Evolutionary ecology of settlement size in planktotrophic neritimorph gastropods

Body size during larval development is one of the most important attributes of aquatic animals. The optimal size for changing form or habitat may vary according to ecological traits of species, while phylogenetic constraints also play a significant role. The major goal of this study is to reveal the patterns in the settlement size of planktotrophic larvae in an archaic gastropod superorder Neritimorpha. We take advantage of the fact that size at various ontogenetic stages of neritimorphs can be rigorously estimated by measuring features of the adult opercula. This unique feature of neritimorphs has allowed us to generate the largest data set so far on larval settlement sizes within a group of marine invertebrates that recruit into very different post-metamorphic habitats. Eighty-eight species that represent most extant genera from rocky shores, seagrass beds, mangroves, estuaries, streams, submarine caves, deep-sea vents or seeps showed negligible intraspecific variation and considerable interspecific differences in settlement size, particularly between genera or families. Settlement size is determined primarily by phylogenetic constraints, while parallel evolution toward smaller sizes was shown to occur exclusively in four independent clades (two living and two extinct) of amphidromous snails with a marine larval period followed by a limnic adult phase. The smaller settlement size may possibly reduce the risk of being wafted away from the estuaries of their natal streams through less time achieving metamorphic competence, while ability to make occasional long-distance trips is retained by the presence of a sufficiently long delay period. This delay period also seems to obscure the possible correlation between settlement size and geographic distribution range of neritimorph species, both fully marine and amphidromous.     

Where are larvae of the vermetid gastropod <Emphasis Type="Italic">Dendropoma maximum</Emphasis> on the continuum of larval nutritional strategies?

This study evaluated whether larvae of the Indo-Pacific vermetid gastropod Dendropoma maximum are obligate planktotrophs, or whether they exhibit an intermediate feeding strategy. Experiments were conducted in Moorea, French Polynesia (149°50′W, 17°30′S), Sep–Oct 2009, to examine D. maximum larval growth and metamorphic responses to different diets and amounts of food. Dendropoma maximum larvae required particulate food to undergo metamorphosis, but were able to survive and grow in the absence of food for up to 20 days. Larvae in Low and Unfed food treatments exhibited phenotypic plasticity by growing a larger velum (the larval feeding structure) compared with those in high food. Unfed D. maximum larvae had a slower initial growth rate; however, by 11-day post-hatch fed and unfed larvae had converged on the same mean shell height (553 μm), which was only 10% larger than the initial size at hatching. Therefore, although the nutritional strategy of D. maximum larvae is best described as obligate planktotrophy, it appears to approach an intermediate feeding strategy.     

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.     

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).     

Dissolved histamine: a potential habitat marker promoting settlement and metamorphosis in sea urchin larvae

Many species of marine invertebrate larvae settle and metamorphose in response to chemicals produced by organisms associated with the adult habitat, and histamine is a cue for larvae of the sea urchin Holopneustes purpurascens. This study investigated the effect of histamine on larval metamorphosis of six sea urchin species. Histamine induced metamorphosis in larvae of three lecithotrophic species (H. purpurascens, Holopneustes inflatus and Heliocidaris erythrogramma) and in one planktotrophic species (Centrostephanus rodgersii). Direct comparisons of metamorphic rates of lecithotrophic and planktotrophic larvae in assays cannot be made due to different proportions of larvae being competent. Histamine (10 μM) induced metamorphosis in 95% of larvae of H. purpurascens and H. inflatus after 1 h, while the coralline alga Amphiroa anceps induced metamorphosis in 40–50% of these larvae. Histamine (10 μM) and A. anceps induced 40 and 80% metamorphosis, respectively, in the larvae of H. erythrogramma after 24 h. Histamine (10 μM) and the coralline alga Corallina sp. induced 30 and 70% metamorphosis, respectively, in the larvae of C. rodgersii after 24 h. No metamorphosis of any larval species occurred in seawater controls. Larvae of two planktotrophic species (Tripneustes gratilla and Heliocidaris tuberculata) did not metamorphose in response to histamine. Seagrasses, the host plants of H. inflatus, induced rapid metamorphosis in larvae of the two Holopneustes species, and several algae induced metamorphosis in C. rodgersii larvae. Histamine leaching from algae and seagrasses may act as a habitat marker and metamorphic cue for larvae of several ecologically important sea urchin species.     

