Types of larval development in marine bottom invertebrates,their distribution and ecological significance: a re-evaluation

The distribution of various types of larval development among marine bottom invertebrates has been discussed on the basis of ecological evidence by Thorson (1936, 1946, 1950, 1952) and Mileikovsky (1961b, 1965). The information at hand is reviewed anew in this paper and is re-evaluated in the light of modern pertinent literature. The interrelationships between certain larval types and their distribution are not as rigid and direct as originally assumed. This can be proved even by the copy book example of the distribution of the various forms of development among species of the coastal gastropod genus Littorina. Especially among species with wide distributional areas, local populations may exhibit greater diversity in larval types than has previously been thought. Different types of larval development have now become known to exist in different populations of opisthobranch gastropods and lamellibranchs, i.e., in invertebrate groups in which such variability had been ruled out by Thorson. Variability in the type of larval development within given species — as a function of geographical, seasonal and other environmental parameters —is also more common in other marine bottom invertebrates than formerly considered. Marine bottom invertebrates are characterized not only by the 3 main different types of larval development proposed by Thorson (pelagic, direct, viviparous), but also by a fourth type: demersal (free non-pelagic) development. This fourth type occurs at all water depths and in all geographic zones of the oceans. The most important of the 4 types is pelagic (planktotrophic) development. Thorson's rule (decrease in numbers of species possessing pelagic development from the Equator towards the Poles, and from shallow-shelf waters to greater oceanic depths) is well substantiated by new data. However, one correction is necessary: pelagic development is not completely absent in the abyssal zone, as was proposed by Thorson (1950, and later), but is represented in it by at least several species belonging to various groups of invertebrates, and is also fairly common in the bathyal zone. A detailed analysis of the distributional pattern of the different types of development of marine bottom invertebrates must further take into consideration asexual reproduction with all its different modifications. Asexual reproduction in benthonic invertebrates is ecologically significant because of its common occurrence in nature; in numerous species it is also important as a biological supplement to sexual reproduction. The vast majority of species inhabiting the shallow-shelf zone and, partly, the higher levels of the slope zone of ocean areas located roughly between the polar circles, reveals development by means of planktotrophic larval stages. In the highest latitudes and on the slopes to abyssal depths—characterized by low water temperatures, scarcity of food, increasing hydrostatic pressure and other environmental peculiarities—other types of larval development prevail and, progressively, replace pelagic development with increasing latitude or depth. The distributional patterns of the various types of development among marine bottom invertebrates form one of the most important factors determining the basic distributional dynamics of the whole benthos in all oceans, both in the geological past and at the present time.Dedicated to the memory of Professor G. Thorson —founder of modern reproductive and larval ecology of marine bottom invertebrates.     

Offshore distributional patterns of Hawaiian fish larvae

An analysis of ichthyoplankton samples based on relative abundance reveals pronounced inshore/offshore distributional gradients for most Hawaiian fish larvae. Larvae of pelagic bay species are found almost exclusively in semi-enclosed bays and estuaries. Larvae of pelagic neritic species are more or less uniformly distributed with distance from shore. The larvae of reef species with non-pelagic eggs are most abundant close to shore, while those of reef species with pelagic eggs are most abundant offshore. Finally, the larvae of offshore (primarily mesopelagic) species show no clear pattern but frequently occur in high numbers nearshore. Within any group, variation in pattern was often evident; for example, although Hawaiian fishes of both the families Labridae and Mullidae spawn pelagic eggs, larvae of the former had not peaked in abundance 12 km from shore while larvae of the latter had peaked between 0.5 and 2 km. Some larvae which occur offshore are highly specialized morphologically for a pelagic existence (e.g. Chaetodontidae, which is illustrated) while others are little modified (e.g. Labridae). These findings indicate ichthyoplankton surveys in tropical areas must sample offshore areas in addition to the inshore adult habitat to obtain a complete picture.Hawaii Institute of Marine Biology Contribution No. 484.     

