Size and structure of a photographically identified population of manta rays <Emphasis Type="Italic">Manta alfredi</Emphasis> in southern Mozambique

The size and structure of a photographically identified population of reef manta ray, Manta alfredi, were examined at aggregation sites over a four-year period in southern Mozambique. The use and standardisation of photo-ID techniques was examined as a minimally-intrusive means to study this species. Using these techniques, we report on the size, structure and seasonality of this population of M. alfredi. In total, 449 individuals were identified during this time period, 40.5% of which were re-sighted on at least one occasion. The longest period between re-sighting events was 1,252 days. During the study period, annual population size estimates for M. alfredi ranged from 149 to 454 individuals. The superpopulation size estimate for the entire study period was 802 individuals, the first reported for M. alfredi at a monitored aggregation site. A highly significant sex bias was evident with a female:male ratio of 3.55:1. The majority of rays (89.9% males; 49.7% females) were considered mature, with most individuals between 3.0 and 4.9 m in disc width. Manta alfredi were observed at the study sites in each month of the calendar year. The maximum number of individual rays seen per dive was 30. Large numbers of rays (20 + per dive) were seen in the months of November, December and January, which coincide with the breeding season. Natural markings were unique to individuals and did not change substantially with time, which provided further support for their use in the identification of individual M. alfredi over multiple years. Multiple re-sightings of individual M. alfredi suggest that many individuals in this population exhibit site fidelity to the examined aggregation sites. As target subsistence fishing for M. alfredi exists along the Mozambican coastline, management efforts to monitor and prevent overexploitation at these critical habitats should be a priority.     

Plasticity in the temporal organization of behaviour in the limpet <Emphasis Type="Italic">Cellana grata</Emphasis>

The behaviour of intertidal consumers is often tightly constrained to tidal movements, although activity patterns can vary within these constraints. Spatio-temporal variability in behaviour of a limpet, Cellana grata, was analysed over different tidal conditions (spring and neap tides) and during different times of the year (one summer and one winter) at sites in Hong Kong. Activity was generally dictated by tidal movements, being concentrated when animals were awash. Plasticity in behaviour was observed, with some limpets anticipating activity during the summer period and delaying activity during winter time. Limpets were active for a time equal, or slightly less, than time awash. As the time awash exceeded ~14–16 h, however, activity duration decreased. Within this general pattern, tidal variation as well as variation among times of the year was noted, with the lowest dependence on time awash being recorded during winter neap tides. Limpets showed a slight preference for being active during nighttime, which was particularly evident when animals were emersed during the summer period. Although the basic activity in C. grata is constrained to a specific temporal window, this limpet is able to modulate its foraging strategies and resting height, according to local, daily changes in environmental conditions.     

Pricing Policy for Tourism in Protected Areas: Lessons from Komodo National Park, Indonesia

Abstract: Protected areas are under increasing pressure to provide economic justification for their existence, particularly in developing countries where demand for land and natural resources is high. Nature-based tourism offers a mechanism to generate substantial benefits from protected areas for both governments and local communities, and ecotourism is increasingly promoted as a sustainable use of protected areas. The extent to which ecotourism offsets the costs of a protected area has rarely been examined. We used financial data from Komodo National Park, Indonesia, and a willingness-to-pay questionnaire of independent visitors to (1) examine the financial contribution of tourism in offsetting the costs of tourism and wider management and (2) assess the effect of hypothetical fee increases on park revenues, visitation patterns, and local economies. Although only 6.9% of park management costs were recovered, visitors were willing to pay over 10 times the current entrance fee, indicating a substantial potential for increased revenue. The potential negative effect of large fee increases on visitor numbers and the resultant effect on local economic benefits from tourism may limit the extent to which greater financial benefits from Komodo National Park (KNP) can be realized. Our results suggest that a moderate, tiered increase in entrance fees is most appropriate, and that partial revenue retention by KNP would help demonstrate the conservation value of tourism to both visitors and managers and has the potential to increase visitors' willingness to pay.     

