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.     

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.     

Feeding aggregations of sea stars (<Emphasis Type="Italic">Asterias</Emphasis> spp. and <Emphasis Type="Italic">Henricia sanguinolenta</Emphasis>) associated with sea urchin (<Emphasis Type="Italic">Strongylocentrotus droebachiensis</Emphasis>) grazing fronts in Nova Scotia

Migrating feeding aggregations (or fronts) of sea urchins can dramatically alter subtidal seascapes by destructively grazing macrophytes. While direct effects of urchin fronts on macrophytes (particularly kelps) are well documented, indirect effects on associated fauna are largely unknown. Secondary aggregations of predators and scavengers form around fronts of Strongylocentrotus droebachiensis in Nova Scotia. We recorded mean densities of the sea stars Asterias spp. (mainly A. rubens) and Henricia sanguinolenta of up to 11.6 and 1.7 individuals 0.25 m⁻² along an urchin front over 1 year. For Asterias, mean density at the front was 7 and 15 times greater than in the kelp bed and adjacent barrens, respectively. There was strong concordance between locations of peak density of urchins and sea stars (Asterias r = 0.98; H. sanguinolenta r = 0.97) along transects across the kelp–barrens interface, indicating that sea star aggregations migrated along with the urchin front at rates of up to 2.5 m per month. Size–frequency distributions suggest that Asterias at the front were drawn from both the barrens (smaller individuals) and the kelp bed (larger individuals). These sea stars fed intensively on mussels on kelp holdfasts and in adjacent patches. Urchin grazing may precipitate aggregations of sea stars and other predators or scavengers by incidentally consuming or damaging mussels and other small invertebrates, and thereby releasing a strong odor cue. Consumption of protective holdfasts and turf algae by urchins could facilitate feeding by these consumers, which may obtain a substantial energy subsidy during destructive grazing events.     

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.     

Surface sequestration of chemical feeding deterrents in the Antarctic sponge<Emphasis Type="Italic"> Latrunculia apicalis</Emphasis> as an optimal defense against sea star spongivory

Latrunculia apicalis is a spherically shaped demosponge that previous investigations have shown is rarely preyed upon by sea stars which are the dominant spongivores in antarctic benthic communities. Prior studies have also demonstrated that L. apicalis produces organic compounds that elicit a tube foot retraction response in the keystone spongivorous sea star Perknaster fuscus that can be used as a reliable assay for feeding deterrence. L. apicalis is known to contain discorhabdin alkaloids which serve, among other roles, as the source of its green coloration. To assess the defensive nature of the discorhabdin alkaloids toward P. fuscus, we have determined discorhabdin G concentrations in discrete sponge layers and evaluated those concentrations in the P. fuscus bioassay. In discorhabdin G-bearing sponges, we found a gradient of discorhabdin G that falls off rapidly toward the center of the sponge. On average, 52% of total discorhabdin G in a given sponge was found within 2 mm of the sponge surface. Tube foot retraction responses to extracts from the surface tissues (0–2 mm depth) of L. apicalis were compared to those of an inner layer (8–10 mm depth) and to a sample comprised of the same inner layer spiked with discorhabdin G at a concentration equivalent to that of the surface tissues. Tube foot retraction response times to extracts of the surface layers and the spiked inner layers were not statistically different, but were significantly greater than responses to the unaltered inner layer and controls. These results support the predictions of the optimal defense theory as L. apicalis sequesters its defensive chemistry (discorhabdin G) in its most vulnerable surface tissues, where the likelihood of predation from sea stars is highest. As antarctic sponges are generally preyed upon by extraoral feeding sea stars rather than deeper biting predators such as fish, surface sequestration may be uniquely adaptive in sessile macroinvertebrates occurring in antarctic marine benthic environments.Communicated by P.W. Sammarco, Chauvin     

Encounters between Antarctic limpets, <Emphasis Type="Italic">Nacella concinna</Emphasis>, and predatory sea stars, <Emphasis Type="Italic">Lysasterias</Emphasis> sp., in laboratory and field experiments

Antarctic limpets, Nacella concinna, from the Admiralty Bay (King George Island, South Shetlands) for at least part of the year (austral winter) co-exist with predatory sea stars Lysasterias sp. Our laboratory and field experiments established that the presence of Lysasterias sp. or its odour had considerable influence upon their behaviour. Limpets’ responses, consisting of shell mushrooming, shell rotation and flight, were distinctly different from their reaction to other stimuli, such as food and conspecific odours, or mechanical stimulation. Moreover, a significant impact of sea star presence on limpets’ activity was observed, with limpets fleeing to a distance of 60 cm from the predator. Such reactions allow limpets to lower the incidence of sea star predation, but at the cost of presumptive disrupting of foraging and an additional energy expended for locomotion. A visible difference was noted between two limpet populations, with the rockpool limpets responding only after physical contact with being touched by a sea star, and the subtidal ones responding at a distance of up to 20 cm.     

