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

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

Larvae of a colonial ascidian use a non-contact mode of substratum selection on a coral reef

The rate at which larvae successfully recruit into communities of marine benthic invertebrates is partially dependent upon how well larvae avoid benthic predators and settle on appropriate substrata. Therefore, to be able to predict recruitment success, information is needed on how larvae search for settlement sites, whether larvae preferentially settle on certain substrata, and the extent to which there are adequate cues for larvae to find these substrata. This article describes how larvae of the colonial ascidian Diplosoma similis find settlement sites on a coral reef. Direct field observations of larval settlement were made on a fringing reef in Kaneohe Bay, Oahu, Hawaii, between September 1985 and April 1986. A comparison of the substrata that larvae contacted prior to settlement relative to the percentage cover of these substrata on the study reef suggests that larvae are using a non-contact mode of substratum identification to locate suitable settlement sites. This mode of substratum identification allowed 74% of larvae to evade predation by benthic organisms who would otherwise have eaten larvae if they had been contacted. Of those larvae that evaded predation, 88% subsequently settled on the same two substrata upon which most adults are found (dead coral or the green alga Dictyosphaeria cavernosa). This pattern of settlement was probably a result of active selection, since the two substrata cover only 14.4% of the reef's surface and currents had little effect on the direction in which larvae swam. An important contributing factor to the high success rate of larval settlement on suitable substrata was the lack of any temporal decay in substratum preference. It is concluded that for Diplosoma similis larval supply is a sufficient predictor of larval settlement rate. However, for marine invertebrates whose larvae are passively dispersed and exhibit a greater temporal decay in substratum preference, larval settlement should generally have a greater dependency on spatial variation in the abundance of benthic predators and suitable substrata.     

Larval period and its influence on post-larval life history: comparison of lecithotrophy and facultative planktotrophy in the aeolid nudibranch Phestilla sibogae

The variable duration of the pelagic phase of metamorphically competent larvae of benthic marine invertebrates is set by an interaction between environmental factors and larval traits that together influence the chance that a larva will encounter and respond to a suitable settlement site. In the Hawaiian aeolid nudibranch Phestilla sibogae Bergh, an extended competent larval phase resulted in a cascade of negative effects on larval and post-larval life-history traits. When raised as fed (i.e., facultatively planktotrophic) larvae, an extended larval period resulted in lower larval survival, slightly lower metamorphic success, and delayed reproduction. When raised as unfed (i.e., lecithotrophic) larvae, an extended larval period resulted in lower larval and post-larval weights, survival, metamorphic success, and reproductive output, and also resulted in a longer juvenile period and delayed reproduction. The chance nature of locating a settlement site generally spreads these negative effects over all larvae of a cohort, and so balances the relative fitness of the genetic lineages within a population.     

Community recruitment: Settlement and juvenile survival of seven co-occurring species of sessile marine invertebrates

Recruitment of a group of co-occurring sessile invertebrates (a serpulid polychaete, an oyster, a bryozoan, and several colonial and one solitary ascidian) that encrust floating docks in Pearl Harbor, Hawaii, was studied in August 1985. Daily photographs were taken of acrylic settling plates over a 14 d period, and daily settlement and juvenile mortality were measured. Settlement rates were compared between species and days, while juvenile mortalities were compared between species, days, juvenile ages and densities of juveniles on the plates. Species differed in abundance of both settlers and juveniles, and in rate and pattern of juvenile mortality. Settlement intensity varied between days for some species. Significant juvenile mortality occurred during the 14 d for most species; it appeared to be caused by fish predation. For two species, mortality varied with juvenile age, with older individuals suffering higher mortality. Mortality was density-dependent for some species, increasing with numbers of juveniles on the plates. Mortality patterns varied even within the colonial species. Assessing larval settlement and juvenile mortality for a single species or type of organism in a community thus may not indicate that similar patterns exist for co-occurring 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.     

