all late-stage embryos were positioned with their mouth and arms pressed against the wall of the bursa, possibly to facilitate the uptake of nutrients from the parent. Thus, A. carchara may be matrotrophic. The largest embryos examined had a rudimentary disk skeleton, and arms with four joints and a terminal plate. Emerging juveniles probably differ in disk diameter. The species appears to brood year round, although differences in gonad size, the incidence of brooding, and the relative numbers of early developmental stages in summer and winter samples indicated that there are seasonal trends in reproduction. Received: 14 February 2000 / Accepted: 3 July 2000     

Non-viable seasonal settlement of larvae of the upper bathyal brittle star Ophiocten gracilis in the Rockall Trough abyssal

Re-examination of ine structure of postlarvae previously identified as the abyssal species Ophiura ljungmani (Lyman) in a time series of samples taken from 1975 to 1980 from a 2900 m deep permanent station in the southern Rockall Trough (Northeast Atlantic Ocean) has shown the majority of the large summertime peak of brittle-star postlarvae to belong to a second species. Most were incomplete and probably corpses when collected. The few intact specimens present were identical to postlarvae collected in benthic samples from the neighbouring continental slope, where a dense population of the bathyal species Ophiocten gracilis (G. O. Sars) is present from 600 to 1 200 m depth. Examination of population size-structures indicates synchronous reproduction in January/February and annual recruitment to the population. The presence both of Ophiopluteus ramosus and intact postlarvae of Ophiocten gracilis in midwater plankton samples from the vicinity of the permanent station in March/April are thought to derive either from the adjacent slope population or, less probably, from larvae transported in the North Atlantic Current from the Flemish Cap area off Newfoundland. In deep water, viability of these benthic postlarvae is low, none surviving longer than the following winter in the Rockall Trough. These observations suggest a large seasonal transfer from the slope of non-viable larvae and postlarvae as food to the abyssal benthic community.Communicated by J. Mauchline, Oban     

Phylogeography of the Antarctic planktotrophic brittle star Ophionotus victoriae reveals genetic structure inconsistent with early life history

In the marine environment, connectivity is influenced by physical oceanography as well as life history and behavioral traits, which in combination with historical events such as geology, physical oceanography, and climate, determine population structure. The Antarctic brittle star Ophionotus victoriae develops via a feeding planktonic larval stage, and therefore has potential for long-distance dispersal throughout its Antarctic/subantarctic range. To evaluate this hypothesis, phylogeography of this ecologically dominant species was elucidated by sequence analysis of two mtDNA genes from individuals collected throughout the Antarctic Peninsula and from two subantarctic islands. Counter to expectations of genetic homogeneity, mtDNA data revealed substantial levels of genetic differentiation as well as diversity. Although there were some genetically homogeneous populations, such as those throughout Bransfield Strait, we found O. victoriae to have significant population structure throughout much of the Antarctic Peninsula, with evidence of potential cryptic speciation between the western and eastern Antarctic Peninsula. Furthermore, Antarctic Peninsula populations were genetically distinct from subantarctic island populations. The low levels of connectivity implied for O. victoriae contrast with those found for many other Antarctic benthic taxa, and suggest a complex interplay between oceanography, recent climate history, and larval ecology.     

Lack of genetically-subdivided population structure in Bullia digitalis, a southern African marine gastropod with lecithotrophic development

Larval shell morphology in fossil and present-day gastropods is often used to infer modes of larval development and levels of dispersal. Dispersal ability influences not only genetic population structure, but also is thought to influence a species' geographical range and evolutionary duration. We tested these predictions in Bullia digitalis, a sandy-beach whelk, by examining genetic variability at 33 protein-coding loci in nine samples (N=739) taken in 1984 to 1985 at localities extending over about three-quarters of the geographical range of this species in southern Africa. Females of this species deposit eggs into benthic or brooded capsules in which larvae develop through the trochophore and veliger stages to emerge as crawling juveniles. Scanning electron micrographs confirmed a protoconch morphology typical for gastropods with lecithotrophic larval development. Contrary to expectations, subpopulations of B.␣digitalis had high levels of variability (H=0.102) and lacked a genetically-fragmented structure (=0.013). The lack of a genetically-subdivided population structure would not have been correctly inferred, if this species were known only from well-preserved fossil shells. Indirect estimates of migration between populations based on and the island model of migration, which assumes drift-mutation equilibrium, ranged between 19 and 23 individuals per generation. Either an undescribed mechanism of dispersal facilitates gene flow between populations, or the geographical range of this species has recently expanded to produce the appearance of high levels of gene flow. Gene-frequency distributions showed that relative to four other species of Bullia, populations of B. digitalis were in mutation-drift disequilibrium, with a significant excess of low-frequency alleles that is consistent with a recent rapid expansion from a small population. Also contrary to expectations, this species has a large geographical range (2 400 km) and an apparently long evolutionary history extending 5 to 20 million years, as estimated from an allozyme phylogeny with four other species of Bullia. Received: 15 January 1997 / Accepted: 28 January 1997     