Egg laying and embryonic-larval development in the snail<Emphasis Type="Italic"> Thais (Stramonita) chocolata</Emphasis> (Duclos, 1832) with observations on its evolutionary relationships within the Muricidae

Oviposition and embryonic-larval development are described for the muricacean snail Thais (Stramonita) chocolata from the Southeast Pacific coast. As with numerous other muricacean snails, this species engages in communal egg laying, with females depositing egg capsules in clusters on subtidal rocks. Each cluster of capsules contains 100–150 pedunculate, ampulliform egg capsules, with each capsule containing an average of 2,600 small (130 m) eggs. Intracapsular development was followed using light and scanning electron microscopy to describe the successive embryonic stages of the species. Free-swimming veliger larvae of about 225 m length were released from capsules after 49 days incubation at 13.6°C. The planktotrophic larvae were cultured in seawater aquaria by feeding with pure cultures of phytoplankton, recording growth and form of the larvae. Larvae reached competence after 4 months at 22°C, at 1,450–1,740 m in size, and a few larvae were observed through metamorphosis and early definitive growth. The embryonic-larval development of T. chocolata coincides with the general characteristics of the ontogeny observed in other Thais species as well as of other genera of the Rapaninae such as Concholepas. This lent support to grouping these genera into a single clade. The lack of knowledge of the development of free larvae of Thais spp. means that we do not know whether these similarities also include an extensive larval phase as generally characteristic of other members of the clade. The mode of development may be useful in characterizing some clades of this family. Thus for example, the transference of some Thais to the genus Nucella (Subfamily Ocenebrinae) is supported by differences in the mode of embryonic development, which differentiates these subfamilies. Paleobiological data reported for Neogastropoda allow postulation of primitiveness in planktotrophic larval development compared to more recent developmental strategies such as direct development of different types, which characterize various clades of this family.Communicated by O. Kinne, Oldendorf/Luhe     

Influence of delayed metamorphosis on survival and growth of juvenile barnacles Balanus amphitrite

Competent cyprid larvae of the barnacle Balanus amphitrite Darwin were prevented from metamorphosing in the laboratory for 3 or 5 d using three different techniques (holding at low temperature, crowding, and detaining on a silanized surface). We then assessed the effects of prolonging larval life on post-metamorphic growth and survival, in comparison with control individuals that metamorphosed soon after they were competent to do so. Seven experiments were conducted over 2 yr (July 1987 to September 1989). In all experiments (each with six replicates per treatment), postponing larval metamorphosis for 3 or 5 d dramatically depressed postmetamorphic growth rate (P<0.05), although metamorphic success and post-metamorphic survival were not affected (P>0.10). The results suggest that B. amphitrite cyprids deferring their metamorphosis in the field may be less successful in competing for space, at least during the first few weeks of postlarval life.     

Metamorphosis and developmental evolution in <Emphasis Type="Italic">Ophionereis</Emphasis> (Echinodermata: Ophiuroidea)

The brittle star Ophionereis schayeri has abbreviated non-feeding development through a reduced ophiopluteus and a vitellaria larva. The metamorphic changes involved with development from a bilateral larva to a radial juvenile were examined in detail. The reduced ophiopluteus has a continuous ciliated band that breaks up into discontinuous ciliary ridges at the vitellaria stage. As the vitellaria develops, the juvenile rudiment forms in the mid-ventral region. The rudiment then undergoes a morphogenetic movement to the left of the larval anterior/posterior axis. This results in a dramatic transformation from bilateral to radial symmetry and is accompanied by development of juvenile structures and settlement, 6–7 days after fertilisation. Ophiuroid development through one larval stage, the ophiopluteus, is termed Type I; whereas development through two larval stages, the ophiopluteus and vitellaria, is termed Type II. We examined the evolutionary changes in the expression of Type II development in Ophionereis by comparing the ontogeny of six species with a range of larval forms. O. fasciata has the ancestral-like planktotrophic ophiopluteus. Vestigial pluteal structures in the larvae of O. schayeri, provide a link between ophiopluteal and vitellarial forms during evolution of non-feeding development. The diversity of larval forms in Ophionereis indicates that evolution of non-feeding development through a vitellaria (Type II) may have involved an increase in egg size, reduction of pluteal structures and shortening of the time to metamorphosis. Assessment of the phylogenetic significance of Types I and II development awaits additional comparative data on the metamorphic stages of other ophiuroid genera.Communicated by G.F. Humphrey, Sydney     

Seasonal variation in the diversity and abundance of pelagic larvae of Antarctic marine invertebrates