Settlement, recruitment and potential predators and competitors of juvenile echinoderms in the rocky subtidal zone

Recruitment patterns of marine invertebrates are affected both by settlement and early post-settlement events. This study examined the settlement and recruitment patterns of echinoderms at three sites in the rocky subtidal zone of Bocabec Cove, Bay of Fundy, Canada using artificial turf collectors and quadrats on the natural substrate. Potential predators were quantified at two of the sites along transects and in 1-m² quadrats. Both potential predators and competitors were quantified in 0.0625-m² quadrats. Settlement varied across sites (1.5–3 km apart) and two years of sampling (2004, 2005). The site of most potential settlement differed for the three groups of echinoderms: sea urchins (Strongylocentrotus droebachiensis), sea stars (Asterias spp.) and sea cucumber (Psolus fabricii). Settlement densities on the artificial turf collectors tended to be greater than the densities of settlers on the natural substrate. On the natural substrate, the only significant difference between densities of juveniles over time was that newly settled sea stars were found in July and were not found the following October. Large lobsters and carnivorous worms were potential predators with densities that varied between sites. Potential competitors that differed in abundance between sites were herbivorous gastropods and conspecifics for sea urchins; and carnivorous worms for sea stars. This study suggests that patterns of recruitment are either set up by patterns of settlement or by events during the first few weeks/months on the benthic substrate for these echinoderms.     

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.     

The types of escape reactions elicited in the scallopPecten maximus by selected sea-star species

Pecten maximus (L.), when stimulated by contact with various sea-star species, displays several fairly distinct types of responses. These range from valve closure, through jumping to the well-known swimming escape reaction. These responses are described in detail. The most violent responses are evoked by contact with the predatory sea starsAsterias rubens,Astropecten irregularis, andMarthasterias glacialis, whereas non-predatory sea stars may evoke minor reactions or none at all. It is suggested that, in this instance, the adaptive value of the various types of responses lies not only in enabling the scallop to escape from predators, but also in preventing needless and even hazardous movement away from the refuge of its prepared depression in the substratum when confronted by the less harmful species of sea stars. Crude chemical extracts of all sea stars always evoked escape reactions, but only those ofAsterias rubens andAstropecten irregularis caused the full swimming response.     

Chemical mediation of invertebrate defensive behaviors and the ability to distinguish between foraging and inactive predators

When the predatory sea star Pycnopodia helianthoides was placed upstream, the sea urchin Strongylocentrotus purpuratus responded defensively by extending and opening its globiferous pedicellariae. No pedicellaria response was given in control seawater or when the sea star was downstream. The snail Tegula funebralis responded by moving up vertical surfaces when Pycnopodia helianthoides or when Pisaster ochraceus were placed upstream. When these sea stars were introduced downstream, the snail's response was not significantly different from that in control seawater. Water collected from an aquarium containing a single sea star was sufficient to trigger the response of S. purpuratus and T. funebralis; the physical presence of the sea star was not essential. This indicated that a chemical stimulus was involved, and the lack of responses when sea stars were downstream argued strongly against the possible additional involvement of visual or vibrational stimuli. S. purpuratus gave stronger pedicellaria responses to water flowing over an active Pycnopodia helianthoides than to water flowing over the same sea star when it was inactive. The significance of the ability to distinguish between actively foraging and inactive predators is discussed, and a mechanism is proposed to explain differences in the amount of stimulatory chemicals released by active and inactive sea stars.     

Emergence pattern of loggerhead turtle (<Emphasis Type="Italic">Caretta caretta</Emphasis>) hatchlings from Kyparissia Bay,Greece

The present study describes the emergence pattern of loggerhead sea turtle hatchlings (Caretta caretta) from a nesting beach in Kyparissia Bay (Greece). We try to establish the role played by hatchling biometry, nest relocation and distance from nest to the sea on this emergence pattern. We surveyed a total of 32 nests, and found long emergence periods (mean = 6.7 nights). The majority of emergences occurred at night, mainly between 0030 and 0100 hours, and in small groups. Most of the hatchlings emerged from the nests the first night. We found no clear trend when we studied the effect of hatchling biometry between successive emergence days. We also found that relocation of the nests did not significantly affect the emergence pattern. However, we noted that in the relocated nests, hatchlings emerged in smaller groups. Emergence periods were inversely related to distance from the sea. In short, factors such as climate conditions, relocation and nest distance to the sea appear to have some effect on the emergence pattern. Therefore, they should be taken into account in both biological studies and management plans for sea turtle nesting beaches. Our results suggest leaving an extended period between the first emergence of hatchlings and the excavation of nests by researchers in future studies in the area.     