Larval ecology of the great barracuda, <Emphasis Type="Italic">Sphyraena barracuda</Emphasis>, and other sphyraenids in the Straits of Florida

The great barracuda (Sphyraena barracuda) is a widespread, ecologically and socioeconomically important coastal fish, yet very little is known about its larvae. We examined spawning and larval ecology of Western Atlantic sphyraenids using monthly ichthyoplankton samples collected over 2 years along a transect spanning the east–west axis of the Straits of Florida (SOF). Samples were dominated by the great barracuda (92.8%) and sennets (Sphyraena borealis and Sphyraena picudilla; 6.6%). While larval sennets and S. barracuda displayed similar vertical distributions (majority in upper 25 m), horizontal and temporal patterns of abundance suggested a spatial and temporal species replacement between larval S. barracuda and sennets that tracks adult ecology. The diet of both taxa consisted largely of copepods, with inclusion of fish larvae at 8 mm SL, and in S. barracuda alone, a switch in the wet season to exclusive piscivory by 12 mm SL (18 days post-hatch). A lack of piscivory in S. barracuda larvae captured in the dry season corresponded to slower larval growth than in the wet season. Larval growth was also related to size-at-hatch and larval age such that larvae that were larger at hatch or larger (older) at capture grew faster at earlier ages, suggesting faster larval growth, and indirectly larger hatch size, conveys a survival advantage. Unlike larval growth, instantaneous mortality rate did not differ with season, and no lunar cyclic patterns in spawning output were identified. Our results provide insight into the pelagic phase of sphyraenids and highlight the importance of both diet and hatch size to the growth and survival of fish larvae in low latitude oceanic environments.     

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.     

Ontogenetic habitat shift,population growth,and burrowing behavior of the Indo-Pacific beach star, <Emphasis Type="Italic">Archaster typicus</Emphasis> (Echinodermata; Asteroidea)

Archaster typicus, a common sea star in Indo-Pacific regions, has been a target for the ornamental trade, even though little is known about its population biology. Spatial and temporal patterns of abundance and size structure of A. typicus were studied in the Davao Gulf, the Philippines (125°42.7′E, 7°0.6′N), from February 2008 to December 2009. Specimens of A. typicus were associated with intertidal mangrove prop roots, seagrass meadows, sandy beaches, and shoals. Among prop roots, specimens were significantly smaller and had highest densities (131 ind. m⁻²) between November and March. High organic matter in sediment and a relatively low predation rate seemed to support juvenile life among mangroves. Size and density analyses provided evidence that individuals gradually move to seagrass, sandy habitats, and shoals as they age. Specimens were significantly larger at a shoal (maximum radius R = 81 mm). New recruits were found between August and November in both 2008 and 2009. Timing of recruitment and population size frequencies confirmed a seasonal reproductive cycle. Juveniles had relatively high growth rates (2–7 mm month⁻¹) and may reach an R of 20–25 mm after 1 year. Growth rates of larger specimens (R > 30 mm) were generally <2 mm month⁻¹. The activity pattern of A. typicus was related to the tidal phase and not to time of day: Specimens moved over the sediment surface during low tides and were burrowed during high tides possibly avoiding predation. This is one of the first studies to document an ontogenetic habitat shift for sea stars and provides new biological information as a basis for management of harvested A. typicus populations.     

Synergistic effects of ultraviolet radiation and conditions at low tide on egg masses of limpets (<Emphasis Type="Italic">Benhamina obliquata</Emphasis> and <Emphasis Type="Italic">Siphonaria australis</Emphasis>) in New Zealand

