The “pelagic larvaton” and its role in the biology of the World Ocean,with special reference to pelagic larvae of marine bottom invertebrates

Ecological subdivision of marine organisms is often based on two characteristics: presence in a defined environment, and types of locomotion (degree of free active movement) in such an environment. The use of these characteristics results in a simple scheme: (1) Inhabitants of the boundary surface “ocean-atmosphere” (a zone including not only the surface film but also the thin subsurface water layer below it and the air layer just above it, i.e., pleuston and neuston). (2) Inhabitants of the deeper water layers of the ocean i.e., excluding the zone mentioned under (1): (a) passively drifting forms with very limited locomotory capacity, moving practically in the vertical plane only (plankton); (b) actively moving forms which migrate both vertically and horizontally (nekton). (3) Inhabitants of the “bottom”-benthos (level-bottom of oceans and coastal waters, tidal zones up to the upper supralittoral, different types of drifting and floating substrata, e.g. ship bottoms, harbour structures, buoys, driftwood, sargassum, whales, etc.). This simple scheme is essentially based on characteristics of adults. If developmental stages are considered, pelagic larvae of bottom invertebrates, eggs and larvae of fishes and other forms, usually present only temporarily in the plankton, neuston, and pleuston, can be distinguished as “mero-plankton”, “mero-neuston” and “mero-pleuston”, from the permanent “holo”-components of these groups. Division into “mero”-subgroups opposes all these larvae to those of planktonic, neustonic and pleustonic forms developing within the “parental” groups and their environments. However, the last category of larvae in the light of world-wide distribution of the seasonal reproductive pattern of marine invertebrates and some other organisms — especially in temperate and high latitudes — can also be rated to some degree as “mero”-(not “holo”-) components. The present paper proposes to unite all larvae of marine invertebrates (and of other organisms) undergoing pelagic development into one biological group, the “pelagic larvaton”. The main characteristic for all forms of this group is the presence of one and the same life-cycle stage in one and the same environment. All forms of the “pelagic larvaton” are, to various degrees, biologically different from their respective adult forms. Even the pelagic larvae of the holoplanktonic species exhibit some differences. Within the “pelagic larvaton”, 3 subgroups can be distinguished on the basis of their ecological peculiarities;

1. Larvae undergoing their whole development in an environment different from that inhabited by their parents and belonging to a group different from that of their parental forms; e.g. the pelagic larvae of bottom invertebrates which develop in the plankton, neuston or pleuston.
2. Larvae undergoing development in the same general pelagic environment, but in “non-parental” ecological groups; e.g. larvae of nektonic species developing in the plankton, neuston or pleuston; larvae of planktonic species in the neuston or pleuston; larvae of neustonic and pleustonic species in the plankton.
3. Larvae undergoing development in the “parental” groups; e.g. larvae of planktonic species in the plankton, of neustonic species in the neuston, or of pleustonic species in the pleuston.

In contrast to the 5 ecological groups: benthos, plankton, nekton, neuston and pleuston, the “pelagic larvaton” represents rather a biological than an ecological group. The “pelagic larvaton” comprises the 5 ecological groups and maintains the permanent turnover of organic substances between water and bottom. This group short-circuits the interrelations between the 5 ecological groups in all possible combinations. The existence of the “pelagic larvaton” presents another illustration of the unity of the biological nature of the oceans. The present paper also discusses the specific distributional patterns of the pelagic larvae of bottom invertebrates and their biological role in the seas.     

Biological investigations of the deep sea. No. 48. Phyllosoma larvae of a scyllarid lobster,Arctides guineensis,from the western Atlantic

A series of scyllarid phyllosoma larvae obtained from plankton in the western Atlantic is described and illustrated. The earliest stages are not represented; the larvae reported range in length from 6 to 59 mm. They are characterized by a narrow, elongated cephalic shield, a broad thorax, a great proportional length of the eyestalk, and by a short dactyl of the fourth pereiopod. There is no direct evidence to establish the parentage of these phyllosomas. By their geographic distribution and the elimination of other species to which larvae have been previously assigned, they are provisionally referred to the scyllarid lobster Arctides guineensis (Spengler).     