Cycles and rates of reproduction of six Baltic Sea bivalves of different zoogeographical origin

All bivalves investigated (Astarte borealis, Astarte elliptica, Macoma calcarea, Cyprina islandica, Macoma baltica, Cardium lamarcki) are dioecious. The mean egg-diameters were determined by a series of measurements for 6 bivalves and compared with data from other authors. M. calcarea does not have a direct non-pelagic larval development as previously assumed, but probably produces pelagic planktotrophic larvae. The arctic species exhibit no modification of their mode of development in the boreal area. The boreal forms spawn over a relatively short period in spring and summer. The arctic forms revealed, over long periods, ripe eggs and sperms. Apparently they begin to spawn in winter or early spring. This extension and shift of the spawning season represents adaptation of the Aretic species to the specific conditions of the Baltic Sea. The number of eggs laid by the most important bivalves were determined and correlated with the larval ecology and length of generation time.     

Hatching mode and latitude in marine gastropods: revisiting Thorson's paradigm in the southern hemisphere

For much of the last century the developmental mode of marine invertebrates, particularly of prosobranch gastropods, has been thought to reflect a latitudinal pattern known as Thorson's rule; the proportion of species with pelagic larvae decreases with increase in latitude. Although the predictions of this rule have been criticized recently, its validity along latitudinal transects in the world ocean still remains poorly tested. In the present work, we compare the frequency of occurrence of contrasting prosobranch hatching modes (pelagic versus benthic development) along two latitudinal gradients of the subequatorial coastline of South America: the southeast Pacific and the southwest Atlantic. The results are clearly contrasting. While the pattern observed along the Pacific (Chilean) coast fits the predictions of Thorson's rule very well, benthic development predominates all along the Atlantic coast, even at subtropical latitudes. This difference in observed patterns is attributable to the different compositions of the gastropod assemblages on each side of the South American continent, which are determined, in turn, by differences in the ecological conditions on the two sides of the continent. The scarcity of pelagic development among the Atlantic prosobranch gastropods reflects the near-continuous soft-bottom habitat there, and the consequent prevalence of predaceous, soft-bottom taxa that had acquired in their evolution more evolved, non-pelagic patterns of development. In contrast, the Pacific coast is much more heterogeneous, with a diverse mixture of benthic habitats, including rocky substrates that, in part, support grazing taxa. Our results indicate that two factors are important for Thorson's rule to be valid. First, the habitat needs to include rocky substrates, as soft-bottom habitats appear to favour non-pelagic development. Second, a diverse assemblage of taxa need to be compared to avoid the problem of phyletic constraints, which could limit the evolution of different developmental modes. Received: 23 May 2000 / Accepted: 13 October 2000     

Types of larval development in Littorinidae (Gastropoda: Prosobranchia) of the World Ocean,and ecological patterns of their distribution

Relationships between the types of larval development in 4 species of Littorina from British shores and their inhabitancy of different tidal levels were established by Woodward (1909). His data have led to the view that the Littorinidae present a “text-book” example illustrating obligate interrelationships between the distribution of marine bottom invertebrates in certain biotopes and the possession by them of definite types of larval development. Analysis of data describing larval development of 39 species of littorinids from various regions of the World Ocean covering all its climatic zones, from the tropics to the Arctic, accumulated in the literature (Tables 1-5) demonstrates, however, that there exists only a relationship between viviparity and the inhabitancy of supralittoral zones, and between direct development and the inhabitancy of littoral zones. In 34 out of 39 littorinid species studied pelagic development, in various modifications, is present at all tidal levels; it is most common in the supralittoral zone, but not in the sublittoral zone, as has been generally assumed. The data presented in Tables 1-5 confirm the author's opinion (Mileikovsky, 1971, 1973) that the concept of obligate interrelationships between definite types of larvel development in marine benthic invertebrate and their existence in definite biotopes, based on Woodward's scheme, is false and must be rejected.     