The effects of ultraviolet radiation (UVR), desiccation and conditions in tidal pools on embryonic survival were examined for two common pulmonate limpets that lay intertidal benthic egg masses on rocky shores in New Zealand: Benhamina obliquata and Siphonaria australis. Field surveys and manipulative experiments were conducted between December 2006 and September 2007 in the Wellington region of New Zealand (41°17′S, 174°47′E). Egg mass deposition sites in the field were species-specific: B. obliquata deposited eggs primarily in shaded crevices, whereas S. australis predominantly deposited egg masses in the sun and in tidal pools. For both species, however, embryonic mortality was greater in egg masses that had been in full sun compared to shade. For S. australis, there was also high mortality in egg masses in tidal pools or desiccated compared to those that remained submerged in flowing seawater at low tide. In outdoor experiments, embryonic mortality was also always greatest for egg masses exposed to full sun, and lowest for those in shaded treatments. Mortality was also higher if egg masses were in simulated tidal pools, and for S. australis, if desiccated, compared to those submerged in flowing seawater. Periods of particularly sunny conditions with high temperatures also resulted in higher overall mortality. Finally, egg masses of both species that were initially deposited in the shade had greater mortality in response to subsequent UV exposure compared to egg masses initially deposited in full sun. Results from this study suggest that the egg masses of these two species are highly vulnerable to UVR, as well as other intertidal stressors. Embryos of both of these species may be at risk of high mortality particularly during summer when extreme conditions of UV intensity and high temperature coincide with low tide cycles. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Laboratory culture of the aeolid nudibranch <Emphasis Type="Italic">Spurilla neapolitana</Emphasis> (Mollusca,Opisthobranchia): life history aspects

Spurilla neapolitana from the Eastern Mediterranean was cultured in a self-sustained, temperature-controlled laboratory culture system, and its life cycle is described. Adults were collected from three field sites situated 120 km apart, along the Israeli Mediterranean coastline, between March 2006 and August 2007. Cultures of the life-cycle stages were raised at 24°C. S. neapolitana deposited white, coiled, spiral egg masses containing zygotes. Veliger larvae hatched 3.0 ± 0.4 days post oviposition. The veliger larvae are obligatory planktotrophs, with a minimal larval phase of 22 days. In the lab, larvae settled and metamorphosed following exposure to metabolites derived from distinct prey sea anemone species. Reproductive maturity was reached 42 ± 5 days post metamorphosis, resulting in a laboratory generation time of 67 days (egg to egg). The average life span of reproductive specimens in this study was 157 ± 13 days post-oviposition and they reached a length of 7–10 cm. During this period, an average adult deposited ca. 40 × 10⁶ zygotes. This species has several characteristics that suggest it will be a useful model for laboratory-oriented research. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Current and historic distribution and abundance of the inarticulated brachiopod, <Emphasis Type="Italic">Lingula reevii</Emphasis> Davidson (1880), in Kaneohe Bay,Oahu, Hawaii,USA

The inarticulated brachiopod, Lingula reevii Davidson (1880) is a filter-feeding invertebrate that burrows vertically in sandy or mixed sediments. Its only recorded occurrence is from Kaneohe Bay, Oahu, Hawaii, southern Japan, and Ambon, Indonesia. Past surveys of Kaneohe Bay populations suggested a distinct decrease in abundance following the diversion of sewage effluent from the bay in 1978/1979. In the summer of 2004 and 2007, visual surveys were conducted in areas of historical L. reevii abundance as well as in areas appearing to have suitable habitat. In 2004, approximately 2,950 m² at 20 sites within the bay were surveyed using quantitative belt transecting methods. A maximum density of 4 Lingula/m² was observed, a decrease from previous maximum estimates of 500 individuals/m² (Worcester, Dissertation, Zoology Department, University of Hawai′i, pp 49, 1969) and 100 individuals/m² (Emig, J Exp Mar Biol Ecol 52:47–61, 1981). When these 20 sites were revisited in 2007, many had fewer or no L. reevii; therefore, broader scale presence/absence surveys were conducted at 16 additional sites in the bay (also surveyed in 2004). The highest density of L. reevii found in 2007 was 0.94 individuals/m². The continued decline in abundance of L. reevii in Kaneohe Bay may be due, in addition to decreased organic enrichment from diversion of sewage discharge almost 30 years ago, to the more recent reduction of suitable habitat by the invasion of mat-forming alien algae species.     