Most marine benthic macroinvertebrate species reproduce via a larval phase but attempts to explain the occurrence of different larval strategies (feeding or non-feeding, pelagic or benthic) in different habitats have been largely inconclusive. There have been very few year-round surveys of meroplankton at any latitude and in consequence fundamental data on the diversity, abundance, and timings of larval life history phases are lacking. There has been considerable debate regarding the viability of pelagic larvae in cold waters with highly seasonal primary production but there has been only one year-round study of meroplankton in the Southern Ocean, and that was outside of the Antarctic Circle. We present data from the first year-round survey of meroplankton assemblages at a location within the Antarctic Circle. We surveyed abundances of meroplanktonic larvae over 1.5 year at Rothera Point, West Antarctic Peninsula (67°34′S, 68°07′W). Larvae were collected in monthly diver-towed net samples close to the seabed at 20 and 6 m total water depths at each of three locations and were identified and counted live immediately after sampling. A total of 99 operationally defined taxonomic types representing 11 phyla were recorded but this is likely to be an underestimate of true diversity because of inherent difficulties of identification. Larvae were present in all months of the year and although planktotrophic larvae were more abundant in summer, both feeding and non-feeding types were present in all months. Comparisons of seasonal larval abundances with data from a settlement study at the same sites and from the literature show that larvae of mobile adults settle in summer regardless of developmental type, whereas sessile taxa settle in all seasons. We suggest that this is a consequence of differences in the food requirements of mobile and sessile fauna and that the availability of food for post-larval juveniles is more critical for survival than factors affecting the larval stage itself.     

Feeding selectivity in larvae of the European hake (<Emphasis Type="Italic">Merluccius merluccius</Emphasis>) in relation to ontogeny and visual capabilities

Feeding ecology was analysed for the first time in the larvae of the European hake (Merluccius merluccius) to determine whether their diet and selectivity were constrained by environmental conditions and how these feeding characteristics were related to ontogeny, prey availability and visual capabilities. Larvae collected during both day and night were analysed, and it was found that feeding incidence was high, regardless of the time of day. Examination of the visual system corroborated the hypothesis that hake larvae should be able to cope with a wide range of photic conditions and to forage even at low light intensity. A clear preference for adult calanoid copepods and, especially, for Clausocalanus spp. was observed in all sizes analysed. Prey number increased with larval size, but prey size did not. This finding indicates that hake larvae behave as selective and specialist predators that consume an increasing number of prey rather than larger prey during larval growth.     

Availability of dissolved organic matter offsets metabolic costs of a protracted larval period for <Emphasis Type="Italic">Bugula neritina</Emphasis> (Bryozoa)

For nearly a century researchers have investigated the uptake and utilization of dissolved organic matter (DOM) by marine invertebrates, but its contribution to their growth, reproduction, and survival remains unclear. Here, the benefit of DOM uptake was assessed for the marine bryozoan Bugula neritina (Linnaeus 1758) through performance comparisons of individuals in the presence and absence of DOM. The experiments were performed using B. neritina collected from floating docks in Beaufort, NC, USA from July to September 2004. Seawater was subjected to ultraviolet irradiation to reduce naturally occurring DOM, and then enriched with either 1 μM of palmitic acid or a mixture containing 1 μM each of glucose, alanine, aspartic acid and glycine. Larvae in DOM-enriched and DOM-reduced treatments were sampled and induced to metamorphose following 1, 6, 12, and 24 h of continuous swimming at 25°C. Sampled larvae were assessed for initiation of metamorphosis, completion of metamorphosis, and ancestrular lophophore size to determine the extent to which energy acquired from DOM uptake could offset the metabolic costs of prolonged larval swimming. DOM treatment had no significant effect on initiation of metamorphosis, but did have a significant effect on completion of metamorphosis and lophophore size. Larvae swimming in DOM-enriched treatments for 24 h experienced a 20% increase in metamorphic completion rate, compared to larvae swimming for 24 h in the DOM-reduced treatment. In addition, larvae in the amino acid and sugar mixture for 24 h had a significantly larger lophophore surface area and volume (23 and 31%, respectively), compared to larvae in DOM-depleted seawater. To ensure that the increases in performance found in larvae with access to DOM were not due to a decrease in metabolic activity, the respiration rates for these larvae were compared to those of larvae in DOM-depleted seawater. There were no significant differences between these treatments, indicating that the increases in performance were due to the energy acquired from DOM. These results clearly show that for B. neritina, DOM uptake results in increased metamorphic success and in the size of the feeding apparatus following an extended larval swimming duration.     

Reproduction and development of Paedoclione doliiformis,and a comparison with Clione limacina (Opisthobranchia: Gymnosomata)

Observations on breeding season, copulation, spawning, and development are described for Paedoclione doliiformis, the only gymnosomatous pteropod which consistently retains external larval features throughout its life. The number of eggs spawned per individual is dependent upon temperature and availability of Spiratella retroversa, its thecosome prey. At 17° to 19°C, planktotrophic veligers hatch from floating egg masses within 3 days, and cast their shells approximately 11 days later. The resultant shell-less veligers could complete metamorphosis to the polytrochous larval stage, in which the body is encircled by 3 ciliary bands, within 12 h. Some polytrochous larvae begin feeding on veligers of S. retroversa within 2 days. Unlike other gymnosome species, there is no further change in external appearance except for growth. A comparison of reproduction and development has been made with Clione limacina, another gymnosome which competes for food with P. doliiformis in Nova Scotian (Canada) coastal waters.     

