Population structure and energetics of the shallow-water antarctic sea star Odontaster validus in contrasting habitats

Individuals and populations of Odontaster validus Koehler differed markedly among different habitats, as revealed in a study from October 1984 through January 1986 in McMurdo Sound, Antarctica. At McMurdo Station, individual sizes (wet weight) and population biomass (g wet wt m^-2 and kJ m^-2) decreased significantly with increasing depth. Individuals from shallow (10 to 20 m) habitats were in superior nutritional condition to those from deeper water (30 and 165 m), as shown by higher gonad and pyloric cecum indexes, and by higher lipid and energetic levels in the pyloric ceca. Moreover, gonadal output (reproductive output) was higher in shallow-water individuals. Higher levels of chlorophyll in the pyloric ceca and richer yellow to red coloration of the body wall in the shallow-water individuals indicate that they utilize the higher levels of primary production at shallow depths. At East Cape Armitage, where nearly permanent, thick, snow-covered ice most of the year resulted in very low levels of benthic primary production, the lowdensity sea stars were all very small and nutritionally similar to the deep-water individuals at McMurdo Station. At Cape Evans, where the generally snow-free sea-ice that broke up in mid-summer resulted in a luxurient benthic cover of diatoms and macroalgae, the sea stars were smaller than at McMurdo Station at comparable depths, but population densities were higher, resulting in 4 to 9 times greater biomass. Growth rates of sea stars fed in the laboratory were very low, especially compared to laboratory-reared temperate and tropical species; well-fed individuals need about 9 yr to reach 30 g wet weight, near the mean size of shallowwater individuals at McMurdo Station. No growth was detected in individuals caged at McMurdo Station for one year, suggesting even lower growth rates in the field. The stable size-frequency distributions at the different sites and depths throughout the year-long study suggest highly stable populations with low temporal variability in recruitment, migration and mortality. These data indicate that individuals and populations of O. validus quantitatively and qualitatively reflect the general level of productivity of a habitat. Differences noted in size, coloration, nutrition, and reproductive effort may be the result of long-term integration of local levels of primary production. These ubiquitous sea stars may serve as a biotic indicator of productivity in localized habitats around the continental shelf of Antarctica.     

A laboratory study of behavioral interactions of the Antarctic keystone sea star Odontaster validus with three sympatric predatory sea stars

The circumpolar sea star Odontaster validus is ubiquitous in the nearshore marine benthos of Antarctica. Despite its ecological importance, little is known of its behavioral interactions with other common sympatric sea stars. To examine these interactions we employed time-lapse video analyses conducted in a large laboratory tank (1.8 m diameter circular tank, 1,629 L). In each experimental trial, 34 adult O. validus were placed in a tight circular grouping on one side of the tank, and one adult individual of one of three common sympatric species of predatory sea star (Labidiaster annulatus, Diplasterias brandti, or Perknaster aurorae) was placed on the opposite side of the tank. Digital images of sea star movements were then captured at one min intervals over a 24 h period and aspects of sea star movements subsequently analyzed. Each 24 h treatment was replicated three times, as was a control treatment consisting only of O. validus. O. validus had significantly elevated levels of activity in the presence of P. aurorae when compared with the other two sea stars (potential chemically mediated response), and displayed a distinct “flight response” (change in direction and twofold to sixfold increase of speed) upon tactile contact with this species. Moreover, an “alarm response” was detected when individuals of O. validus that encountered a fleeing conspecific also fled the vicinity. In contrast, our results indicated that O. validus displays virtually no chemical or tactile behavioral responses to the large multi-armed L. annulatus and only weak tactile responses to D. brandti.     