Breeding and larval distribution of the pteropod Clione limacina in the North Atlantic,Subarctic and North Pacific Oceans

The pteropod Clione limacina (Phipps, 1774) is an arcticboreal, circumpolar species, which is widely distributed in the North Atlantic and Subarctic Oceans; it also occurs in the North Pacific Ocean (in the Oyashio and neighbouring waters) and along the Atlantic coast of North America in the waters of the cold Labrador current to the Cape Hatteras region (35° N). The distribution of C. limacina larvae in the plankton of the Norwegian, Barents and White Seas, the Bear Island-Spitsbergen region of the Greenland Sea, the Newfoundland Grand Bank and the Flemish-Cap Bank region of the North-western Atlantic Ocean, and the Kurile-Kamchatka region of the North-western Pacific Ocean has been studied, and information from literature concerning the reproduction and larval occurrence of the species is summarized. Throughout its distributional are, spawning of C. limacina is characterized by the same general ecological pattern. This species breeds and spawns in all types of water masses occurring within the vertical range which it commonly inhabits — from surface layers to 500 m water depth. In all local populations of the species, the most intensive spawning is correlated with the spring/summer period of annual heating of the local waters, and the highest abundance parallels maximum growth of phytoplankton which serves as food for veligers and early polytrochous larvae. After the end of this period, spawning intensity in all local C. limacina populations declines sharply, but spawning continues at low intensity during the autumn/winter season, being practically continuous throughout the year. Distribution patterns of C. limacina larvae are determined by those of their parental forms (the parental forms spawn in the zones permanently inhabited). The earliest larval stages of C. limacina (veligers) are present predominantly in the upper 100 or 200 m water layer, i.e. in the zone of high phytoplankton abundance. Polytrochous larvae, after becoming predaceous feeders, are distributed throughout the whole water column from the surface to 500 m depth, similar to adult C. limacina. As with the adults, larvae are present (within the species' distribution area) in all types of water masses. Since the beginning of the twentieth century, in the course of the warming of the Arctic Ocean, the southern race of C. limacina (formerly a summer/autumn seasonal invader in the Norwegian Sea) has become a permanent component of the plankton fauna of the Norwegian and Barents Seas in regions influenced by the Norwegian-Northcape Current System.     

Vertical and horizontal distribution of fish larvae near coral reefs at Lizard Island,Great Barrier Reef

Consistent patterns of horizontal distribution of fish larvae from plankton tows were found in shallow waters around Lizard Island, Great Barrier Reef during 1979 and 1980. Few types of larvae were most abundant in Lizard Lagoon, and none of these were old larvae. Forty percent of the 57 types of larvae studied differed in abundance between windward and downwind sides of the island. More types of old larvae were found in greatest abundance off the windward side of the island than the downwind side. Most types of larvae preferred deeper water (>3 m) during the day and moved upward at night, although a few types preferred upper (<3 m) or middle portions of the water column. These latter were more likely to descend at night or to maintain their day-time distribution than to move upward. Windward larvae [those more abundant off the windward (SE) side of the island] were more shallow-living than were downwind larvae, and were more likely to maintain their day-time distribution at night. The current patterns around Lizard Island were favourable for retention of larvae in both Lizard Lagoon and off the windward side of the island, if combined with certain vertical distributions of the larvae. However, while there was evidence for retention on the windward side of the island, there was no evidence for retention in Lizard Lagoon. Currents on the downwind side of the island were not favourable for retention of larvae and there was little evidence that larvae were retained there. Retention may be an accidental result of interaction between currents and larval behaviour, or the result of a strategy of retention by the larvae. These could not be distinguished in the present study.     