Scales of variation in otolith elemental chemistry of juvenile staghorn sculpin (<Emphasis Type="Italic">Leptocottus armatus</Emphasis>) in three Pacific Northwest estuaries

Although distinct otolith elemental signatures are often observed in fish collected from different estuaries, significant differences are also observed among sites within estuaries. Variation at these smaller spatial scales is not well quantified and has the potential to lead to inappropriate interpretations of otolith elemental data. To quantify variation at multiple scales, the otolith elemental composition (Mg:Ca, Mn:Ca, Sr:Ca, Ba:Ca, and Pb:Ca) of juvenile staghorn sculpin (Leptocottus armatus Girard, 1854) collected from five sites within three estuaries, the Columbia River (two sites) and Coos Bay (one site), Oregon, and Humboldt Bay, California (two sites), was examined. Using laser ablation-inductively coupled plasma mass spectrometry, each otolith was sampled at three zones: (1) within the primordium, which represents the egg and early larval periods; (2) at the outer edge, which represents the juvenile period just prior to collection; and (3) midway between the primordial and edge samples, which represents the late larval and early juvenile period. There were significant differences in otolith metal-to-calcium ratios at all scales examined. Using multi-element otolith signatures, fish were classified to estuary and site within estuary with relatively high levels of accuracy (av = 70–90%). The largest differences in metal-to-calcium ratios were observed between sites within estuaries (<5 km apart) and the smallest differences were observed among otolith zones. Variation in otolith chemistry may be used to provide information on probable habitat use by estuarine fish but studies must be carefully designed. Electronic supplementary material  Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Habitat influences on reproductive allocation and growth of the mummichog (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>) in a coastal salt marsh

To determine whether life history differences can occur in salt marsh fishes that occupy different habitats within the same marsh, we compared reproductive allocation in female mummichog (Fundulus heteroclitus) inhabiting creeks and ponds of a coastal salt marsh in southern New Jersey, USA during the spring and summer of 2001 and 2002. Females were collected in phase with the lunar spawning cycle from four sites of each habitat type, and assessed for gonad-to-body-mass ratio and growth increment. Annual reproductive allocation, expressed as a percent of somatic mass, was estimated for each site and year from the gonadosomatic indices of individuals collected during each spawning period. Mummichogs from creeks showed little change in annual reproductive allocation from 2001 to 2002, whereas those inhabiting ponds showed a significant increase between these years. Seasonal reproductive patterns indicated that pond females cease spawning at least one lunar cycle earlier than creek fish. While ponds experienced considerably higher maximum summer temperatures than creeks as well as near-anoxic pre-dawn conditions, neither of these variables explained a significant amount of variation in annual reproductive allocation. In contrast, annual reproductive allocation of mummichogs in a pond correlated with its flood frequency in both years of study. Our results suggest that while the length of the spawning season differs in mummichogs inhabiting marsh creeks and ponds, annual reproductive allocation depends more upon the hydrodynamic conditions of the particular waterbody than its habitat type per se.     

Relation of spawning of the giant scallop,Placopecten magellanicus,to temperature fluctuations during downwelling events

We examined the relation of spawning to biological and physical factors for the scallop Placopecten magellanicus (Gmelin, 1791) over 8 yr (1984 to 1991) in the Baie des Chaleurs, southwestern Gulf of St. Lawrence, Canada. Spawning was always abrupt and occurred between July and mid-September. It did not appear to be related to the abundance of phytoplankton or particulate organic carbon and nitrogen in the water. It further showed no relationship to lunar or tidal phascs or to current velocity. Spawning consistently occurred during the summer temperature maximum, but did not coincide with any critical temperature or cumulative temperature threshold. All but one of the 33 spawning events, for which temperature data were recorded, were associated with temperature changes; 25 of these were sharp temperature increases and 7 were during strong temperature fluctuations when the mean temperature was 9 to 14°C. Both types of temperature changes were caused by downwelling of warm surface water. The delay by about 1 d in time of spawning between sites coincided with the rate at which downwelling events propagated into the bay. Virtually all of the spawning events resulted in gametes being ejected into warm water masses where conditions are likely to favour larval development.     

