Fuels for development: evolution of maternal provisioning in asterinid sea stars

For marine invertebrates, larval developmental mode is inseparably linked to the nutritional content of the egg. Within the asterinid family of sea stars there have been multiple, independent, evolutionary transitions to lecithotrophic development from the ancestral, planktotrophic state. To investigate the evolution of maternal investment and development within the Asterinidae, we quantified individual lipid classes and total protein for eggs and larval stages of closely related species representing three developmental modes (planktotrophy, planktonic lecithotrophy and benthic lecithotrophy). Within species, maternal provisioning differed between females indicating that egg quality varied with parentage. Maternal investment was related to egg size but, after correcting for egg volume, we identified two major oogenic modifications associated with the evolution of lecithotrophic development: (1) a reduction in protein deposition that probably reflects the reduced structural requirements of nonfeeding larvae, (2) an increase in deposition of a single class of energetic lipid, triglyceride (TG). The exception was Parvulastra exigua, which has benthic, lecithotrophic development and lays eggs with a lipid to protein ratio close to that of planktotrophs. This oogenic strategy may provide P. exigua larvae with a protein “weight-belt” that assists in maintaining a benthic existence. Asterinids with planktotrophic development used a significant portion of egg TG to build a feeding bipinnaria larva. For Meridiastra mortenseni, female-specific differences in egg TG were still evident at the bipinnaria stage indicating that egg quality has flow-on effects for larval fitness. In lecithotrophic asterinids, TG reserves were not depleted in development to the larval stage whereas protein stores may help fuel early larval development. Available data indicate that there may be two evolutionarily stable egg lipid profiles for free-spawning, temperate echinoderms.     

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.     

Giant larvae: Prolonged planktonic larval phase in the asteroidLuidia sarsi

The bipinnaria larva of the shelf-dwelling asteroidLuidia sarsi Duben and Koren, 1845, can attain (exceptionally) a length of 35 mm; metamorphosis to the adult benthic phase usually occurs well before this size is reached. A seastar rudiment develops at the oral end of the bipinnaria and, under normal circumstances, it detaches itself from its larval body before reaching a diameter of 5 mm. Midwater trawl samples taken from the surface to 1650 m in the Porcupine Seabight during 1978 and 1979, however, includedL. sarsi larvae with post-larval rudiments of 15 mm diameter and greater. It is shown that the bipinnariae and attached post-larvae both continued to grow until a point was reached when it appeared that the young sea-star could no longer be sustained by the bipinnaria. There is evidence that the largest post-larvae had absorbed the bipinnaria tissue and this, it is concluded, is the final stage before enforced settlement. From the estimated growth rate it is likely that many specimens had been in the plankton for >1 yr, giving this species a capability for long-distance dispersal.     

The effects of zooplankton swimming behavior on prey-capture kinematics of red drum larvae, <Emphasis Type="Italic">Sciaenops ocellatus</Emphasis>

Most marine fishes undergo a pelagic larval phase, the early life history stage that is often associated with a high rate of mortality due to starvation and predation. We present the first study that examines the effects of prey swimming behavior on prey-capture kinematics in marine fish larvae. Using a digital high-speed video camera, we recorded the swimming velocity of zooplankton prey (Artemia franciscana, Brachionus rotundiformis, a ciliate species, and two species of copepods) and the feeding behavior of red drum (Sciaenops ocellatus) larvae. From the video recordings we measured: (1) zooplankton swimming velocity in the absence of a red drum larva; (2) zooplankton swimming velocity in the presence of a red drum larva; and (3) the excursion and timing of key kinematic events during prey capture in red drum larvae. Two-way ANOVA revealed that: (1) swimming velocity varied among zooplankton prey; and (2) all zooplankton prey, except rotifers and ciliates, increased their swimming velocity in the presence of a red drum larva. The kinematics of prey capture differed between two developmental stages in S. ocellatus larvae. Hyoid-stage larvae (3–14 days old) fed on slow swimming B. rotundiformis (rotifers) while hyoid-opercular stage larvae (15 days and older) ate fast moving A. franciscana. Hyoid-opercular stage red drum larvae had a larger gape, hyoid depression and lower jaw angle, and a longer gape cycle duration relative to their hyoid-stage conspecifics. Interestingly, the feeding repertoire within either stage of red drum development was not affected by prey type. Knowledge of the direct relationship between fish larvae and their prey aids in our understanding of optimal foraging strategies and of the sources of mortality in marine fish larvae.     