Ecological investigations of the late-stage phyllosoma and puerulus larvae of the western rock lobster Panulirus longipes cygnus

The distribution and abundance of the late-stage phyllosoma larvae of Panulirus longipes cygnus George and the distribution and densities of the final larval stage, the puerulus, both in the plankton and at settlement along the coast, were investigated. A total of 3,617 late-stage phyllosoma (Stages VI to IX) and 301 puerulus larvae were caught at 187 plankton stations during the July to November periods 1974, 1975 and 1976 off the west coast of Australia between 29°00 to 32°30S and 113°30 to 115°00E. The depth range sampled was 0 to 35 m on the continental shelf and 0 to 90 m off the shelf. During onshore/offshore cruises with similar sampling effort on and off the shelf, 1,169 late-stage phyllosoma larvae were taken, of which only 9 were caught on the shelf, and these near the outer edge. A series of cruises sampling two areas beyond the shelf near 29°30 and 32°00S yielded 2448 late-stage phyllosoma, with greater densities of larvae in the northern location. The settlement of puerulus-stage larvae along the coast in the same geographical range was also greater in the north than in the south. The data from the onshore/offshore cruises showed a definite effect of moon phase on numbers of puerulus larvae caught on the shelf, with higher catches near new moon. The low numbers of puerulus larvae (usually 0, 1 or 2 individuals) caught at all stations showed that the puerulus stage is sparsely distributed in the plankton. Fewer puerulus larvae were present at the surface than at lower depths, but it was not possible to determine a depth preference for the puerulus between 10 m and the lowest depths sampled because of the low catch numbers. No relationships were found between puerulus larvae density and surface-water temperature, salinity, or plankton biomass at each station. Data on the larval distributions indicate that, near the end of their planktonic existence, the majority of the late-stage phyllosoma larvae of P. longipes cygnus are not carried onto the shelf, where mixing of oceanic and continental shelf waters occurs only on the outer third, but are transported southward by oceanic circulation beyond the shelf. The puerulus moults from the last phyllosoma stage beyond the shelf and completes the larval cycle by swimming across the shelf and settling in the shallow reef areas.     

Morphology and development of benthic and pelagic life stages of North Sea jellyfish (Scyphozoa, Cnidaria) with special emphasis on the identification of ephyra stages

Jellyfish blooms or invasions could be detected in an early phase of development if the youngest medusa stages (ephyrae) and their early growth stages (post-ephyrae) were identifiable in plankton samples but a useful identification key for ephyrae in early growth stages is lacking for most species. In the present study, the identification characteristics of adult North Sea scyphomedusae (Aurelia aurita, Cyanea capillata, Cyanea lamarckii, Chrysaora hysoscella, Rhizostoma octopus) collected around the island of Helgoland (German Bight) in July?CAugust 2003 and 2004 are described. Planula larvae were measured and reared to polyps in the laboratory. The process of ephyrae development asexually produced by the polyps (strobilation) was photo-documented. Photographs of the ephyrae growth stages were combined with drawings of features useful for the species identification. The provided identification key allows discrimination among post-ephyrae from plankton samples, probably leading to conclusions on the development of jellyfish blooms and their causes.     

Distribution and seasonal abundance of Polygordius spp. (Class: Polychaeta; Family: Polygordiidae) exo- and endolarvae in the southern Mid-Atlantic Bight,USA

The importance life history plays in understanding population dynamics and the functional roles of species for predicting climate change scenarios are well established. Yet, in the marine environment, the complete life history is unknown for many species, especially the link between morphologically and ecologically distinct planktonic larvae, and their corresponding benthic adult forms. Integration of meroplankton abundance, benthic adult species, larval morphology, and molecular data was employed to unravel the complete life history of Polygordius, a dominant polychaete in sandy shelf sediments of the Mid-Atlantic Bight. Polygordius species are unusual, having two distinct planktonic larval forms: an exolarva and an endolarva. Extensive sampling in the southern Mid-Atlantic Bight with careful preservation of meroplankton (2006–2007) revealed the identity of multiple stages of exo- and endolarvae, and their spatial, seasonal, and vertical distribution. Molecular and morphological evidence indicated exolarvae are Polygordius jouinae and endolarva are an undescribed species. Structure and development of these larvae differed greatly. P. jouinae exolarvae were found off Delaware Bay to North Carolina. At some stations, they were abundant, with densities up to 4,013 m^?3, comprising >90 % of the total meroplankton. Exolarvae spent up to a month in the plankton starting in March/May depending on year, settlement began in July when larvae were at least 2 mm in length, and by October were no longer observed in the plankton. These findings are consistent with the distribution patterns and life cycle known for adults. This is the first report of endolarvae north of Cape Hatteras.     