Feeding by larvae of two different developmental modes in Streblospio benedicti (Polychaeta: Spionidae)

Females of the spionid polychaete Streblospio benedicti (Webster) produce either small eggs (60–70 μm diameter) and planktotrophic larvae, or large eggs (100–200 μm) and lecithotrophic larvae that reportedly do not feed. This intraspecific polymorphism, a form of poecilogony, is potentially useful in studies of larval ecology and evolution, but necessary data on larval form and function are lacking. This study describes the morphology and nutritional biology of larvae obtained from Atlantic (South Carolina) and Pacific (California and Washington) populations from 2003 to 2005. The two types of larvae produced by Atlantic S. benedicti differed greatly in length (229±22 μm SD for planktotrophs vs. 638±40 μm for lecithotrophs) and chaetiger number (2–5 vs. 10–11) at release from the female’s brood pouch. Planktotrophic larvae bore long provisional chaetae on their first chaetiger; provisional chaetae were absent in lecithotrophic larvae. Larvae from Pacific populations were all of the lecithotrophic form, and were similar to their Atlantic counterparts in all respects. High-speed video microscopy revealed that both types of larvae used opposed bands of cilia to capture suspended particles and transport them to the mouth, where they were often ingested. Lecithotrophic larvae reared with suspended phytoplankton (Rhodomonas sp., 10⁴ cells ml⁻¹) for 2 days grew significantly faster than sibling larvae reared without added food, indicating that these larvae can digest and assimilate ingested food. Larvae of S. benedicti that develop from large eggs are thus facultative planktotrophs instead of obligately non-feeding lecithotrophs, a result that affects the interpretation of comparative studies of the ecology and evolution of larvae in S. benedicti and certain other marine invertebrates.     

Shading facilitates sessile invertebrate dominance in the rocky subtidal Gulf of Maine

Dramatic shifts in community composition occur between vertical and horizontal rocky surfaces in subtidal environments worldwide, yet the forces mediating this transition are poorly understood. Vertical rock walls are often covered by lush, diverse communities of sessile suspension-feeding invertebrates, while adjacent horizontal substrates are dominated by algae, or corals in the tropics. Multiple factors, including light, sedimentation, water flow, and predation have been proposed to explain this pattern, but experimental tests of these hypotheses are lacking. We manipulated light level and predation to test whether variation in these mechanisms could be responsible for the shift in composition of sessile communities between vertical and horizontal surfaces in the rocky subtidal Gulf of Maine. Shaded horizontally oriented granite plots were dominated by invertebrates (e.g., ascidians, barnacles, bryozoans) after 25 months. Unshaded plots were dominated by macroalgae, which was virtually absent in shaded plots. Exclusion of grazers with cages had no effect on percent cover of invertebrates or algae. Preferential settlement of invertebrate larvae to shaded plots, due to larval behaviors such as negative phototaxis, did not seem to play a role. Shading likely affects post-settlement mortality of invertebrates by alleviating competition for space with algae, although greater abundance of micropredators in algal-dominated communities could also be important. Communities on shaded plots lacked many taxa present on natural wall communities, likely due to greater disturbance on horizontal substrates and/or lack of sufficient time for colonization of these taxa. These results suggest that light plays a key role in controlling the structure, composition, and function of shallow subtidal communities.     

Low genetic differentiation between isolated populations of the colonial ascidian Symplegma rubra Monniot, C. 1972