Divide and broadcast: sexual reproduction in the West Indian brittle star Ophiocomella ophiactoides and its relationship to fissiparity

It is known that asexual reproduction by fission is the dominant mode of propagation in the small (disc diameter <5.0 mm) ophiocomid brittle star Ophiocomella ophiactoides (H. L. Clark). This species is, however, able to reproduce sexually as well. Sexual reproduction occurs throughout much or all of the year in a population of O. ophiactoides at Jamaica. Gonads are often present in recently split individuals, indicating that sexual and asexual reproduction could occur simultaneously. Regeneration of new gonads in the new disc half is underway by the time an individual is competent to split again. Unsexable individuals are significantly smaller than males, which in turn are significantly smaller than females. The unsexable individuals appear to be below a threshold size for gonad production. Fecundity is low, with the largest females producing a maximum of about 7 400 eggs of small (mean diameter 80 m) size. Fertilized eggs develop into planktotrophic ophioplutei which are typical of the larvae of other species of ophiocomid brittle stars in morphology, rate of development and duration of planktonic period. Egg size and larval type for O. ophiactoides are characteristic of brittle stars with the planktotrophic mode of development, but egg number is uncharacteristically low. Fissiparity in O. ophiactoides may have evolved in association with small body size, concomitant low fecundity and a high-risk planktotrophic larva as a means of ensuring reliable recruitment. Available data, although sparse, suggest that this hypothesis may be applicable to fissiparous brittle stars in general. Fission in brittle stars has a broad systematic incidence at the familial level which suggests that it has evolved recurrently. The ability both to divide and broadcast appears to be a useful combination for O. ophiactoides: it can recruit reliably at large size to occupy favourable habitats quickly, but, it can still maintain low level dispersal of larvae which could provide escape from deteriorating habitats and the potential to colonize new habitats.Contribution No. 328 of the Discovery Bay Marine Laboratory, University of the West Indies     

Effect of temperature and food availability on reproductive responses of Streblospio benedicti (Polychaeta: Spionidae) with planktotrophic or lecithotrophic development

Streblospio benedicti (Webster) from Tar Landing North Carolina (NC), USA with either planktotrophic or lecithotrophic development were reared under two food levels and three temperature regimes (two mimicking seasonal cycles in NC and one at constant 20°C). During the eight-month experiment no females switched reproductive mode and no significant differences in survivorship or reproductive activity were observed between reproductive types. However, reproductive activity and fecundity-related parameters were subject to influence by food and temperature. Survivorship, body size, and larval production was greater in winter-spring than summer-fall regimes. Higher food levels produced increased survivorship, reproductive activity and egg production in adults with lecithotrophic development but no change in those with planktotrophic development. Body size, egg size, egg number, numbers of larvae per brood pouch, and brood size were strongly correlated in female S. benedicti and most correlation coefficients were similar (or identical) in individuals having planktotrophic and lecithotrophic development. A comparison of egg size and brood size in females from Tar Landing suggests that individuals with the two forms of development package offspring differently but expend approximately equivalent reproductive effort. Larval trophic mode is best viewed as a genetic polymorphism in S. benedicti. Individuals with planktotrophic and lecithotrophic development exhibit similar reproductive responses to environmental variation and there is no evidence for speciation.     