Growth and average growth rates of tanner crab zoeae collected from the plankton

Growth of the two zoeal stages of Tanner crabs, Chionoecetes bairdi and C. opilio, was estimated by changes in total dry weight of individuals sampled from plankton of the southeastern Bering Sea during the springs of 1977, 1978, 1980 and 1981. Since the age of larvae within a stage cannot be determined, approximate beginning and end weights of each stage were used. Epidermal retraction, conspicuous under low magnification of a dissecting microscope, was used to identify larvae presumed to be in advanced stages of growth (biomass) for their respective larval stage. This assumption was corroborated by holding retracted larvae that subsequently molted on board ship, by comparison of dry weight measurements from extensive field sampling, and by the low proportion of larvae showing such conspicuous retraction. An exponential growth model was used to calculate the average daily growth rate based on estimates of stage duration in the field. The adequacy of this model for describing the average rate, rather than the pattern, of growth is discussed. Carbon-specific rates of respiration from discrete measurements and of growth averaged for each zoeal stage indicate average net growth efficiencies comparable to laboratory results for other species. The method used to estimate growth should be applicable to other larval decapods and offers the advantage of estimating this parameter from data gathered from natural populations.Contribution No. 652 of the School of Fisheries, University of Washington     

Two methods of sampling fish larvae over reefs: a comparison from the Gulf of California

I compared the sampling properties of two methods for collecting fish larvae over reefs: nighttime collecting with a light trap, and daytime collecting with a small plankton net that could be steered by a diver. Samples were collected in the Gulf of California during summer, 1989 and 1990. The 90 light-trap samples yielded 9406 larvae from 31 families, while the 75 plankton-net samples yielded 17852 larvae from 43 families plus unidentified anguilliforms. Four families were collected only in the light trap, and 16 families plus the anguilliforms were collected only with the plankton net. With one exception, the families that were collected by only one method were rare. Twenty-seven families were collected by both methods, but only 13 were collected at least five times by each. The average catch per sample differed significantly between methods for 9 of these 13 families. In each case, the plankton net yielded more larvae per sample. The distribution of larvae among families was less equitable in light-trap samples than in planktonnet collections, primarily because clupeids were so dominant in the former. However, the taxonomic composition of light-trap and plankton-net collections was broadly similar. Seven families were shared among the ten most abundant families for each method, and the relative abundances of taxa (47 families plus anguilliforms) were strongly correlated between methods. A comparison of larval size-distributions for 12 families indicated that the size structure of catches usually differed between collecting methods. In four families there was little overlap in the size classes collected, in five families the distributions overlapped broadly but had different shapes, and in three families the size distributions were similar. Although the light trap collected larger larvae on average, its catches were not limited to settlement-stage or transition larvae. Larvae of at least ten families were present over reefs in all size classes, but the combination of both sampling methods was usually required to detect this. Based on their abundance and wide size distribution over reefs, at least some larvae from these ten families may remain over reefs throughout development. However, additional data are required to determine the importance of water over reefs as a larval habitat.     