Seasonality of reproduction in the cushion star Odontaster validus at Signy Island, Antarctica

The reproductive and larval biology of the common Antarctic cushion-star Odontaster validus Koehler (Echinodermata: Asteroidea) were studied in a shallow-water population at Signy Island in the maritime Antarctic. The gonad index peaked in April/May, and injection with 1-methyladenine indicated that competence to spawn peaked in May/June. Gastrulae were first detected in May, and maximal water-column larval densities were measured in July. Reproduction was therefore highly seasonal, although not all individuals appeared to spawn each year. The pyloric caeca index increased in summer (typically November to March, but variable between years) and decreased during winter (June to October), suggesting that feeding activity was seasonal. All aspects of biology investigated showed significant interannual variability. Comparison with previous studies in the high Antarctic at McMurdo Sound indicated a similar if more pronounced seasonality at Signy, but the warmer seawater temperatures in the maritime Antarctic (particularly in late winter/early spring from September onwards) resulted in faster larval development at Signy. Received: 21 April 1997 / Accepted: 28 January 1998     

Temperature effects on the metabolism of larvae of the Antarctic starfish Odontaster validus, using a novel micro-respirometry method

Oxygen consumption of individual larvae of the Antarctic sea-star Odontaster validus was measured during the 50-day period following fertilisation. Values ranged from 0.76 pmol O₂ h^-1 for one specimen at the coeloblastula stage to 77.6 pmol O₂ h^-1 for one bipinnaria larva. At 0°C the mean oxygen consumption rate of an individual larva increased from 10.9 pmol O₂ h^-1 (standard error of the mean, SEM, 0.13) for a gastrula larva, 13 days post-fertilisation, to 25.4 pmol O₂ h^-1 (SEM 3.5) at the bipinnaria stage (50 days post-fertilisation). Gastrulae reared at -0.5°C did not have significantly different oxygen consumption rates between days 13 and 45 post-fertilisation (mean=11.4 pmol O₂ h^-1). Individual metabolic rates were highly variable, covering more than a 40-fold range. At 2°C gastrula oxygen consumption was on average 45% higher (17.35 pmol O₂ h^-1), giving a Q₁₀ temperature effect of 4.4. For bipinnaria, mean oxygen consumption in 2°C larvae (31.4 pmol O₂ h^-1) was not significantly different from that in larvae at -0.5°C, suggesting bipinnaria metabolism may be less sensitive to temperature change than earlier stages. At 2°C the bipinnaria stage was reached at 30-35 days compared with 45-50 days at 0°C, giving a Q₁₀ of 4.5 for temperature effects on development. The method here used a new, highly sensitive micro-respirometry method that is inexpensive and straightforward in design. Individual larvae of O. validus were held in 35- to 50-µl respirometers. These larvae have very low metabolic rates, and published work on such organisms have utilised at least 25 individuals per chamber. The oxygen content of the respirometers was measured using a 25-µl sample injected into a couloximeter. Oxygen consumption rates down to -1 pmol h^-1 can be detected. Under optimum conditions oxygen consumption of a single larva of -4 pmol O₂ h^-1 was measured with an accuracy of ᆨ%. Values of ~15 pmol h^-1 could routinely be measured with this accuracy. This method would allow oxygen consumption to be evaluated in individual field-caught larvae of most marine ectotherms.     

Chemical defense in the eggs and embryos of antarctic sea stars (Echinodermata)

Fifteen species of reproductively mature echinoderms (11 sea stars, 3 sea urchins, 1 sea cucumber) were collected from McMurdo Sound, Antarctica, during austral spring and summer of 1985 and 1986; eggs and embryos were obtained, and were tested for ichthyonoxicity using the common marine killifishFundulus grandis as a model predator. Chemical deterrents occurred in the large, yolky eggs of the pelagic lecithotrophic sea starPerknaster fuscus and the planktotrophic sea starPorania antarctica. Brooded embryos of the sea starsDiplasterias brucei andNotasterias armata were also noxious. Significant ichthyonoxicity was not detected in the remaining 7 species of sea stars, 3 sea urchins, and 1 sea cucumber. Chemical deterrents were generally effective at concentrations below a single egg or embryo per agar test-pellet. Although chemicals found in these eggs and embryos are noxious to an allopatric fish, they may not be noxious to sympatric fish.     