Re-appraisal of age composition,growth and survivorship of the deep-sea brittle star Ophiura ljungmani from size structure in a sample time series from the Rockall Trough

Samples were taken from a 2 900 m deep permanent station (centred on 54°40N; 12°16W) in the southern Rockall Trough, from November 1975 to September 1980, inclusive. Revision of juvenile size frequencies was necessary after removal of postlarvae of an annual non-viable settlement of Ophiocten gracilis (G. O. Sars) that were mistakenly included in a previous analysis. The present study of the revised data shows that settlement of Ophiura ljungmani (Lyman) occurs in summer when disk-size frequencies are dominated by the newly metamorphosed postlarvae. The likely age composition in larger sizes was suggested from the progression of modes along the size axis in juvenile size classes in consecutive were developed from Gaussian mixtures fitted by constrained computer search using, as starting values, component parameter values suggested from the frequency histograms. Choice between competing solutions was made on the basis both of goodness-of-fit and biological plausibility. However, identification of year-class structure amongst sizes >3 to 4 mm disk diameter remained conjectural owing to the low frequencies of adults in the samples. Comparison of the fitted component-means in the time series suggested a marked seasonal growth pattern, with maximal growth in spring. Probable age composition amongst adult sizes was extrapolated from von Bertalanffy growth parameters estimated from the means of juvenile year classes.Communicated by J. Mauchline, Oban     

Physiological and biochemical changes during lecithotrophic larval development and early juvenile growth in the northern stone crab,Lithodes maja (Decapoda: Anomura)

Larvae of the northern stone crab, Lithodes maja L., were reared in the laboratory from hatching to the second crab stage. complete larval development (at constant 9°C) lasted about 7 wk, invariably consisting of three pelagic zoeal stages and a semibenthic Megalopa; only two zoeal stages have been described in the literature. All larval stages are lecithotrophic. First feeding was consistently observed only after metamorphosis, in the first juvenile crab stage. In short intervals (every 1 to 5 d), developmental changes in biomass, B (expressed as: dry weight, W; carbon, C; nitrogen, N; hydrogen, H) and oxygen consumption (respiration, R) were measured in larvae and early juveniles; additionally, protein and carbohydrates were measured, but only in the zoeal stages and early Megalopa. Unusually high C contents (varying between 56 and 61% of W in eggs and freshly hatched Zoea I larvae from 12 different females) and high C:N weight ratios (8 to 11) indicate enhanced initial lipid stores, which are utilized as the major metabolic substrate during both embryonic and lecithotrophic larval development. Predominant degradation of lipids is shown indirectly; the C:N ratio decreased significantly, from 10 (at hatching) to 6 (at metamorphosis), while larval protein decreased only little, from ca. 55% of W (at hatching) to 48% (in the Megalopa). From hatching to metamorphosis, about 27% of the initially present W, 48% of C, 18% of N, and 52% of H were lost. This decrease in larval biomass can be described as an exponential function of development time. The major part of these losses were associated with metabolic energy requirements, while exuvial losses were comparably small. In each of the zoeal stages, only about 1 to 2% of late premoult (LPM) B was shed with the exuvia. The Megalopa, which produces a much thicker, calcified exoskeleton, lost 20% of LPM W, but only 5 to 8% of organic constituents (C, N, H). Much higher exuvial losses were measured in the Crab I stage (51% in W, 21% in C, 5% in N, and 7% in H). Maximum respiration was found in the actively swimming zoeal stages, a minimum in the predominantly benthic, mostly inactive Megalopa. The Crab I stage exhibits also a sluggish behaviour and low R, in spite of beginning food uptake and growth. Immediately after metamorphosis, the juvenile crab gained rapidly in W, in particular in its C fraction. A transitorily steep increase in the C:N ratio indicates a replenishment of partially depleted lipid stores, but also a rapid initial increase of inorganic C in the heavily calcified exoskeleton. Instantaneous rates of growth, assimilation, and net growth efficiency (K ₂) were high during the initial (postmoult) phase in the first juvenile crab stage (C-specific growth rate: 6% d^-1; K ₂:70%), but decreased towards zero values during laterstages of the moulting cycle; metabolism remained practically constant during the Crab I stage. Entirely lecithotrophic larval development from hatching to metamorphosis in L. maja is considered an adaptation to seasonally short and limited planktonic food production in subarctic regions of the northern Atlantic.     