Seasonal and daily dynamics in pelagic larvae of marine shelf bottom invertebrates in nearshore waters of Kandalaksha Bay (White Sea)

Seasonal and daily population dynamics have been studied in pelagic larvae of littoral and upper-sublittoral bottom invertebrates in the plankton of the shallow, narrow Velikaya Salma Sound, which connects the inner and outer areas of the Kandalaksha Bay in the western part of the White Sea. Hydrologically, this Sound is characterised by a clearly defined cycle of great seasonal variations in water temperature coupled with more or less stable salinities and regular, pronounced semi-diurnal tides corresponding to daily and lunar monthly tidal cycles. The seasonal dynamics of larvae in the Sound reflect differences in occurrence of spawning periods in local waters of various species and systematic groups of bottom invertebrates. These differences are caused by the correlation of spawning periods of local species of different zoogeographical origin with the different water temperatures. They reflect, also, lunar periodicities of spawning and larval hatchings. The daily dynamics of larval abundancies are related to the daily spawning rhythms of many species with pelagic development affected by the daily tidal cycles of the Velikaya Salma Sound. A daily invasion of the Sound by pelagic larvae of bottom invertebrates from the inner and the outer parts of the Kandalaksha Bay occurs at ebb tide, and also at flood tide; the rhythms of the invasions coincide with the daily spawning rhythms of the Sound's invertebrates. From literature data summarized by Mileikovsky (1958a, b, 1960a, b, c, 1961, 1965, 1968, 1970), it is concluded that seasonal, lunar and daily (tidal) reproductive periodicities for the marine shallowshelf bottom invertebrates concerned, follow world-wide ecological patterns. It is evident that the effects of these rhythms upon the population dynamics of pelagic invertebrate larvae, as demonstrated by the present data on the Velikaya Salma Sound (White Sea), must also follow world-wide regularities.     

Sympatric species of threespine stickleback differ in their performance in a spatial learning task

Increasing evidence suggests that cognitive function is under selection in diverse taxa and that this results in different cognitive abilities in taxa experiencing different selective environments. For example, species inhabiting spatially complex environments might be expected to have good spatial learning ability. We investigated whether local habitat conditions influence learning by comparing the performance of two recently diverged species of threespine stickleback (Gasterosteus aculeatus complex) in a spatial learning task. The two species reside sympatrically in several lakes. Benthics occupy the spatially structured vegetated littoral zone, whereas limnetics occupy the spatially homogenous open-water pelagic zone. We trained fish to locate a hidden reward in a T-maze and asked whether they did so by learning a body-centred pattern of movement (turn left or right) or by using plant landmarks. Both benthics and limnetics used turn and landmark cues, but benthics learnt the task almost twice as quickly as limnetics. This difference is consistent with the hypothesis that benthic and limnetic sticklebacks are equipped with spatial learning abilities well suited to the spatial complexity of their littoral and pelagic habitats. Our findings add to the understanding of the evolution of learning.     

Spatial variability in habitat associations of pre- and post-settlement stages of coral reef fishes at Ishigaki Island,Japan

Successful settlement of pelagic fish larvae into benthic juvenile habitats may be enhanced by a shortened settlement period, since it limits larval exposure to predation in the new habitat. Because the spatial distribution of marine fish larvae immediately prior to settlement versus during settlement was unknown, field experiments were conducted at Ishigaki Island (Japan) using light trap sampling and underwater visual belt transect surveys to investigate the spatial distribution patterns of selected pre- and post-settlement fishes (Acanthuridae, Pomacentridae, Chaetodonidae and Lethrinidae) among four habitats (seagrass bed, coral rubble, branching coral and tabular coral). The results highlighted two patterns: patterns 1, pre- and post-settlement individuals showing a ubiquitous distribution among the four habitats (Acanthuridae) and pattern 2, pre-settlement individuals distributed in all habitats, but post-settlement individuals restricted to coral (most species of Pomacentridae and Chaetodontidae) or seagrass habitats (Lethrinidae). The first pattern minimizes the transition time between the larval pelagic stage and acquisition of a benthic reef habitat, the latter leading immediately to a juvenile lifestyle. In contrast, the second pattern is characterized by high settlement habitat selectivity by larvae and/or differential mortality immediately after settlement.     