Observations in captivity of the activity patterns and resources utilization of the spider crab <Emphasis Type="Italic">Inachus phalangium</Emphasis> (Decapoda,Majidae)

The spider crab Inachus phalangium is common in the sublitoral fringe of the Mediterranean Sea and north-eastern Atlantic Ocean, where it can be found in association with the snakelocks sea anemone Anemonia viridis. Studies concerning its activity patterns and the role of the host sea anemone are lacking. Our study aimed at investigating activity rhythms and resources utilization of I. phalangium reared in captivity. The main behavioral traits exhibited by I. phalangium are performed mostly at night. Two experiments were designed, one examined the time budget of various behavioral acts and the degree of association with the sea anemone, the other analyzed the behavioral response to algae and anemones. We showed that algae have a crucial role in the biology of I. phalangium and that crabs are ready to leave the protection of their host to obtain them. Algae represent both the major component of the diet and one of the most utilized sources of masking material of I. phalangium, which provide, together with specialized cryptic behaviors, protection against predators. Although our data suggest that the association with A. viridis is not obligatory, but the role of the snakelocks sea anemone in the life of I. phalangium is still central, both as an anti-predatory defense and as a nutritional source. The association of I. phalangium with algae and the anemone is a facultative biotrophic commensalistic symbiosis.     

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.     

Pleistocene speciation of sister taxa in a North Pacific clade of brooding sea stars (<Emphasis Type="Italic">Leptasterias</Emphasis>)

Although numerous coastal marine species show intra-specific lineage splitting and population divergence times that date to the period of glacial cycles during the Pleistocene epoch, reported instances of recent speciation in the coastal marine environment are relatively rare. Marine organisms with brood-protection and other reproductive modes with limited dispersal potential have been suggested to experience more frequent speciation and extinction events than related species with higher dispersal rates, but few studies have actually estimated divergence times of sister species in these organisms. Here, two mitochondrial gene regions (cytochrome oxidase subunit I, putative control region and upstream tRNAs) and a nuclear gene region (Elongation factor 1α subunit intron 4) provide evidence of recent (0.5–1.2 Mya) cladogenetic events in four pairs of putative sister taxa in a predominantly North Pacific brooding subgenus of sea stars (Leptasterias subgenus Hexasterias). Calibration is obtained from a trans-arctic migration in a related clade of sea stars (Leptasterias subgenera Hexasterias and Nesasterias) that is timed to the opening of the Bering Strait at 3.5 ± 0.25 Mya, and uncertainty in the calibration point is accommodated with a normally-distributed Bayesian prior probability. Similar estimates of population splitting times for two of the pairs of putative sister taxa were obtained by a multilocus coalescent analysis. Estimates of mitochondrial mutation rates (0.01/My) were approximately 50% of the values calibrated for sister species pairs in tropical sea stars and sea urchins. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Differences in relative abundance and size structure of the sea stars Pisaster ochraceus and Evasterias troschelii among habitat types in Puget Sound, Washington, USA

We surveyed patterns in the relative abundance and size structure of the sea stars Pisaster ochraceus and Evasterias troschelii in five habitat types of varying structural complexity and prey availability (sand/cobble, boulder, and rocky intertidal; pilings; and floating docks) in Puget Sound and the San Juan Islands, Washington. For both species, small sea stars were most abundant in the most structurally complex habitat type (boulder), where they occurred almost exclusively under boulders during low tide. Larger individuals became more abundant as structural complexity decreased, occurring more frequently in open habitat types (rocky shores, pilings, and docks) known to have greater abundances of prey resources. Gull foraging observations and experiments demonstrated that exposed small sea stars of both species were highly vulnerable to predation, suggesting that small sea stars require structural complexity (crevice microhabitat) as a predation refuge. Large sea stars, once attaining a size refuge from predation, appear to migrate to more exposed habitat types with more abundant food resources. These results suggest parallel ontogenetic habitat shifts in two co-occurring consumer species related to a shared predation risk at early life stages and demonstrate how the relative importance of top-down and bottom-up processes may differ with ontogeny.     