Diel relationships of microbial trophic groups and in situ dissolved carbohydrate dynamics in the caribbean sea

Dissolved total carbohydrate (TCHO), polysaccharide (PCHO), monosaccharide (MCHO) and organic carbon (DOC) were determined at 3-h intervals over 5 diel cycles in the mixed layer of the northwestern Caribbean Sea while following a drogued buoy. These data have been compared to populations of phototrophic (PNAN) and heterotrophic (HNAN) nanoplankton (2–20 m diameter) and heterotrophic bacteria (HBAC) (0.2–2.0 m diameter) estimated by epifluorescence counts, as well as to CO₂, phosphate, chlorophyll a and phaeopigment data determined simultaneously. Two different types of apparent diel dissolved carbohydrate (CHO) patterns were found. On 3 d when no sustained net CO₂ uptake was evident, TCHO and PCHO generally declined during the afternoon and early evening while MCHO tended to increase. On two other days when apparent sustained CO₂ uptake occurred during the day, there were large evening TCHO and PCHO peaks with constant or declining MCHO levels. These accumulations probably resulted from the release of recently produced PCHO from phototrophs. As was found earlier in the Sargasso Sea, PNAN populations were inversely related to PCHO concentrations. The sample to sample fluctuations of PNAN also were inversely related to the apparent rates of change of TCHO and PCHO, possibly due to an inverse relation between the rates of PNAN cell division and CHO excretion. Fluctuations in HBAC populations were inversely correlated with PCHO dynamics and directly related to MCHO variations, possibly due to extracellular hydrolysis of PCHO to MCHO during periods of rapid bacterial growth as well as to net heterotrophic PCHO uptake. A direct relationship between HNAN and TCHO fluctuations suggests the importance of HNAN excretion in the release of dissolved organics. The combined PNAN and HBAC fluctuations accounted for a more significant fraction of the variance in the apparent rates of change of PCHO than did any single population parameter indicating that intimate interactions between the microbial plankton groups are important in the in-situ regulation of CHO dynamics. Total system net TCHO release and uptake rates for 5 d averaged 56 and 53 g C l^-1 d^-1 respectively, assuming that the observed fluctuations resulted from temporal planktonic processes in homogeneous water masses. While the data contain indications that this was the case, this assumption is not definitive.     

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

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

Feeding by a “nonfeeding” larva: uptake of dissolved amino acids from seawater by lecithotrophic larvae of the gastropod Haliotis rufescens

Larvae of the red abalone (Haliotis rufescens Swainson) are functionally incapable of capturing particulate foods. The aim of this study was to determine whether these larvae could acquire energy from seawater in the form of dissolved organic material. Trochophore and veliger larvae were shown to acquire energy by transporting dissolved organic material from seawater. Both larval stages took up all classes of amino acids tested. The influx of radiolabeled alanine represented the net substrate flux, as determined by direct chemical measurement for both trochophore and veliger larvae. Although veliger larvae have a transport system to take up taurine from seawater, a net efflux was observed for this amino acid. The release of taurine occurred independently of the presence of either taurine or other amino acids in the medium. Transported alanine was used in both anabolic and catabolic pathways. The percent of ¹⁴C-alanine in the trichloroacetic acid-insoluble fraction (macromolecules) of veliger larvae ranged from 21 to 56% of the total radioactivity in the larvae. No lipid biosynthesis was detected from ¹⁴C-labeled alanine. Veliger larvae catabolized 15 to 19% of the total alanine taken up and released it as ¹⁴CO₂. The metabolic rate (oxygen consumption) and the rate of amino acid uptake were both determined for the same group of veliger larvae. The percent contribution that the uptake of amino acids, from a total concentration of 1.6 M, made to the metabolic demand of abalone larvae ranged from 39 to 70%. Thus, these lecithotrophic larvae are not energetically independent of their environment, a result which differs from the current view of energy allocation to nonfeeding larvae.Please address all requests for reprints to Dr. Manahan at the University of Southern California     

Post-larval development of the common north-west European brittle stars Ophiura ophiura,O. albida and Acrocnida brachiata (Echinodermata: Ophiuroidea)

The post-larval skeletal plate ontogeny of the ophiuroids Ophiura ophiura (Linnaeus) and Acrocnida brachiata (Montagu) from the recently metamorphosed 0.3 mm disc diameter size to a juvenile size of 2.0 mm disc diameter has been examined using scanning electron microscopy. Post-larval development of O. albida Forbes is briefly described for sizes between 1.0 and 2.0 mm disc diameter. Early post-larval development in the 0.2 to 1.0 mm discsize range involves considerable morphogenetic change and necessitates the use of a developmental series in the identification of newly settled ophiuroids. Comparison of these ophiuroids with other closely related species reveals that the post-larvae are readily distinguishable at even the smallest sizes by their transient post-larval morphologies. Abbreviated larval development in A. brachiata is suggested from the egg size and post-larval form. The ophiuroid post-larvae were collected from the shallow sublittoral of Oxwich Bay, Bristol Channel, UK, during the years 1980 to 1982.     