Phylogeography, historical demography, and the role of post-settlement ecology in two Hawaiian damselfish species

Coral reef fish generally have relatively sedentary juvenile and adult phases and a presumed highly dispersive pelagic larval phase, yet previous studies that have tried to relate pelagic larval duration (PLD) to population structure have given inconsistent results. In the present study, the population structures of two damselfishes, Stegastes fasciolatus and Dascyllus albisella, were examined using mitochondrial control region sequences. The two species have similar PLDs (∼25 and 27 days respectively), but consistently differ in their settlement preferences, habitat, and densities in populations throughout the Hawaiian Archipelago, from Hawaii north to Kure Atoll, and south to Johnston Atoll. Information on habitat preferences and population densities were collected between September 2000 and October 2002, and tissue samples for the genetic studies were collected between January and April 2004. Based on the differences in habitat and abundance of the two species, the expectation was that S. fasciolatus would have high genetic variability but little population structure compared to D. albisella, and this was largely confirmed. Stegastes fasciolatus had little population structure in most of the Hawaiian Islands, and D. albisella showed evidence of strong population structure throughout its range. An exception to this pattern was the large difference between the Kure Atoll population of S. fasciolatus and all others. These results suggest that the interaction of several biological factors (e.g. species-specific spawning habitat and season) with environmental factors (e.g. seasonal wind and current patterns) may have more influence on population structure than single life history characteristics, such as the PLD.     

Phylogeography of the diamond turbot (Hypsopsetta guttulata) across the Baja California Peninsula

We compared morphology and sequenced nuclear and mitochondrial genes from 11 populations of a previously genetically unstudied “Baja California disjunct” species, the diamond turbot (Hypsopsetta guttulata). This species exhibits very limited adult movement and restriction to soft-bottom habitats but has a moderately long pelagic larval duration. Therefore, if pelagic larval duration is correlated with gene flow between Gulf of California and Pacific populations, we expect a reduced level of genetic and morphological differentiation. However, if adult habitat and ecology have more effect on gene flow, we expect the populations in the two bodies of water to be more highly differentiated. We used logistic regression to compare morphological features and phylogenetic and population genetic analyses to compare nucleotide sequence data. Gulf of California H. guttulata are different from Pacific populations in morphology and both mitochondrial and nuclear gene sequences. MtDNA shows reciprocal monophyly, and nuclear sequences from the Gulf of California formed a monophyletic group. Population genetic analyses also suggest further population subdivision within the Pacific and within the Gulf of California. We argue that adult ecology has a significant effect on migration rates among populations in the Pacific Ocean and the Gulf of California.     

Settlement patterns and post-settlement survival in two Mediterranean littoral fishes: influences of early-life traits and environmental variables

We examined the relationships between daily pattern of settlement and environmental parameters during two consecutive years in two littoral fishes, Lipophrys trigloides (Blenniidae) and Chromis chromis (Pomacentridae), in the NW Mediterranean Sea. We also used individual early-life traits (pelagic larval duration, size at hatching and size at settlement) calculated from otoliths, to study the proximate causes of settlement variability and size-selective mortality after settlement. Several early-life characteristics of L. trigloides (planktonic larval duration and size at hatching), and environmental variables averaged during the whole planktonic period (e.g. water temperature, wave height, solar radiation) were related with the magnitude of settlement. In contrast, C. chromis showed no significant relationships between early-life traits and the magnitude of settlement, and a weak relationship between settlement magnitude and environmental variables. Furthermore, juvenile survivors showed larger size at hatching than settlers, indicating that size at hatching affected the juvenile survival of the two species. These results suggest that survival was linked largely to conditions at hatching for both species.     