The effect of queen-worker conflict on caste determination in the bumblebee Bombus terrestris

Endocrine analyses were used to investigate the well-known association between queen production and the onset of worker reproduction (termed the competition phase, CPh) in Bombus terrestris. Larvae that reached the age of 5 days before the CPh had a worker-like profile: low juvenile hormone (JH) biosynthesis rates and low JH hemolymph titers. In contrast, larvae that reached the age of 5 days during the CPh had a queen-like profile: high JH biosynthesis rates and high hemolymph JH levels. Larval fate could be manipulated by transplanting egg cells into host colonies with different social structures. There was a steep rise in JH production in larvae transplanted into colonies near or during the CPh. This indicates that during colony development, larvae switch from the ”worker developmental pathway” to the ”queen developmental pathway,” and that the switch is socially regulated. In small rearing groups, larvae reared with queens before the CPh developed into workers, whereas those reared with queens after the CPh developed into queens. Variation in worker type (naive or experienced) did not affect caste determination. Therefore, we hypothesize that queens produce a pheromone that directly inhibits queen differentiation by larvae. We also present two alternative scenarios that explain the timing of gyne production in B. terrestris, one based on ecological constraints and the other based on queen-worker competition. Received: 20 October 1999 / Received in revised form: 18 December 1999 / Accepted: 23 January 2000     

Temporal constraints and female preference for burrow width in the fiddler crab, <Emphasis Type="Italic">Uca mjoebergi</Emphasis>

We studied sampling behaviour and mate choice in the fiddler crab Uca mjoebergi. Once a female selects a mate, she copulates in his burrow and remains there until releasing her aquatic larvae. U. mjoebergi occurs in habitats that are inundated only by the highest amplitude spring tides. Females can only release their larvae during these tides, and release before or after will result in complete failure of reproductive effort. Matings occur over a 5-day period near the end of neap tides. Our results suggest that within the mating period, females adjust their larval developmental rates by selecting specific burrows in which to incubate their clutches. We found that at the start of the mating period, females chose larger males as mates. Since male size was positively correlated to burrow width, females were selecting wide burrows and effectively incubating at lower temperatures. This would slow down the developmental rates of larvae. In contrast, females that mated late in the mating period selectively chose small males. By incubating in narrower, warmer burrows, these females may increase the developmental rates of larvae. We propose that females are selecting burrows to influence incubation rate and ensure timely release of their larvae. Female U. mjoebergi appear to adjust their preference for the direct benefits of mate choice to increase their reproductive success.     

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

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

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.     

Permanence of Euscorpius carpathicus (L.) larvae on the mother's back (Scorpiones,Chatidae)

Summary The tendency of Euscorpius carpathicus larvae to remain on immobilized live or dried scorpions of the same or different species (Euscorpius flavicaudis) has been tested. Dried scorpions are less attractive than live ones, which are equally attractive regardless of species, sex or reproductive phase. The larvae find old (one year) dried specimens less attractive than freshly or recently (15 days) killed females, which are equally as attractive as live mothers (Table 1). A substratum treated with a chloroform extract of the mother's cuticle is consistently preferred by the larvae. The maternal behaviour (tolerating the larvae on the back) usually disappears about a week after the moult of the larvae, but it can be protracted if they are continually replaced by younger larvae (Table 2). The survival rate of larvae on live mothers is higher than on dried specimens.     

Pelagic larval development in the brooding Antarctic brachiopod Liothyrella uva

Larvae of the Antarctic brachiopod Liothyrella uva (Broderip, 1833) were released by brooding females between 10 and 15 January 1992. They were collected and cultured for the ensuing 45 d. At release they were at the gastrula stage, and were recognisable as the usual brachiopod three-lobed larva from around 18 d after release. At 22 d post-release, larvae began to congregate under shell fragments placed in culture vessels as potential settlement substrata. However, the larvae showed no signs of settlement which, combined with the absence of many of the attributes of mature larvae seen in previous studies, was taken as evidence that these larvae had not reached fully competent stages. Previous work had shown that L. uva spawns in October, indicating that the developmental period lasts for a minimum of between 115 and 160 d. The release and subsequent development of larvae also shows that what was previously thought to be a brooding species combines a long brooding period with a free-swimming phase. If this is true of other brachiopod species, then there may need to be a recategorization of some species from brooding to combined brooding and free-swimming developmental types. Even though the development period in L. uva was likely to have been underestimated because a fully competent stage was not reached, a comparison with development rates for temperate species indicated a slowing of between 8 and 70 times. This dramatic slowing of development rate in brachiopods now joins similar data previously reported for echinoderms and nemerteans, suggesting that this is a very widespred, perhaps universal, attribute of Antarctic marine invertebrates.     