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)     

Rates of fission, somatic growth and gonadal development of a fissiparous sea star, Allostichaster insignis, in New Zealand

Seasonal changes in population structure and incidence of fission were measured in intertidal and subtidal populations of Allostichaster insignis, a fissiparous sea star. Population size structure was stable over the course of the 1-year study. Sea stars in the subtidal zone attained greater maximum size (mean arm length, R = 35 mm) than those in the intertidal population (20 mm). Fission rates were greatest among small individuals (R < 20 mm). The frequency of fission ranged from 5 to 32% with peaks in early austral summer in the intertidal zone, and in autumn and winter in the subtidal zone. Sexual reproduction occurred in early spring in sea stars larger than 12 mm. The populations were heavily biased toward males. In the laboratory, A. insignis of three size classes (small, R = 9–13 mm; medium, 19–21 mm; and large, 29–31 mm) were fed mussels ad libitum or starved (not fed macroscopic food) for ∼1 year in a 3 × 2 factorial experiment. Small and medium-sized sea stars divided throughout the experiment and the ramets of most individuals regenerated sufficiently to divide again after 6–9 months. Unfed sea stars did not undergo fission (with one exception), had a higher mortality rate, and did not grow. Small, fed sea stars grew significantly faster than medium-sized or large individuals. At the end of the experiment, the pyloric caeca index (a measure of nutritional condition) was greater in fed than in unfed animals. Gonads (only testes were observed) developed in medium-sized and large, fed sea stars. Our field and laboratory results indicate that asexual reproduction in A. insignis predominantly occurs in small, well-nourished individuals. Ramets grow gradually through repeated fission and regeneration to a size (mean length of regenerating arms, R _r ∼ 20 mm) at which they begin to switch to sexual reproduction as the dominant reproductive mode.     

Effects of temporary starvation on larvae of the sea star Asterina miniata

Patchy food distribution may force temporary starvation conditions on planktonic larvae. This potential food limitation may affect survivorship, duration of larval period, and post-metamorphic succes. In this study, larvae of the asteroid Asterina miniata were subjected to temporary food deprivation of several durations and at different stages. Developmental effects were documented by quantification of larval stage, total length, time to metamorphosis, initial juvenile radius, range of settling times, and percent survival to metamorphosis. All starved treatments were significantly affected in settling time and most in percent survival. However, larvae starved later in development demonstrated tremendous tolerance of food deprivation (e.g. the total number of settlers in the treatment starved for 28 d was not significantly different from the fed control). Survival was lower in treatments starved earlier in development than those starved later. Food is apparently required until late in larval development to facilitate metamorphosis. The range of settling times was large; for example, the continuously-fed control treatment produced juveniles from Days 58 through 136. Temporary starvation had no effect on initial juvenile radius.     

Slow developing demersal embryos and larvae of the antarctic sea star Odontaster validus

The early development of Odontaster validus at McMurdo Sound, Antarctica, is indirect and includes equal cleavage, a convoluted blastula, a free-swimming coeloblastula, a gastrula, and a feeding bipinnaria larva. Development differs from that of other asteroids in two respects: (1) The developmental rate is extremely slow; blastulae form nearly 2 days after fertilization, gastrulation begins after 7 days, and the bipinnaria develops in about 40 to 55 days. The slow developmental rate appears to be only partly related to the low environmental temperature (-1.5°C). (2) The embryos and larvae are largely demersal. Such behavior may be an adaptation to keep the larvae out of antarctic surface waters, as does brooding in many other polar echinoderms.     

Photoperiodic regulation of gametogenesis and gonadal growth in the sea star Pisaster ochraceus

A photoperiod regime 6 mo out of phase between December 1978 and August 1980 resulted in gametogenesis, gonadal growth, and spawning in laboratory-maintained sea stars [Pisaster ochraceus (Brandt)] 6 mo out of phase with individuals in the field or laboratory on normal celestial photoperiods at Santa Cruz, California (USA). The seasonal fluctuation of pyloric cecum size also was shifted when the sea stars were held on the out-of-phase photoperiod regime. Phase shift of all these events was evident within 6 to 9 mo. Long daylengths (or short nightlengths) in spring and summer apparently synchronize or entrain the initiaion of gametogenesis and gonadal growth in fall.