Pelagic larvae are highly important for maintaining the gene flow among populations of sessile marine invertebrates. Colonial ascidians consist, exclusively, of brooding species, with lecithotrophic larvae that have a limited dispersal. As a result, there is a marked differentiation among populations. In this work, we used allozyme electrophoresis to access the genetic variation in four populations of Symplegma rubra, a colonial ascidian frequently found in the intertidal zone of Southeastern Brazilian coast. High variability was found at three of the four sites sampled, the exception being Praia Grande in the State of Rio de Janeiro. At this site, there was a great preponderance of clones, which possibly reflected the enclosed nature of the location and its low water circulation that reduce the dispersal capabilities of these animals. S. rubra did not conform to expectations for random mating (Hardy–Weinberg equilibrium): there was a deficit of heterozygotes that was more related to the small population size than to inbreeding processes, since F _is analysis per locus revealed a deficiency of heterozygotes at only one locus—MDH*. The greatest variation in allele frequency was found for GPI-2*. Analyses of genetic variability revealed moderate differentiation among the populations (F _ST=0.051), which was unexpected for a species with a low dispersal capability. Rafting, a frequently underestimated means of dispersal, may be the main mode of gene flow in this species over large areas, since colonies of S. rubra are frequently seen growing on drift material and there is no evidence that the larva survive for a long time in the plankton.     

Reduced planktotrophy in larvae of <Emphasis Type="Italic">Clypeaster</Emphasis><Emphasis Type="Italic">rosaceus</Emphasis> (Echinodermata,Echiniodea)

Phylogenetic analyses have demonstrated that nonfeeding larvae have evolved from feeding larvae many times among marine invertebrates. In light of this observation, it is surprising that an intermediate strategy, a larva that can feed but is provisioned with enough energy to metamorphose without acquiring exogenous food (i.e., facultative planktotrophy), is rare. A hypothesis for the lack of facultative planktotrophic species among marine invertebrates is that the transition from feeding to nonfeeding is rapid due to this intermediate stage being evolutionarily unstable. Evidence that would support this hypothesis is if species with facultative planktotrophy have reduced food assimilation when compared with obligate planktotrophs. We studied a species with facultative planktotrophic larvae, Clypeaster rosaceus, that is very near the boundary between facultative and obligatory planktotrophy, to answer two questions: (1) does feeding during the larval stage result in energy gains in larval or juvenile stages and (2) if not, are larvae capable of assimilating exogenous food at all. Our measurements of energetics in larval and juvenile stages show that C. rosaceus larvae accumulate very little if any energy when fed, but stable isotope data indicate that larvae are able to assimilate some food. Our results are consistent with similar studies on facultative planktotrophic larvae suggesting poor food assimilation and rapid loss of larval feeding after a population evolves the ability to reach metamorphosis without feeding (lecithotrophy).     

Embryonic,larval, and post-metamorphic development of the sea urchin <Emphasis Type="Italic">Cassidulus mitis</Emphasis> (Echinoidea; Cassiduloida): an endemic brooding species from Rio de Janeiro,Brazil

Cassiduloids are currently rare irregular echinoids with a highly conserved adult morphology. Aristotle’s lantern is present only during the post-metamorphic stage, and little is known about the early development of species in this group. Cassidulus mitis produces eggs of about 375 μm in diameter, lecithotrophic larvae with four reduced arms with skeletal fenestrated rods, cilia along the body surface, and a ciliated band on arms and lobes. Offspring is brooded among the female spines from embryo to settler’s stage. The echinopluteus larval stage is reached 6 days after fertilization, and the settler’s stage is formed at the age of 17 days. Aristotle’s lantern appears around the thirteenth day of development. The lantern is well developed and functional in settlers. It remains until at least 62 days after fertilization and can be used to acquire food from the environment. The early development of C. mitis is unusual concerning features of typical lecithotrophic larvae (such as reduced arms), but retains some features of planktotrophic larvae (such as skeletal rods and a ciliated band). Regarding egg size, early development in C. mitis seems to be transitioning from facultative lecithotrophic to typical obligate lecithotrophic pattern in echinoid larval evolution.     