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.     

Symbiosis between bacteria and oceanic clonal sea star larvae in the western North Atlantic Ocean

Bacteria were present in the majority of clonal sea star bipinnariae (Luidia sp.) collected between June 1987 and August 1990 from oceanic populations in the subtropical western North Atlantic Ocean. Light and electron microscopy revealed dense aggregations of bacteria within lobes of the epidermal cuticle in the gastric region of larvae. Gram-negative rods and less-common elongate spiral bacteria were observed, always in close association with branching epidermal microvilli. Intact and partially digested bacteria occurred in vesicles and phagosomes within epidermal cells, a probable indication of phagocytotic activity by the host. The association of larvae of Luidia sp. and bacteria ranged widely on both geographical and temporal scales. During July and August 1988, larvae were abundant (1.6 to 11.1 m³) in the mixed surface layer (<10 to 100 m) of stations in the Gulf Stream, the Sargasso Sea and the North Equatorial Drift, and 96% (mean of 5 stations, n=8 to 10 larvae in each station) of the individuals surveyed harbored bacteria beneath their cuticle. In a single station on the Florida Current sampled during winter, spring, and summer months between 1989 and 1991, 79 to 90% of the larvae harbored bacteria. The incidence of symbiosis was higher in actively cloning larvae than in non-cloning larvae. This is the first documentation of a symbiosis in field populations of echinoderm larvae.     

Calorespirometry of developing embryos and yolk-sac larvae of turbot (Scophthalmus maximus)

The rates of oxygen consumption and heat dissipation were simultaneously measured and related to contents of glucose, glycogen and lactate in order to determine whether anaerobic processes contributed significantly to the energy metabolism of developing turbot embryos and larvae. The results suggest that metabolism is fully aerobic between Days 0 and 12 post fertilisation. The data further suggest that glycogen is the sole metabolic fuel during the first 18 to 19 h post fertilisation. After the commencement of epiboly, carbohydrates play an insignificant role in the energy metabolism of the developing embryo and yolk-sac larva.     

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)     

Dislocation experiments in the migrating sea star Astropecten jonstoni

Movements of tagged specimens of the sea star Astropecten jonstoni (D. Ch.), both spontaneous and after displacement, were studied by means of SCUBA diving in shallow coastal waters of Sardinia (Mediterranean Sea). In spring, the entire population nigrates towards the shore. Displaced sea stars were able to return to their original depth zone. Both migration and return to the original depth zone occur in a unidirectional way.     

Reproductive pattern in the brooding and broadcasting sea star Pteraster militaris

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

Reproduction and genetic variation in the deposit-feeding sea star Ctenodiscus crispatus

The gonad index in the deposit-feeding asteroid Ctenodiscus crispatus (Retzius) in the Gulf of Maine (USA) is seasonally less variable than in any other sea star, ranging from 2.99 to 4.98% of dry body weight in females and from 2.28 to 3.42% in males, and varies in concert with, rather than reciprocal to, the pyloric caecum index. Biochemical composition and, hence, caloric content, of the gonads also show little seasonal change, suggesting that reproduction is aseasonal and continuous in this population. Oocyte development is asynchronous, all females having a full size range of oocytes (from less than 30 m to greater than 400 m diameter) throughout the year. Seasonally determined size-frequency distributions of juveniles, oocyte cytology and size-frequency distributions, responsiveness of adult females to 1-methyladenine, and oxygen uptake rates indicate that variations in reproductive intensity are superimposed on continuous reproduction, and seem related to changes in phytoplankton production rather than to temperature. The rich neutral lipid content (ca. 50% of total lipid) and large egg size (>400 m) in the ovaries suggest that development is direct. The population is extremely variable genetically, polymorphism among 13 enzyme-coding genes being 77% and average heterozygosity being 0.174. The reproductive pattern and genetic variation in the eurybathic C. crispatus are similar to those in deep-sea echinoderms. This may be related to the constancy of the population's detrital food source, to small-scale heterogeneity of its physical environment, and to low individual vagility.     