Ethologie du stade infestant du copépode ascidicole Notodelphyidae Pachypygus gibber. I. Réactions à la lumière au cours de la vie larvaire pélagique

The phototropic reactions of pelagic larvae of Pachypygus gibber (Thorell) under laboratory conditions change according to their stage and their age. The behaviour of these larvae is described by 4 experimental parameters: threshold of the reaction, orientation towards the light or the opposite, speed and form of motility. Specimens were tested under different conditions of illumination. In the nauplii, the threshold reaction to light is higher than that in Copepodids 1 and 2. In the early planktonic stages (nauplii, Copepodids 1 and young Copepodids 2, from emergence to 3 days), most specimens respond positively to light, however the response is negative in older Copepodids 2 (4 to 10 days). There is a parallel evolution between the development of P. gibber and the change from positive to negative phototropism. The inversion of this response occurs in the developmental period between the 3-day old Copepodids 2 and the Copepodids 2 of 4 to 6 days of age. This reversal in vitro could correspond to the moment of penetration into the host. The average swimming speed is fastest in the positive phase of Copepodids 1, and is very slow in the older Copepodids 2. The form of the motility, rectilinear for most Copepodids 1, becomes increasingly winding as the Copepodids 2 grow. This random exploration of the environment would increase the possibility of finding the host. The evolution of phototropism is apparently related to the search for the host and its infestation.     

Development of the echinothurid sea urchin Asthenosoma ijimai

Development of an echinothurid sea urchin is described for the first time; eggs of Asthenosoma ijimai (Yoshiwara) have been fertilized in the laboratory, and development has been followed light-microscopically up to the early juvenile stage. The eggs, which are orange and float, are 1.2 mm in diameter, by far the largest echinoid eggs on record. The embryos, larvae and early juveniles are lecithotrophic, and no exogenous food is needed for development. The embryonic stages include a remarkable biscuit-shaped late blastula, which has never been described for any echinoid before. The larva, with its para-arms unsupported by skeletal ossicles, is unlike the echinoid prism or pluteus and more resembles the early bipinnaria or early auricularia of asteroids and holothurians, respectively. All stages through late larvae float just beneath the surface until settlement occurs during the third and fourth weeks at 20°C. Metamorphosis from late larva to juvenile is gradual and no part of the larval body appears to be cast off.This paper is dedicated to Professor Juro Ishida on the occasion of his seventieth birthday.     

Otolith microstructure and daily increment validation of marbled sole (<Emphasis Type="Italic">Pseudopleuronectes yokohamae</Emphasis>)

We examined the daily deposition of otolith increments of marbled sole (Pseudopleuronectes yokohamae) larvae and juveniles by rearing experiments, and estimated the growth pattern of wild larvae and juveniles in Hakodate Bay (Hokkaido Island, Japan). At 16°C, prominent checks (inner checks; ca. 19.8 µm in diameter) were observed on the centers of sagittae and lapilli extracted from 5-day-old larvae. On both otoliths, distinctive and regular increments were observed outside of the inner checks, and the slopes of regression lines between age and the number of increments (n_i) (for sagittae: n_i=0.98×Day–5.90; for lapillus: n_i=0.96×Day–5.70) did not significantly differ from 1. Inner check formations were delayed at lower temperature, and the inner checks formed 13 days after hatching at 8°C. Over 80% of larvae, just after their yolk-sac has been absorbed completely (stage C), had inner checks on both their otoliths. On the lapilli, other checks (outer check) formed at the beginning of eye migration (stage G). To validate the daily deposition of increments during the juvenile stage, wild captured P. yokohamae juveniles were immersed in alizarin complexone (ALC)-seawater solutions and reared in cages set in their natural habitat. After 6 days, the mean number of rings deposited after the ALC mark was 5.7. The age–body length relationship of wild P. yokohamae larvae and juveniles caught in Hakodate Bay was divided into three phases. In the larval period, the relationship was represented by a quadratic equation (notochord length=–0.010×Age²+0.682×Age–2.480, r²=0.82, P<0.001), and the estimated instantaneous growth was 0.38 mm day^–1 at 15 days, 0 mm day^–1 at 34 days and –0.12 mm day^–1 at 40 days. The age–body length relationship in the early juvenile stage (<50 days) and the late juvenile stage (>50 days) were represented by linear equations (standard length=0.055×Age+5.722 and standard length=0.345×Age–9.908, respectively). These results showed that the growth rates in the late larval periods and the early juvenile stage were lower than those in the early larval stage and late juvenile stage; during the slow growth period, energy appears to be directed towards metamorphosis rather than body growth. This study provided the information needed to use otolith microstructure analysis for wild marbled sole larvae and juveniles.Communicated by T. Ikeda, Hakodate     