Uncommon diversity in developmental mode and larval form in the genus <Emphasis Type="Italic">Macrophiothrix</Emphasis> (Echinodermata: Ophiuroidea)

Development mode in the ophiuroid genus Macrophiothrix includes an unusual diversity of planktonic larval forms and feeding types. The modes of development for seven congeners that coexist in coral reef habitats at Lizard Island, Australia were compared using larvae generated from crosses over several reproductive seasons from 1999 to 2003. Three species (Macrophiothrix koehleri Clark, Macrophiothrix longipeda Lamarck, Macrophiothrix lorioli Clark) develop from small eggs (<170 μm) into typical obligately feeding planktonic (planktotrophic) pluteus larvae with four larval arm pairs. The remaining four species develop from larger eggs (≥230 μm) into either facultatively-feeding or non-feeding (lecithotrophic) larval forms. The facultative planktotroph (Macrophiothrix rhabdota Clark) retains the ability to digest and benefit from food but does not require particulate food to complete metamorphosis. Among the lecithotrophic species, Macrophiothrix caenosa Hoggett retains the pluteus morphology with four pairs of larval arms, but is incapable of feeding, depending instead on maternal provisions for larval development. The remaining two lecithotrophs have simplified larval morphologies with only a single pair of full length (Macrophiothrix nereidina Lamarck) or highly reduced (Macrophiothrix belli Doderlein) larval arms and no functional mouth or gut. This genus includes the first example of facultative planktotrophy in ophiuroids, the first example in echinoderms of a complete pluteus morphology retained by a lecithotrophic larva, and three degrees of morphological simplification among lecithotrophic larval forms. Egg volume varies 20-fold among species and is related to variation in feeding mode, larval form, and development time, as predicted for the transition from planktotrophic to lecithotrophic development.     

Implications of life history for genetic structure and migration rates of southern African coastal invertebrates: planktonic,abbreviated and direct development

The amount of genetic structure in marine invertebrates is often thought to be negatively correlated with larval duration. However, larval retention may increase genetic structure in species with long-lived planktonic larvae, and rafting provides a means of dispersal for species that lack a larval dispersal phase. We compared genetic structure, demographic histories and levels of gene flow of regional lineages (in most cases defined by biogeographic region) of five southern African coastal invertebrates with three main types of larval development: (1) dispersal by long-lived planktonic larvae (mudprawn Upogebia africana and brown mussel Perna perna), (2) abbreviated larval development (crown crab Hymenosoma orbiculare) and (3) direct development (estuarine isopod Exosphaeroma hylecoetes and estuarine cumacean Iphinoe truncata). We hypothesized that H. orbiculare, having abbreviated larval development, would employ a strategy of larval retention, resulting in genetic structure comparable to that of the direct developers rather than the planktonic dispersers. However, regional population structure was significantly lower in all species with planktonic larvae, including H. orbiculare, than in the direct developers. Moreover, nested clade analysis identified demographic histories resulting from low levels of gene flow (isolation by distance and allopatric fragmentation) in the direct developers only, and migration rates were significantly higher in all three species having planktonic larvae than in the direct developers. We conclude that the amount of genetic structure within marine biogeographic regions strongly depends on the presence or absence of free-swimming larvae. Whether such larvae are primarily exported or retained, whether they have long or short larval duration, and whether or not they are capable of active dispersal seems to have little effect on connectivity among populations.     

Influence of contrasting larval developmental types upon the population-genetic structure of cheilostome bryozoans

For many sedentary or sessile benthic marine invertebrates the planktonic duration of the larval stage is believed to be a key determinant of the magnitude of genetic differences between populations. An obvious dichotomy in dispersal potential exists within cheilostome bryozoans that develop from either (1) a cyphonautes larva that spends several weeks in the plankton or (2) a brooded coronate larva that settles soon after release from the adult colony. This study characterises the pattern of variation at allozyme loci among British populations of four species of bryozoan—two species with cyphonautes and two with coronate larvae. There is some variation in the estimates of genetic differentiation among similarly separated populations that may be a consequence of non-equilibrium genetic conditions arising from sporadic migration, possibly through dispersal by rafting on macroalgae by mature colonies. Despite this, however, both the level of genetic differentiation between populations and the pattern of migrant exchange correlate with the larval developmental mode. Bryozoan species that brood coronate larvae show higher levels of genetic heterogeneity between populations and significant isolation by distance genetic structure while, by contrast, distance has little or no effect upon the amount of genetic differentiation among populations of bryozoans with cyphonautes larvae. For cheilostome bryozoans, therefore, it appears that genetic differentiation between populations is directly associated with the type of larval development. These data are discussed also with respect to levels of gene diversity and the geological pattern of cheilostome bryozoan species diversity.     