Long-term individual identification and site fidelity of white sharks, <Emphasis Type="Italic">Carcharodon carcharias</Emphasis>, off California using dorsal fins

Mark-recapture techniques can be used to estimate white shark (Carcharodon carcharias) population abundance. These frameworks are based on assumptions that marks are conserved and animals are present at the sampling location over the entire duration of the study. Though these assumptions have been validated across short-time scales for white sharks, long-term studies of population trends are dependent on these assumptions being valid across longer periods. We use 22 years of photographic data from aggregation sites in central California to support the use of dorsal fin morphology as long-term individual identifiers. We identified five individuals over 16–22 years, which support the use of dorsal fins as long-time individual identifiers, illustrate strong yearly site fidelity to coastal aggregation sites across extended time periods (decades), and provide the first empirical validation of white shark longevity >22 years. These findings support the use of fin morphology in mark-recapture frameworks for white sharks.     

Spawning patterns in the leopard grouper, <Emphasis Type="Italic">Mycteroperca rosacea</Emphasis>, in comparison with other aggregating groupers

We documented the spawning patterns of the leopard grouper, Mycteroperca rosacea, from April to June 2005 in the central Gulf of California, Mexico to draw comparisons with other aggregate-spawning groupers and provide information useful for management of their fishery. Adults formed spawning aggregations of 150 to >700 individuals at specific sites, and spawning occurred daily at these sites from late April through early June. Courtship occurred throughout the day, but spawning was restricted to the evening hours. Adults spawned in groups of 6–40 fish, and pair-spawning was not observed. The group-spawning behavior of adults and the gonosomatic indices of mature males (maximum = 7.2%) suggest that sperm competition was present. The site-specificity of leopard grouper spawning aggregations and diel spawning period were typical of most aggregating groupers, and the size and structure of these aggregations was similar to other species in the genus Mycteroperca. Leopard grouper behavior patterns were unusual in that spawning aggregations persisted for extended periods, spawning was not synchronized with the lunar cycle, and adults aggregated during non-spawning periods. The extensive duration and site-specificity of spawning aggregations and the propensity of M. rosacea to form aggregations year-round increases the vulnerability of the species to overfishing. Policies that limit harvest from these aggregations are needed to improve the management of leopard grouper fisheries in the Gulf of California.     

Relationship of daily and circatidal activity rhythms of the fiddler crab,<Emphasis Type="Italic"> Uca princeps</Emphasis>, to the harmonic structure of semidiurnal and mixed tides

Intertidal organisms may employ circatidal rhythms to track the tidal cycle, but tidal patterns may vary within a species range and necessitate adaptation to the local tides. Circatidal rhythms were examined in populations of the eastern Pacific fiddler crab Uca princeps (Smith) from four sites with differing tidal characteristics, La Paz (24°10N; 110°21W), San Blas (21°33N; 105°18W) and Manzanillo (19°6N; 104°24W), Mexico (lower amplitude, mixed semidiurnal tides) and Mata de Limon, Costa Rica (9°55N; 84°43W) (high-amplitude, semidiurnal tides). Local tides were characterized by harmonic constants of M₂, S₂, K₁, and O₁, partial tides that largely determine their semidiurnal and diurnal features. Rhythmic structure in continuously recorded locomotor activity of individual crabs held under laboratory conditions was described by cosinor and periodogram methods of time-series analysis. Both daily and circatidal rhythms were found in crabs studied in light–dark cycles set to local conditions at the time of collection. Crabs at all four sites shared a tendency toward bimodality, with a mid-morning activity peak and varying degrees of nocturnal activity. Circatidal rhythms closely matching the period of the 12.42-h M₂ partial tide were consistently present at all sites except Manzanillo. At Mata de Limon, the circatidal rhythm clearly dominated locomotor activity, but was strongly modulated by a daily rhythm in a repeating pattern at a semilunar interval. In contrast, the amplitude of the daily rhythm was higher than that of the circatidal rhythm in crabs from the three mixed tide sites on the Mexican coast, where the tidal pattern is dominated by a diurnal inequality arising from the diurnal K₁ and O₁ partial tides. These results suggest that populations of U. princeps use both daily and circatidal timing systems to track local forms of the tide generated by their M₂, S₂, K₁, and O₁ geophysical counterparts.Communicated by J.P. Grassle, New Brunswick     

Temporal and spatial use of stranded wrack by the macrofauna of a tropical sandy beach