On predation of pelagic larvae and early juveniles of marine bottom invertebrates by adult benthic invertebrates and their passing alive through their predators

The dynamic quantitative balance between prey and predator invertebrate species inhabiting the same shallow-shelf (sublittoral level bottom) benthic communities was first discussed by Thorson (1953). Thorson considered the exact timing of larval settlement of prey and predator species possessing pelagic development and temporal supression of the adult predators' feeding activities during reproduction at the time of the preys' settlement to constitute the major factors which facilitate survival of the prey species in such communities. However, information obtained demonstrates that Thorson's “mechanism of balance between predator and prey species of benthic communities” is not always effective in securing survival from predation not only of the prey's spat but even sometimes of the predator's spat also. Because of this, the “mechanism” can not be rated as universally effective in all situations. Analysis of the data so far published demonstrates that, in marine benthic communities, especially in shallow-shelf waters, it is not uncommon for gametes, larvae, or early juveniles of different prey species to pass alive through suspension (filter)-feeding and deposit-feeding adult invertebrates preying on them. Sometimes development can even continue after excretion by predators. The hypothesis of Voskresensky (1948) and Goycher (1949) of the importance of this phenomenon for the maintenance and recruitment of the mussel Mytilus edulis and other filter-feeding lamellibranchs of nearshore waters preying on their own and other lamellibranch pelagic larvae must be rejected on the basis of accumulated data on their feeding and general biology and on the adverse influence of the mucous of their faecal pellets and pseudofaeces on the larvae excreted by them alive. The data considered here demonstrate that, although the passing alive of larvae and spat of benthic invertebrates through benthic predators is not uncommon in shallow-shelf bottom-communities, it plays no important role in the processes of maintenance and recruitment of the species and communities involved nor of the marine benthos as a whole. The actual ecological significance of predation on pelagic larvae and bottom spat of benthic invertebrate prey species by all three main trophic groups of marine benthos (suspension or filter-feeders, deposit-feeders, carnivores) and its importance to predator-prey dynamics in marine benthic communities remains open to debate until more reliable quantitative data become available.     

Impacts of ocean acidification on marine shelled molluscs

Over the next century, elevated quantities of atmospheric CO₂ are expected to penetrate into the oceans, causing a reduction in pH (?0.3/?0.4 pH unit in the surface ocean) and in the concentration of carbonate ions (so-called ocean acidification). Of growing concern are the impacts that this will have on marine and estuarine organisms and ecosystems. Marine shelled molluscs, which colonized a large latitudinal gradient and can be found from intertidal to deep-sea habitats, are economically and ecologically important species providing essential ecosystem services including habitat structure for benthic organisms, water purification and a food source for other organisms. The effects of ocean acidification on the growth and shell production by juvenile and adult shelled molluscs are variable among species and even within the same species, precluding the drawing of a general picture. This is, however, not the case for pteropods, with all species tested so far, being negatively impacted by ocean acidification. The blood of shelled molluscs may exhibit lower pH with consequences for several physiological processes (e.g. respiration, excretion, etc.) and, in some cases, increased mortality in the long term. While fertilization may remain unaffected by elevated pCO₂, embryonic and larval development will be highly sensitive with important reductions in size and decreased survival of larvae, increases in the number of abnormal larvae and an increase in the developmental time. There are big gaps in the current understanding of the biological consequences of an acidifying ocean on shelled molluscs. For instance, the natural variability of pH and the interactions of changes in the carbonate chemistry with changes in other environmental stressors such as increased temperature and changing salinity, the effects of species interactions, as well as the capacity of the organisms to acclimate and/or adapt to changing environmental conditions are poorly described.     

Vertical structure of very nearshore larval fish assemblages in a temperate rocky coast

Small-scale vertical patterns of larval distribution were studied at a very nearshore larval fish assemblage, during the spring–summer period of several years, at two depth strata (surface and bottom) using sub-surface and bottom trawls. A total of 4,589 larvae (2,016 from surface samples and 2,573 from bottom samples) belonging to 62 taxa included in 22 families were collected. Most larvae belonged to coastal species. Although inter-annual variations in larval density and diversity could be found, total larval abundance was always higher near the bottom whereas diversity was higher at the surface. A marked distinction between the structure of surface and bottom assemblages was found. Sixteen taxa explained 95% of the similarity among surface samples. Larvae which contributed most to this similarity included species like clupeiformes, sparids and serranids, and also blenniids, tripterygiids and some labrids. In the bottom samples, fewer species were present, with only six taxa, almost exclusively from species which lay demersal eggs, contributing to 95% of the similarity between samples. Larvae present at the surface were significantly smaller than at the bottom. For some of the most abundant species found at the bottom, only small larvae occurred at the surface while the whole range of sizes was present at the bottom, indicating that larvae may be completing the entire pelagic phase near the adults’ habitat. These results indicate that larval retention near the reefs probably occurs for these species, although for others dispersal seems to be the prevailing mechanism.     