RNA-DNA ratio: an index of larval fish growth in the sea

Data on water temperature, RNA-DNA ratio, and growth of eight species of temperate marine fish larvae reared in the laboratory were fit to the equation: $$G_{pi} = 0.93{\text{ }}\operatorname{T} + 4.75{\text{ RNA - DNA}} - 18.18$$ where G_pi is the protein growth rate in % d^-1 and T is the water temperature. Water temperature and larval RNA-DNA ratio explained 92% of the variability in growth rate of laboratory-reared larvae. The model is useful over the entire range of feeding levels (starvation to excess), temperatures (2° to 20°C) and fish species studied. Estimates of recent growth of larval cod, haddock, and sand lance caught at sea based on water temperature and RNA-DNA ratio ranged from negative to 26% d^-1. These data demonstrate the importance of food availability in larval fish mortality and suggest that short-term growth under favorable conditions may be considerably higher than expected from long-term indicators. RNA-DNA ratio analysis offers new possibilities for understanding larval growth and mortality, and their relation to environmental variability.     

Waterborne chemical compounds in tropical macroalgae: positive and negative cues for larval settlement

Settlement sites of marine invertebrate larvae are frequently influenced by positive or negative cues, many of which are chemical in nature. Following from the observation that many shallow-water, Hawai'ian marine macroalgae are free of fouling by sessile invertebrates, we predicted that the algae are chemically protected and dependent on either surface-bound or continuously released soluble compounds to deter settling invertebrate larvae. To address the importance of waterborne algal compounds, we experimentally determined whether larvae of two of Hawai'i's dominant hard-surface fouling organisms, the polychaete tube worm Hydroides elegans and the bryozoan Bugula neritina, would settle in the presence of waters conditioned by 12 species of common Hawai'ian macroalgae (representing the Phaeophyta, Chlorophyta, Rhodophyta and Cyanophyta). The results included a full spectrum of biological responses by each larval species to waterborne algal compounds. Larval responses to conditioned water were consistent for each algal species, but the outcomes were not predictable based on the taxonomic relationships of the algae. For example, among the species of Phaeophyta examined, different conditioned waters were: (1) toxic, (2) inhibited settlement, (3) simulated settlement, or (4) had no effect, compared to larvae in control dishes containing filtered seawater. Additionally, larval responses to aged (24 h) conditioned waters could not be predicted from the results of assays run with conditioned waters utilized immediately after preparation. Finally, settlement by larvae of one species did not predict outcomes of tests for the other species. Four of 12 shallow-reef Hawai'ian macroalgae tested released compounds into surrounding waters that immediately killed or inhibited settlement by both H. elegans and B. neritina (toxic: Dictyota sandvicensis; inhibitory: Halimeda discoidea, Sphacelaria tribuloides, Ulva reticulata); the remaining 8 algal species prevented settlement by one of these fouling organisms but for the other had no effect or, in some cases, even stimulated settlement     

Stronger predation in the tropics shapes species richness patterns in marine communities