Physiological energetics of developing embryos and yolk-sac larvae of Atlantic cod (Gadus morhua). I. Respiration and nitrogen metabolism

This paper provides the basis for a general model of catabolic metabolism for developing embryos and yolk-sac larvae of Atlantic cod (Gadus morhua L.). Yolk-dependent routine rates of oxygen consumption, ammonia excretion, and accumulation of ammonium ions were related to quantitative changes in contents of glucose, glycogen, lactate, free amino acids, proteins and lipid classes (lipid classes published separately) in order to determine the rate and sequence of catabolic substrate oxidation that occurs with development at 6.0°C, 34.5 S. The stoichiometric relation of the oxygen consumption and total ammonia production to substrate utilisation indicated that during the first 2 to 3 d of development, glycogen was the sole substrate of oxidative metabolism. After formation of the syncytium, free amino acids (75%) together with polar lipids (13%, mainly phosphatidyl choline) and neutral lipids (9%, mainly triacylglycerol) comprised the metabolic fuels of embryonic development. Following hatch (Day 16 post fertilisation), the fuels were free amino acids (32%), polar lipids (20%, mainly phosphatidyl choline), neutral lipids (17%, mainly triacylglycerol) and proteins (31%). Thus, the catabolic metabolism of endogenously feeding Atlantic cod larvae was predominantly fuelled by amino acids (67%) and lipids (32%), while glycogen only accounted for 1% of the total enthalpy dissipated. It is proposed that the above sequence of catabolic substrate oxidation is also generally applicable to other cold-water fishes which spawn eggs that do not contain oil glubules.     

Physiological energetics of developing embryos and yolk-sac larvae of Atlantic cod (Gadus morhua). II. Lipid metabolism and enthalpy balance

This paper presents quantitative data for the changes in the contents of total lipids, lipid classes and their associated fatty acids, together with the changes in caloric contents of developing eggs and yolk-sac larvae of Atlantic cod (Gadus morhua L.). During development between Day 0 to 28 post fertilisation, 32% of catabolic metabolism was fuelled by lipids. On a mass-specific basis, polar lipids (mainly phosphatidyl choline) contributed 60%, and neutral lipids (mainly triacyl-glycerol) contributed 40% to this catabolic component, with each class supplying similar amounts of fatty acids as fuel. The fatty acids supplied by phosphatidyl choline were catabolised non-selectively (i.e. in proportion to their presence in the egg), with about half of them being polyunsaturated. However, of the fatty acids esterified in triacylglycerol, the larvae showed an apparent oxidation preference for monoenes over polyunsaturates or saturates. Routine rates of oxygen consumption and ammonia production were related to the caloric contents of the eggs and larvae in order to derive an enthalpy balance equation (of the form P=C+R+E) for an Atlantic cod larva during its period of endogenous nutrition. For the interval of Day 0 to 25 post fertilisation (the period of yolk dependence), integration of the physiological and caloric data revealed that Atlantic cod larvae conserved 53% of yolk enthalpy (C) for growth (P), 42% was dissipated due to metabolism (R) while only 5% was lost via excretion (E).     