The relationship between population genetic structure and pelagic larval duration in coral reef fishes on the Great Barrier Reef

Pelagic larval duration (PLD) is a commonly used proxy for dispersal potential in coral reef fishes. Here we examine the relationship between PLD, genetic structure and genetic variability in geographically widespread and ecological generalist species from one coral reef fish family (Pomacentridae) that differs in mean larval duration by more than a month. The genetic structure was estimated in eight species using a mitochondrial molecular marker (D-loop) and in a sub-set of five species using nuclear molecular markers (ISSRs). Estimates of genetic differentiation were similar among species with pelagic larvae, but differed between molecular markers. The mtDNA indicated no structure in all species except one, while the ISSR indicated some structure between the sampling locations in all species. We detected a relationship between PLD and genetic structure using both markers. These relationships, however, were caused by a single species, Acanthochromis polyacanthus, which differs from all the other species examined here in lacking a larval phase. With this species excluded, there was no relationship between PLD and genetic structure using either marker despite a range of PLDs of more than 20 days. Genetic diversities were generally high in all species and did not differ significantly among species and locations. Nucleotide diversity and total heterozygosity were negatively related to maximum PLD but again these relationships were caused by A. polyacanthus and disappeared when this species was excluded. These genetic patterns are consistent with moderate gene flow among well-connected locations and indicate that at this phylogenetic level (i.e., within family) the duration of the pelagic larval phase is unrelated to the patterns of genetic differentiation.     

Salinity favorable for early development and gamete compatibility in two sympatric estuarine species of the genus Hediste (Polychaeta: Nereididae) in the Ariake Sea, Japan

Early development was examined, under various salinities, for two sympatric nereidid polychaetes, Hediste japonica and H. diadroma, which participated in a simultaneous reproductive swarming in an estuary of the Omuta-gawa River in Ariake Sea, Japan. The eggs of both species were isotonic to the medium of 27.5–30 psu salinity. The egg diameter in the isotonic salinity was 180–205 μm in H. japonica, and 130–160 μm in H. diadroma. Successful development of most embryos was observed in a salinity range of 22.5–30 psu in both species, while successful fertilization occurred in wider ranges of salinity, i.e., 10–34 psu in H. japonica and 10 to 30 psu in H. diadroma. In both species, free-swimming larval life started from the indistinct hatching of trochophores out of a jelly layer capsule. The lecithotrophic development appeared to run to the 4-setiger nectochaetes in H. japonica, while to 3-setiger nectochaetes in H. diadroma, resulting in a shorter pelagic larval life in H. japonica. In a comparison of larval morphology among Hediste species, we found a definite negative correlation between the prostomium width, which represents the larval size and depends on egg size, and relative length of chaetae to the prostomium width: the relative length of chaetae was the longest in H. diadroma (with the smallest egg size and long pelagic life), intermediate in H. japonica (intermediate egg size, short pelagic life), and the shortest in H. atoka (largest egg size, no true pelagic life). We also examined the possibility of hybridization between H. japonica and H. diadroma through cross-insemination experiments. The gametes of the two species were reciprocally compatible, and viable hybrid offspring were produced by the laboratory crosses. The hybrid larvae expressed intermediate phenotypes, but with a greater maternal influence in characteristics such as the relative length of chaetae and the lecithotrophic larval duration.     

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.     

Open and closed seascapes: where does habitat patchiness create populations with high fractions of self-recruitment?