Diel and tidal cycles regulate larval dynamics in salt marshes and mangrove forests

Variation in the release and recruitment of larvae of estuarine invertebrates affects the distribution and abundance of adults, as well as trophic interactions in both the plankton and the benthos. Larval release and supply are often timed to environmental cycles such as the diel and tidal cycles. Here, we determined using plankton tows whether the abundance of larvae spanning salt marsh and mangrove habitats across the intertidal landscape varied with diel and tidal cycles. Using three different sampling designs across two sites and within each of two estuaries over a 12-month period, we covered a range of spatial and temporal scales. This allowed us to test the general prediction that densities of meroplankton in the water column would be greater during nocturnal ebb tides than during other phases of the diel or tidal cycle. As predicted, nocturnal ebb tides yielded the highest densities of meroplanktonic larvae and were dominated by first-stage crab zoeae and this finding was most pronounced in the salt marsh. Throughout the course of the year, greater numbers of meroplankters consistently occurred during the ebb tide compared with the flood tide. The densities of other taxa (e.g. gastropods and polychaetes) showed no clear trends with diel or tidal cycles. This study highlights the effects of these pervasive physical cycles on the timing of larval release and supply in the salt marsh–mangrove complex, and emphasises their contribution to the trophic interactions and the dynamics of benthic populations within estuaries.     

How does cod (Gadus morhua) cope with variability in feeding conditions during early larval stages?

Response of mesocosm-reared cod (Gadus morhua L.) larvae to different feeding conditions was investigated in 1988 in two mesocosms: a large basin and a smaller bag enclosure within the basin. The basin was filled with seawater, and a community of naturally occurring plankton developed. Plankton concentrations were monitored, and cod larvae stocked in the enclosures were sampled for determination of growth, survival, and gut content. In the bag, insufficient amounts of energetically favourable prey, as copepod nauplii, led to non-selective ingestion of plankton from a broad range of sizes, including considerable amounts of protozoans (tintinnid and oligotrich ciliates). Growth of larvae from the bag was low, with daily specific growth rates (SGR) less than 2.8% the first 3 wk post-hatch. This was followd by rapid increase of SGR to 21.7%, which coincided with a large increase in availability of copepod nauplii. In the basin, high nauplii concentrations led to SGR of 13.7 to 21.7% from onset of feeding to 16 d post-hatch, respectively. Under such conditions, the larvae were highly selective feeders. At 3 wk post-hatch, survival was 36.7 and 38.3% in the basin and bag enclosure, respectively. To cope with variations in the feeding conditions, the cod larvae were shown to be opportunists when nauplii were scarce, and included plankton from several trophic levels in their diet. When nauplii were abundant, cod larvae realized their high potential for growth. Both opportunism and realization of a high growth potential may enhance survival of the larvae.     

Growth maximization in early sardine larvae: a metabolic approach

Sardine larvae are forced to grow as fast as possible to reduce larval mortality. Thus, changes in biochemical composition during the first steps of sardine larval growth are intended to maximize larval growth rate efficiency and survival. Protein and RNA weight-specific growth rates were the highest and their corresponding doubling times the shortest among all the biomolecules, reflecting the importance of fast growth during early stages of larval development. The protein percentage increased and the carbohydrate and lipid percentages decreased during early growth until they reached, respectively, a percentage of 73.7, 3.1 and 18.0%. These percentages would represent the optimal proportion of biochemical components in sardine early larvae and they are the result of the trade-off between, in the short term, the protein proportion necessary to optimize larval movement and growth and, in the long term, the minimum lipid percentage necessary to guarantee energy reserves to fuel metamorphosis. RNA/DNA ratio increases during larval growth up to an asymptotic optimal value of ≈3.5 in postflexion larvae. Nutritional condition of sardine larvae was good and was influenced by the parental effect through the egg biochemical composition and by the growth trajectory determined by the actual environmental conditions.     