Diet composition and feeding intensity of horse mackerel,<Emphasis Type="Italic"> Trachurus trachurus</Emphasis> (Osteichthyes: Carangidae) in the eastern Adriatic

Diet composition and feeding intensity of the horse mackerel, Trachurus trachurus, collected in the eastern Adriatic Sea, were examined. Stomach contents of 1,200 specimens, taken at monthly intervals (January–December 1996), were analyzed. The stomachs contents consisted of five major prey groups: Crustacea (Euphausiacea, Mysidacea, Decapoda), Cephalopoda and Teleostei. Euphausiacea constituted the most important food resource by weight, number and frequency occurrence. Teleosts were the second most important food category, while mysids, decapods and cephalopods were occasional food. There was little seasonal variation in diet. Euphausiids were dominant prey during all seasons, and were especially abundant from spring and autumn. Feeding intensity of horse mackerel varied during the year. The lowest intensity of feeding was recorded in winter (February, March) and early spring (April) during the lower sea temperature as well as at the time of intensive spawning. Feeding activity increased upon spawning period (May and June) and was also higher during summer. Feeding intensity and diet composition changed during the diurnal cycle. The highest feeding intensity was recorded at night and during early morning hours. Euphausiids, mysids and a greater part of teleosts dominated night and morning diet, while decapods and cephalopods were most frequently in the daily and evening diet.Communicated by O. Kinne, Oldendorf/Luhe     

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.     

Tidal-linked behaviour of the shrimp Ligur uvea in a land-enclosed marine pool

At Aldabra Atoll, the shrimp Ligur uvea inhabits a land-locked marine pool which is connected to the sea by a subterranean fissure. The shrimps migrate daily with the tide from a hole in the subterranean labyrinth to a feeding area at the bottom of the pool. The timing of the migration is tidal-linked and independent of light and temperature. With each incoming tide, tagged individuals return to a definite feeding ground. Such homing has not been reported from other L. uvea pools. Feeding activity was quantitatively measured, differences between day and night activity are interpreted as adaptations against predators (terrestrial wading birds). Compared to other L. uvea pools, the shrimps at Aldabra Atoll are exposed to a more rigorous environmental situation, which probably influences the tidal-linked behaviour of the local shrimp population.     

Time-lapse cinematography of feather stars (Echinodermata: Crinoidea) on the Great Barrier Reef,Australia: demonstrations of posture changes,locomotion, spawning and possible predation by fish

Aspects of feather star behavior and ecology were recorded by time-lapse cinematography approximately 1 frame min^-1 on the Great Barrier Reef, Australia over a 1 mo period in 1983. The current regime influenced body postures of most species studied, whether nocturnal or not. Moreover, feather stars of several species crawled on the substratum with their arms; each crawling episode lasted roughly 10 min, and the maximum speed attained was about 1 arm length min^-1. Nocturnal feather stars crawled to their nighttime feeding perches around dusk and crawled back to their daytime hiding places around dawn. Surprisingly, some species of feather stars living on the reef surface both day and night also crawled around at dawn and dusk for reasons that are not known. In the time-lapse films, and individual of Comanthus bennetti (sex undetermined) spawned for about 2 min just after dark on 5 July 1983. Another film showed possible predation on a feather star (Himerometra robustipinna) by a saddled coralfish (Chaetodon ephippium).     

Brooding and development of <Emphasis Type="Italic">Anasterias minuta</Emphasis> (Asteroidea: Forcipulata) in Patagonia,Argentina

Brooding, embryonic and larval development, and the influence of environmental and biological factors in tidepool habitats were studied in the sea star, Anasterias minuta, at various sites along ~220 km of the Patagonian coast. This species has a benthic, lecithotrophic development that includes eight distinct developmental stages. A larval organ, the connection cord, is developed from a small preoral lobe at early stages of development and becomes larger and thinner at advanced stages. Fecundity and average egg size increased with female body size. The regression of log egg number to log sea-star size and weight at different sites had a slope significantly less than 3.0, resulting in negative allometry and indicating that brood capacity was limited in large females. Development was generally synchronous among sites, but varied within each brood at advanced stages, with more developed brooded larvae located at the periphery of the brood mass. Brooding was synchronous among various populations at different years and spatial scales, and extended over a period of 8 months. The highest proportion of brooding females occurred during May and June (austral winter). Juveniles were released mainly during September. The likelihood of finding brooding sea stars decreased with increasing sea water temperature, tidal height, and wave exposure, and increased with increasing body size. Both body size of brooding females and brooding rate were higher in the infralittoral fringe than at midlittoral levels. A revision of the current model of brooding behavior and development among forcipulate sea stars is given.