Which populations are replenished primarily by immigrants (open) and which by local production (closed) remains an important question for management with implications for response to exploitation, protection, and disturbance. However, we lack methods for predicting population openness. Here, we develop a model for openness and show that considering habitat isolation explains the existence of surprisingly closed populations in high-dispersal species, including many marine organisms. Relatively closed populations are expected when patch spacing is more than twice the standard deviation of a species'. dispersal kernel. In addition, natural scales of habitat patchiness on coral reefs are sufficient to create both largely open and largely closed populations. Contrary to some previous interpretations, largely closed marine populations do not require mean dispersal distances that are unusually short, even for species with relatively long pelagic larval durations. We predict that habitat patchiness has strong control over population openness for many marine and terrestrial species with a highly dispersive life stage and relatively sedentary adults. This information can be used to make initial predictions about where populations will be more or less resilient to local exploitation and disturbance.     

Spawning and larval development of the black turban snail Tegula funebralis (Prosobranchia: Trochidae)

An understanding of spawning and larval development can be fundamental to interpreting the abundance, distribution, and population structure of marine invertebrate taxa. Tegula funebralis (A. Adams, 1855), the black turban snail, has been the focus of numerous ecological studies on the Pacific coast of North America. To date, there are only conflicting and anecdotal reports of spawning, and there is no information on larval or juvenile development for this conspicuous and abundant species. On 19 September 1995, two individuals of T. funebralis were observed free-spawning gametes into seawater in tanks at the Oregon Institute of Marine Biology. Embryos and larvae were subsequently reared to metamorphosis and beyond. Development was pelagic and similar to development described for other trochids, and larvae were observed not to feed at any stage. Larvae began to metamorphose at 5.7 to 6.7 d and settled at 260 μm shell length. Juveniles grew ≃ 10 μm in shell length per day and appeared to feed on detritus. Juveniles lacked some adult diagnostic shell characters, including two columellar nodes and a closed umbilicus. In the field, small (<3 mm) juveniles occurred in the adult habitat on all sampling dates between October and March. Small juveniles were found only under rocks and were most abundant under rocks partially buried in coarse sand, suggesting that juveniles may utilize a specific microhabitat within the adult T. funebralis habitat. Received: 7 October 1996 / Accepted: 17 October 1996     

Evaluating the importance of demographic connectivity in a marine metapopulation

Recently researchers have gone to great lengths to measure marine metapopulation connectivity via tagging, genetic, and trace-elemental fingerprinting studies. These empirical estimates of larval dispersal are key to assessing the significance of metapopulation connectivity within a demographic context, but the life-history data required to do this are rarely available. To evaluate the demographic consequences of connectivity we constructed seasonal, size-structured metapopulation matrix models for two species of mytilid mussel in San Diego County, California, USA. The self-recruitment and larval exchange terms were produced from a time series of realized connectivities derived from trace-elemental fingerprinting of larval shells during spring and fall from 2003 to 2008. Both species exhibited a strong seasonal pattern of southward movement of recruits in spring and northward movement in fall. Growth and mortality terms were estimated using mark-recapture data from representative sites for each species and subpopulation, and literature estimates of juvenile mortality. Fecundity terms were estimated using county-wide settlement data from 2006-2008; these data reveal peak reproduction and recruitment in fall for Mytilus californianus, and spring for M. galloprovincialis. Elasticity and life-stage simulation analyses were employed to identify the season- and subpopulation-specific vital rates and connectivity terms to which the metapopulation growth rate (lambda) was most sensitive. For both species, metapopulation growth was most sensitive to proportional changes in adult fecundity, survival and growth of juvenile stages, and population connectivity, in order of importance, but relatively insensitive to adult growth or survival. The metapopulation concept was deemed appropriate for both Mytilus species as exchange between the subpopulations was necessary for subpopulation persistence. However, highest metapopulation growth occurred in years when a greater proportion of recruits was retained within the predominant source subpopulation. Despite differences in habitat and planktonic duration, both species exhibited similar overall metapopulation dynamics with respect to key life stages and processes. However, different peak reproductive periods in an environment of seasonal current reversals led to different regional (subpopulation) contributions to metapopulation maintenance; this result emphasizes the importance of connectivity analysis for spatial management of coastal resources.