Real-time PCR assay for detection and relative quantification of <Emphasis Type="Italic">Liocarcinus </Emphasis><Emphasis Type="Italic">depurator</Emphasis> larvae from plankton samples

Accurate species identification of decapod crustacean larvae is required to understand their population distributions, life cycle dynamics and interactions with their habitats. Analysis of plankton samples using morphological taxonomic methods and microscopy is time-consuming, requires highly skilled and trained operatives and may often be inaccurate. As complementary tools to classical identification methods, recent work has focused on the development of molecular approaches and shows their feasibility for species-specific identification. This study has developed real-time PCR assays utilising species-specific Taqman^® probes designed in the cytochrome oxidase I (COI) gene of Liocarcinus depurator, Necora puber, Carcinus maenas and Cancer pagurus. Our study then employed the probe and primers designed for L. depurator to obtain accurate identification and relative abundance estimates of L. depurator larvae in plankton samples collected between March 2005 and October 2006. Ranges of larval abundances were derived from a standard curve created from plankton samples spiked with a known number of larvae reared in the laboratory. Inhibition of the PCR reaction was shown to be an important factor and our results suggested that 0.1 ng of DNA as template provided accurate identification and avoided inhibition. Real-time PCR was shown to provide accurate species identification on unsorted plankton samples and could be suitable for the estimation of larval abundances in the plankton, although more work must be done to improve the accuracy of those estimations.     

Vertical distribution of eggs and larvae ofEngraulis encrasicolus in stratified waters of the western Mediterranean

The vertical distribution of the eggs and larvae of the European anchovy (Engraulis encrasicolus) in the western Mediterranean Sea in June 1984 and Agust 1985 was analyzed based on multiple plankton tows carried out at varying depths and using nets equipped with opening and closing mechanisms. Hydrographic parameters such as temperature, salinity, and chlorophylla were recorded simultaneously. Maximum abundance of anchovy eggs and larvae always occurred above the thermocline, even when maximum chlorophylla concentrations were located below the thermocline. Larval distribution appeared to be associated with the availability of suitable food organisms. As in other clupeoid species, the anchovy larvae carried out vertical migrations related to the photoperiod.     

Specificity and sensitivity of microsatellite markers for the identification of larvae

The identification of larval marine invertebrates to species or even higher taxonomic levels by morphological examination is notoriously difficult. Many diagnostic features are absent or poorly formed at early stages in development. This is particularly true for the larvae of bivalve molluscs, for which a routine and accurate method of identification would prove valuable to both ecologists and fishery managers. A simple molecular genetic method to identify specifically larvae of the European oyster, Ostrea edulis L., 1758, is presented. The test is based on PCR amplification of highly species-specific microsatellite loci and is sensitive enough to register the presence of a single larval individual (~200 µm width) in a mixed sample of 20 mg wet weight plankton (approximately 250 larval animals). This work demonstrates that microsatellite loci can be used as highly sensitive and specific taxonomic indicators, for studies of planktonic larvae. Details of three novel microsatellite loci are also given for O. edulis, increasing the suite of molecular tools available for use in population genetic studies of this commercially important species.     

Fine-scale distribution of larval fishes: patterns and processes adjacent to coral reefs in Kaneohe Bay,Hawaii

Plankton samples were taken from January to June 1987 in Kaneohe Bay, Oahu, Hawaiian Islands, with a free-fall plankton net, to investigate the fine-scale distribution of larval fishes around coral reefs. Daytime samples indicated that the postflexion larvae of two gobiids (Psilogobius mainlandi and an unidentified species) were significantly more abundant at stations immediately adjacent to reefs (near-reef) than at stations in open water off the reef (off-reef). These postflexion gobiid larvae appeared to be capable of resisting advection and dispersal while remaining in the water column near suitable adult habitats. The larvae of Foa brachygramma (Apogonidae) and Encrasicholina purpurea (Engraulidae) were significantly more abundant at off-reef stations than at near-reef stations. Nighttime samples indicated that the gobiid larvae depend on visual cues to remain near the reef. The horizontal distributions of F. brachygramma and E. purpurea larvae appeared to be related to their vertical positioning. These data suggest that typical ichthyoplankton surveys which do not sample close to adult fish habitats would greatly underestimate the abundances of larvae such as the gobiids.     