Palatability of invertebrate larvae to corals and sea anemones

Risk of larval mortality is an underlying theme in debates and models concerning the ecology and evolution of the differing reproductive characteristics among marine benthic invertebrates. In these discussions, predation is often assumed to be a major source of larval mortality. Previous studies, focused primarily on planktotrophic larvae, suggested that marine larvae generally were susceptible to, and poorly defended against, planktivorous fishes and invertebrates. Larval-planktivore interactions involving larger and more conspicuous lecithotrophic larvae that are typical of many brooding sessile invertebrates have not been well studied. This lack of data for diverse larval types has hindered testing broad generalities about marine larvae and planktivore prey-preferences. This study demonstrates that lecithotrophic larvae of many Caribbean and temperate western Atlantic invertebrates are distasteful to co-occurring corals and anemones. These larval predators frequently rejected larvae of sponges (6 of 9 species), gorgonians (7 of 9 species), corals (3 of 3 species), hydroids (2 of 2 species) and a bryozoan. Larvae of three temperate colonial ascidians were readily consumed. Frequencies of survivorship for larvae captured but rejected by corals and anemones were generally high and, in 20 of 24 assays, were not statistically different from those of unattacked control larvae. Levels of metamorphosis (when it occurred) of rejected larvae also rarely differed significantly from those of unattacked controls. These results provide further evidence that larval palatability to predators may not be as high as once thought, particularly for brooded larvae of sessile colonial invertebrates. The means by which larvae may avoid or deter predators, and the demographic consequences for marine invertebrates and for the evolution of invertebrate life-history patterns, need to be assessed.     

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.     

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.     

In situ ontogeny of behaviour in pelagic larvae of three temperate, marine, demersal fishes

The ontogeny of behaviour relevant to dispersal was studied in situ with reared pelagic larvae of three warm temperate, marine, demersal fishes: Argyrosomus japonicus (Sciaenidae), Acanthopagrus australis and Pagrus auratus (both Sparidae). Larvae of 5–14 mm SL were released in the sea, and their swimming speed, depth and direction were observed by divers. Behaviour differed among species, and to some extent, among locations. Swimming speed increased linearly at 0.4–2.0 cm s⁻¹ per mm size, depending on species. The sciaenid was slower than the sparids by 2–6 cm s⁻¹ at any size, but uniquely, it swam faster in a sheltered bay than in the ocean. Mean speeds were 4–10 body lengths s⁻¹. At settlement size, mean speed was 5–10 cm s⁻¹, and the best performing individuals swam up to twice the mean speed. In situ swimming speed was linearly correlated (R ²=0.72) with a laboratory measure of swimming speed (critical speed): the slope of the relationship was 0.32, but due to a non-zero intercept, overall, in situ speed was 25% of critical speed. Ontogenetic vertical migrations of several metres were found in all three species: the sciaenid and one sparid descended, whereas the other sparid ascended to the surface. Overall, 74–84% of individual larvae swam in a non-random way, and the frequency of directional individuals did not change ontogenetically. Indications of ontogenetic change in orientated swimming (i.e. the direction of non-random swimming) were found in all three species, with orientated swimming having developed in the sparids by about 8 mm. One sparid swam W (towards shore) when <10 mm, and changed direction towards NE (parallel to shore) when >10 mm. These results are consistent with limited in situ observations of settlement-stage wild larvae of the two sparids. In situ, larvae of these three species have swimming, depth determination and orientation behaviour sufficiently well developed to substantially influence dispersal trajectories for most of their pelagic period.     

Effects of oil dispersants on the developing embryos of marine fish

The effects of oil dispersants BP1002¹, Finasol ESK and Corexit 7664 on the development of herring, Clupea harengus L., plaice, Pleuronectes platessa L. and sole, Solea solea (L.) have been studied. Treatment of developing embryos for 100 h with BP1002 and Finasol ESK gave rise to abnormalities in cell division and differentiation, to reductions in heart-rate, eye pigmentation, growth rate and hatching success above concentrations of 10 parts/10⁶. Larvae with abnormal flexures of the spine, which prevented them from feeding successfully, resulted from exposure to 5 parts/10⁶. Treatment of the embryos with dispersants throughout the period from fertilisation to hatching (15 to 20 days) produced similar abnormalities at slightly lower concentrations. Corexit did not produce any demonstrable deleterious effects on embryos exposed to concentrations up to 5000 parts/10⁶, the highest tested.