Dissolved saxitoxin causes transient inhibition of sensorimotor function in larval Pacific herring (<Emphasis Type="Italic">Clupea harengus pallasi</Emphasis>)

Herring (Clupea harengus pallasi) spawning sites in Puget Sound, Washington overlap spatially and temporally with blooms of Alexandrium catenella, a toxic dinoflagellate species responsible for paralytic shellfish poisoning. Consequently, newly hatched herring larvae may be regularly exposed to the suite of dissolved paralytic shellfish toxins that are released into the water column from toxic cells during blooms. To date, virtually nothing is known about the impacts of these neurotoxins on early developmental stages of marine fish. In the present study, herring larvae at three ages, 0 days post hatch (dph), 4 dph, and 11 dph, were exposed to dissolved saxitoxin (STX) in 24-h and multi-day exposures. All larvae were examined for sensorimotor function (i.e. spontaneous swimming behavior and touch response). Significant reductions in spontaneous and touch-activated swimming behavior occurred within 1 h of exposure. EC₅₀s at 1 h of exposure were 1,500, 840, and 700 μg STX equiv. l⁻¹ for larvae introduced to STX at 0, 4, and 11 dph, respectively. This progressive age-specific increase in STX-induced paralysis suggests that older larvae were more sensitive to the toxin than younger larvae. Interestingly, herring larvae at all ages exhibited a significant degree of neurobehavioral recovery within 4–24 h of continuous exposure relative to the 1-h time point. This recovery of normal motor behaviors was not observed in previous studies with freshwater zebrafish (Danio rerio) larvae under the same continuous exposure conditions, suggesting that an adaptive detoxification or toxin sequestration mechanism may have evolved in some species of marine fish larvae. Our data reveal that (1) dissolved STX is bioavailable to marine finfish larvae, (2) the toxin is a paralytic agent with potencies that differ between developmental stages, and (3) STX-induced sensorimotor inhibition occurs rapidly but is transient in marine larvae. Collectively, these results suggest that dissolved algal toxins may have important sublethal effects on marine fish populations.     

Synchronous recruitment and growth pattern of planktonic larvae of the amphioxus Branchiostoma belcheri in the Seto Inland Sea, Japan

Planktonic larvae of the amphioxus Branchiostoma belcheri were collected in the western part of the Seto Inland Sea, Japan, in order to document their seasonal occurrence, recruitment, and growth patterns. The larvae appeared from mid-July to early October and their size ranged from 0.6 to 5.5 mm. Three distinct cohorts were observed during the summers, indicating multiple, synchronous spawning within the population. The length increment pattern of a cohort demonstrates slow growth for at least a week during the early larval stage. July 10, 20, and 30 are the inferred dates when spawnings started to produce the three cohorts in 2001. Favorable temperature range for spawning was 21–23°C. Bias in population structure due to advection can be assumed to be negligible; therefore the length increment of the cohort, about 0.05 mm day⁻¹, is regarded as the larval growth rate after the initial slow-growth stage. The planktonic stage of the first cohort in 2001 is estimated to have lasted around 55 days. The relative proportions of the second and third cohorts in samples from surface and mid-depth waters suggest downward migration of the planktonic larvae, as they age.     

Feeding,conversion efficiencies,and growth of larval mummichogs,Fundulus heteroclitus

Feeding rates, conversion efficiencies and growth of larvae of the mummichog Fundulus heteroclitus, an extremely abundant estuarine fish, were measured at temperatures ranging from 18° to 30°C. The food used was Artemia salina nauplii. At the time of total yolk sac absorption (5 to 7 days after hatching), the feeding rate decreased for a short time, an indication of a shift in metabolism. Higher feeding rates and growth occurred at higher rearing temperatures. The highest conversion efficiency (gross growth efficiency) was 1.1%, at 22°C. Mummichog larvae may be energetically inefficient compared with other fish species, but efficiency might not be critical for this fish, which is an opportunistic omnivore in an energy-rich environment.Contribution No. 291 of the Belle W. Baruch Institute for Marine Biology and Coastal Research, supported by DOE contract No. EY-76-5-09-0869.     