Planktonic larval duration of one hundred species of Pacific and Atlantic damselfishes (Pomacentridae)

The plankton larval duration for 100 species of Pacific and Atlantic damselfishes was estimated from daily growth increments on the otolith of juvenile fish collected at various localities between July 1987 and September 1988. For newly-settled fishes, larval duration was determined by counting the entire number of increments present on the otolith, while for older juveniles estimates were made by counting the number of increments between the center of the otolith and a mark corresponding to settlement. We document the development of otolith formation during the period when eggs are incubated on the reef and show that daily increments are only accreted after larvae hatch and enter the planktonic phase. The planktonic larval duration for damselfish is shorter and less variable, both between and within species, compared to other groups of reef fishes such as wrasses and surgeonfishes. Larval duration ranged from 12 to 39 d. Average duration between species ranged from 13.1 to 35.2 d. The time spent in the plankton was not significantly correlated with geographic distribution when evaluated among species, however, genera with confined regional distribution have a shorter mean larval life than do widely distributed genera. Size at settlement was positively correlated with time spent in the plankton among species, but a significant correlation between these variables was only evident within one of ten species. The low variance in planktonic larval duration within species indicates that most damselfish are unable to delay metamorphosis following competency. This inability to postpone settlement limits the potential for dispersal, especially when dispersal time between suitable habitats is greater than about 30 d.     

Larval abundance and recruitment of the sand dollar Dendraster excentricus in Monterey Bay,California, USA

Seasonal abundance of planktonic larvae of the sand dollar Dendraster excentricus was determined in Monterey Bay, California, USA. Larvae were counted from two offshore stations and also over a coastal sand dollar bed, and these data were compared with settlement in the sand dollar bed, with adult population structure and with adult reproductive condition. These measurements were made during the period July 1978 to October 1980 and in October 1981. Sand dollar larvae were most abundant in the plankton during the summer, a period when phytoplankton productivity tended to be high and currents were relatively slow and variable. In some years, small-scale current variations appeared to prevent many larvae in the open bay from reaching the nearshore sand dollar beds; however, in other years, massive shoreward transport of the larvae evidently did occur, since the adult population in the sand dollar bed exhibited a mode in size, indicating a large settlement. A comparison of settled individuals in 1980 and the adult size-frequency distribution in 1981 gives an estimated mortality rate of 88% yr^-1 for early juveniles of D. excentricus.     

Dispersal of phytoplanktotrophic shipworm larvae (Bivalvia: Teredinidae) over long distances by ocean currents

Shipworms or Teredinidae may be dispersed either as adults in floating wooden objects or as pelagic larvae drifting near the sea surface. Five shipworm species, i.e., half of those having an amphi-Atlantic geographical distribution, are known also to have pelagic phytoplanktotrophic larvae which can be carried by ocean currents. From a series of 742 plankton samples taken from throughout the temperate and tropical North Atlantic Ocean, it can be shown that shipworm larvae are not uncommon in the open sea. Teredinid veligers were found in 19% of all samples taken. One species of larvae, identical in all details to that described by Rancurel (1965), is particularly prevalent and is tentatively identified as Teredora malleolus (Turton). A definitive identification will be possible only after the pelagic larvae of the other Atlantic species are known. The larvae of Teredora malleolus are found throughout the North Atlantic Gyre and the adjacent temperate and tropical seas, and from scattered records in the South Equatorial Current. Larvae of other unidentified Teredinidae species were also found. The distance that larvae may be transported depends upon the length of pelagic larval development and the velocity of the currents. From the known current velocities it can be shown that, even in a few weeks, larvae may be dispersed many hundreds of kilometers. The geographical distribution of shipworm larvae suggests that they are carried along the coasts of continents and even across ocean basins, and that this dispersal must be an important factor in the geographical distribution of the adult forms and in the maintenance of genetic continuity between populations otherwise isolated from one another.Contribution No. 2555 from the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, USA