Species interactions are widely assumed to be stronger at lower latitudes, but surprisingly few experimental studies test this hypothesis, and none ties these processes to observed patterns of species richness across latitude. We report here the first experimental field test that predation is both stronger and has a disproportionate effect on species richness in the tropics relative to the temperate zone. We conducted predator-exclusion experiments on communities of sessile marine invertebrates in four regions, which span 32 degrees latitude, in the western Atlantic Ocean and Caribbean Sea. Over a three-month timescale, predation had no effect on species richness in the temperate zone. In the tropics, however, communities were from two to over ten times more species-rich in the absence of predators than when predators were present. While micro-and macro-predators likely compete for the limited prey resource in the tropics, micropredators alone were able to exert as much pressure on the invertebrate communities as the full predator community. This result highlights the extent to which exposure to even a subset of the predator guild can significantly impact species richness in the tropics. Patterns were consistent in analyses that included relative and total species abundances. Higher species richness in the absence of predators in the tropics was also observed when species occurrences were pooled across two larger spatial scales, site and region, demonstrating a consistent scaling relationship. These experimental results show that predation can both limit local species abundances and shape patterns of regional coexistence in the tropics. When preestablished diverse tropical communities were then exposed to predation for different durations, ranging from one to several days, species richness was always reduced. These findings confirmed that impacts of predation in the tropics are strong and consistent, even in more established communities. Our results offer empirical support for the long-held prediction that predation pressure is stronger at lower latitudes. Furthermore, we demonstrate the magnitude to which variation in predation pressure can contribute to the maintenance of tropical species diversity.     

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.     

Comparison of laboratory rates of predation of five species of marine fish larvae by three planktonic invertebrates: effects of larval size on vulnerability

Rates of predation by the invertebrates Aurelia aurita, Thysanoessa raschi and Euchaeta norvegica on larval stages of cod (Gadus morhua L.), flunder (Platichthys flesus L.), plaice (Pleuronectes platessa L.), herring (Clupea harengus L.), and turbot (Scophthalmus maximus L.) were determined. Experiments were conducted in late winter and early spring 1982 with predators collected in Loch Etive, Scotland and prey obtained from several locations in Great Britain. Early stages of the smallest species, cod, flounder and turbot, tended to be most vulnerable to all three predators, while the early stages of the larger species, plaice and herring, and older stages of all species, were less vulnerable. For all stages and species of larvae, predation rates by the three predators were most closely related to larval length and escape swimming speed. Larval length itself was closely correlated to indices of larval escape ability. Low predation rates on large larvae by E. norvegica could be due to handling difficulties, whereas for A. aurita and T. raschi these low rates were due to escape abilities of the larger larvae. Prey movement is an important stimulus eliciting predation in E. norvegica but not in A. aurita or T. raschi.     

Feeding behavior and acquisition of zooxanthellae by planula larvae of the sea anemone Anthopleura elegantissima

Symbiotic associations between cnidarians and photosynthetic dinoflagellates (i.e., zooxanthellae) are common in the marine environment. Many symbiotic cnidarians produce offspring that are initially nonsymbiotic. These new hosts must acquire symbiotic algae from environmental sources. We examined zooxanthella acquisition by laboratory-reared planula larvae of the temperate sea anemone Anthopleura elegantissima. Larvae ingested zooxanthellae while they were feeding. However, the signal that prompted larval feeding behavior did not originate from the symbiotic algae; the addition of algal cells to larval cultures never elicited a feeding response. In contrast, the addition of macerated animal tissue from several sources invariably generated a strong feeding response, which resulted in the larvae indiscriminately ingesting any particulate matter that was present, including zooxanthellae or other unicellular algae. Ingested zooxanthellae were incorporated into endodermal cells, where they remained undigested, while all other ingested material was digested or expelled within 24 h. Our results provide evidence that one source of zooxanthellae likely to serve as a route of infection in the natural environment is zooxanthella-laden mucus egested by anemones. This egested material fulfilled both of the criteria necessary for successful infection: it prompted larvae to begin feeding and provided an abundant supply of zooxanthellae that were ingested and taken up into endodermal cells of the new host.     

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.