Effect of delayed first feeding on larval performance of the spider crab Maja brachydactyla assessed by digestive enzyme activities and biometric parameters

The effect of food deprivation on larval performance of the spider crab Maja brachydactyla was studied in terms of survival, moulting capacity, size, weight and enzymatic activities. Five feeding treatments that differed in the initial age of first feeding larvae (fed from hatching, 2, 4 and 6 days post-hatching and unfed) were tested for 20 days. Newly hatched larvae kept without food supply lasted for 10 days and did not moult; with 50% survival observed at 6 days post-hatching. Larvae (zoea I stage) were only able to tolerate 2 days of food deprivation after the onset of exogenous feeding without their performance being compromised. Multivariate analyses suggest that digestive enzyme activities may be good indicators of the nutritional condition of larvae.     

The influence of temperature on the development of Baltic Sea sprat (Sprattus sprattus) eggs and yolk sac larvae

In spring 2004 and 2005 we performed two sets of experiments with Baltic sprat (Sprattus sprattus balticus Schneider) eggs and larvae from the Bornholm Basin simulating ten different temperature scenarios. The goal of the present study was to analyse and parameterise temperature effects on the duration of developmental stages, on the timing of important ontogenetic transitions, growth during the yolk sac phase as well as on the survival success of eggs and early larval stages. Egg development and hatching showed exponential temperature dependence. No hatching was observed above 14.7°C and hatching success was significantly reduced below 3.4°C. Time to eye pigmentation, as a proxy for mouth gape opening, decreased with increasing temperatures from 17 days post hatch at 3.4°C to 7 days at 13°C whereas the larval yolk sac phase was shortened from 20 to 10 days at 3.8 and 10°C respectively. Maximum survival duration of non-fed larvae was 25 days at 6.8°C. Comparing the experimental results of Baltic sprat with existing information on sprat from the English Channel and North Sea differences were detected in egg development rate, thermal adaptation and in yolk sac depletion rate (YSDR). Sprat eggs from the English Channel showed significantly faster development and the potential to develop at temperatures higher than 14.7°C. North Sea sprat larvae were found to have a lower YSDR compared to larvae from the Baltic Sea. In light of the predictions for global warming, Baltic sprat stocks could experience improved conditions for egg development and survival.     

A “source” for asteroid larvae?: recruitment of Pisaster ochraceus,Pycnopodia helianthoides and Dermasterias imbricata in Nootka Sound,British Columbia

A major recruitment of the forcipulate asteroid Pisaster ochraceus was observed in September 1987 in the channel leading into Boca del Infierno, a semi-enclosed bay on the southeastern shore of Nootka Island, on the west coast of Vancouver Island, British Columbia, Canada. Newly settled recruits were observed at high densities subtidally in the channel (mean maximal density=1.23x10⁴m^-2) and at nearby sites. Subsequent surveys in 1988 to 1991 and size-frequency distributions of adults indicated strong yearly recruitment of P. ochraceus. Recruits of P. ochraceus were found on all available substrata, including hard and soft bottoms and on benthic algae. The initial food of the recruits included newly settled mussels (Mytilus sp.), snails and barnacles. Recruitment of the forcipulate Pycnopodia helianthoides was observed in 1987 to 1989 and in 1991, while recruitment of the spinulosan Dermasterias imbricata occurred only in 1988. High densities of planktonic bipinnaria larvae were found in the bay of Boca del Infierno in late May. We hypothesise that the high density population of adult P. ochraceus found in the channel of Boca del Infierno spawns synchronously and most of the resulting embryos and larvae are retained within the bay. This area may act as a source of larvae that then disperse locally. Post-recruitment mortality was estimated by comparing the density of recruits of P. ochraceus in 1987 with the numbers of juveniles presumed to be 1 yr old in 1988. Post-recruitment mortality was in excess of 97% with few individuals surviving to 1 yr. Moreover, even after the first year, mortality or emigration of the juveniles resulted in the almost complete loss of the 1-yr cohort at North Saavedra between 1988 and 1989. This study provides evidence that the P. ochraceus population in this area of Nootka Island may not be open, but may be regularly supplied from a source of larvae in the bay of Boca del Infierno. Post-settlement processes may, however, have significant effects on the local population, resulting in a poor correlation between the rate of recruitment and the incorporation of yr-1 + individuals into the adult population.