During two field studies the macrofauna associated with wrack stranded on a tropical sandy shore was analysed. During the first period all animals present in small wrack deposits were randomly collected with cores. During the second field study active animals were sampled in larger wrack deposits for an entire lunar period, and tube traps were used. After a comparison between the two studies, only the data from the tube traps were examined from a temporal and spatial point of view. The aim of the present work was to gain information on how, when and where wrack was colonised by invertebrates, keeping in mind cyclical aspects such as lunar, diel and tidal phases. Predatory taxa, such as Staphylinoidea and the amphipods Talorchestia martensii, were most abundant, whereas Diptera larvae and Tenebrionidae were scarce. Successional changes of beach wrack colonisation were evident throughout a semi-lunar period, with molluscs invading wrack during the first days of deposition and histerids during the last ones. Strictly nocturnal or diurnal surface-active species were found to be active in the wrack during both the day and the night. The analysis of the tidal component of species activity in the wrack showed that some species moved at ebbing tide, and others moved at rising tide. Differences were found also in the mean hours of tidal activity, calculated separately during day and night periods. The study of the zonation of each species showed that in some cases wrack deposits were closely followed by the fauna as their position changed during the semi-lunar phase. In other cases differences occurred between species zonation during day and night periods and tidal phases. Cluster analysis indicates that most species exploit the wrack in different ways (as refuges and/or feeding site), both in space and in time. Received: 11 July 1999 / Accepted: 20 January 2000     

Recruitment and ontogenetic habitat shifts of the yellow snapper (<Emphasis Type="Italic">Lutjanus argentiventris</Emphasis>) in the Gulf of California

We examined recruitment and ontogenetic habitat shifts of the yellow snapper Lutjanus argentiventris in the Gulf of California, by conducting surveys and collections in multiple mangrove sites and major marine coastal habitats from 1998 to 2007. Over 1,167 juvenile individuals were collected and 516 otoliths were aged to describe the temporal pattern of the settlement. L. argentiventris recruits in mangroves, where juveniles remain until they are approximately 100 mm in length or 300-days-old. Back-calculated settlement dates and underwater surveys indicated a major recruitment peak during September and October, around 8 days before and after the full moon. The majority of mangrove sites in the Gulf of California had a similar L. argentiventris average size at the beginning of the settlement season for the cohort of 2003; although there were significant differences in individual sizes at the end of the nursery stage. When sub-adults leave mangroves, they live in shallow rocky reefs and later become abundant in deeper rocky reefs. The density of migratory individuals (10–20 cm SL) decreased exponentially as the distance between a reef and a nearby mangrove site increased. This finding has important implications for local fishery regulations and coastal management plans.     

Mantle-to-shell CaCO<Subscript>3</Subscript> transfer during shell repair at different hydrostatic pressures in the deep-sea vent mussel <Emphasis Type="Italic">Bathymodiolus azoricus</Emphasis> (Bivalvia: Mytilidae)

Calcium carbonate transfer was experimentally examined in hydrothermal mussels Bathymodiolus azoricus, which were collected from 850 m depth at Menez Gwen hydrothermal vent site (31°31′W, 37°50′N) on the Mid Atlantic Ridge in May 2007. In each of four 10-day experiments, groups of mussels were maintained at atmospheric pressure or re-pressurised to depths relevant to their site of occurrence, i.e. 850 m depth at Menez Gwen, 1,750 m at Lucky Strike (31°31′W, 37°18′N) and 2,300 m at Rainbow (31°31′W, 36°13′N). The shells of experimental mussels were perforated and mantle tissue was fixed for light and TEM studies at days 7 and 10 following the injury. Simultaneously, haemocytes from the extrapallial fluid (EPF) at the site of induction were studied. At day 7 the response was most intense in the middle fold of the mantle margin and possibly proportional to hydrostatic pressure. At day 10 the epithelial cells on the mantle surface facing the body cavity produced copious organic secretions that avidly bound calcium. Haemocyte migration was noted within the mantle tissue, and the haemocytes at the mantle surface facing the shell had a Ca-positive granular content. Large haemocytes were detected in the EPF at the injury site, and some showed evidence of an immune reaction while others showed Ca-positive granular content. These results suggest that haemocytes are involved in shell repair in these deep-sea mussels just as in some freshwater and shallow marine molluscs.