Prolonged exposure to low dissolved oxygen affects early development and swimming behaviour in the gastropod Nassarius festivus (Nassariidae)

Effects of low dissolved oxygen on early development and swimming behaviour of veliger larvae of the scavenging gastropod Nassarius festivus were studied. Embryonic development was significantly delayed when dissolved oxygen level was reduced to 3.0 mg O₂ l⁻¹ and no embryo hatched successfully at 0.5 mg O₂ l⁻¹. Veliger larvae hatched at 4.5 mg O₂ l⁻¹ had significantly smaller velar lobe, shell length and shell width. Median 48-h LC₅₀ value of the veliger larvae was estimated at 1.25 mg O₂ l⁻¹ with lower swimming speed (swimming velocity and dispersal velocity) being recorded for the survivors exposed to reduced oxygen levels. The percentage of veliger larvae that developed into crawling juveniles was significantly reduced and metamorphosis was delayed at 4.5 mg O₂ l⁻¹ whereas all larvae at 3.5 mg O₂ l⁻¹ died before they underwent metamorphosis. Juveniles developed at 4.5 mg O₂ l⁻¹ were also smaller than those at 6.0 mg O₂ l⁻¹. Results indicated that dissolved oxygen levels well above hypoxia levels (2.8 mg O₂ l⁻¹) have already had significant impact on the hatching success and larval development in gastropods, which may lead to long-term decreases in population growth.     

Food limitation stimulates metamorphosis of competent larvae and alters postmetamorphic growth rate in the marine prosobranch gastropodCrepidula fornicata

The effects of food limitation on growth rates and survival of marine invertebrate larvae have been studied for many years. Far less is known about how food limitation during the larval stage influences length of larval life or postmetamorphic performance. This paper documents the effects of food limitation during larval development (1) on how long the larvae ofCrepidula fornicata (L.) can delay metamorphosis in the laboratory after they have become competent to metamorphose and (2) on postmetamorphic growth rate. To assess the magnitude of nutritional stress imposed by different food concentrations, we measured growth rates (as changes in shell length and ash-free dry weight) for larvae reared in either 0.45-m filtered seawater or at phytoplankton concentrations (Isoehrysis galbana, clone T-ISO) of 1 × l0³, 1 × 10⁴, or 1.8 × 10⁵ cells ml^–1. Larvae increased both shell length and biomass at 1 × 10⁴ cells ml^–1, although significantly more slowly than at the highest food concentration. Larvae did not significantly increase (p > 0.10) mean shell length in filtered seawater or at a phytoplankton concentration of only 1 × 10³ cells ml^–1, and in fact lost weight under these conditions. To assess the influence of food limitation on the ability of competent individuals to postpone metamorphosis, larvae were first reared to metamorphic competence on a high food concentration ofI. galbana (1.8 × 10⁵ cells ml^–1). When at least 80% of subsampled larvae were competent to metamorphose, as assessed by the numbers of indlviduals metamorphosing in response to elevated K⁺ concentration in seawater, remaining larvae were transferred either to 0.45-m filtered seawater or to suspensions of reduced phytoplankton concentration (1 × 10³, 1 × 10⁴, or 5 × 10⁴ cells ml^–1), or were maintained at 1.8 × 10⁵ cells ml^–1. All larvae were monitored daily for metamorphosis. Individuals that metamorphosed in each food treatment were transferred to high ration conditions (1.8 × 10⁵ tells ml^–1) for four additional days to monitor postmetamorphic growth. Competent larvae responded to all food-limiting conditions by metamorphosing precociously, typically 1 wk or more before larvae metamorphosed when maintained at the highest food ration. Surprisingly, juveniles reared at full ration grew more slowly if they had spent 2 or 3 d under food-limiting conditions as competent larvae. The data show that a rapid decline in phytoplankton concentration during the larval development ofC. fornicata stimulates metamorphosis, foreshortening the larval dispersal period, and may also reduce the ability of postmetamorphic individuals to grow rapidly even when food concentrations increase.     

Larval alarm pheromones as a potential control for invasive cane toads (Bufo marinus) in tropical Australia

Novel approaches to control invasive species are urgently needed. Cane toads (Bufo marinus) are large, highly toxic anurans that are spreading rapidly through tropical Australia. Injured toad larvae produce an alarm pheromone that elicits rapid avoidance by conspecifics but not by frog larvae. Experiments in outdoor ponds show that repeated exposure to the pheromone reduced toad tadpole survival rates (by >50%) and body mass at metamorphosis (by 20%). The alarm pheromone did not induce tadpoles to seek shelter, but accelerated ontogenetic differentiation. Perhaps reflecting mortality of weaker individuals during larval life, growth rates post-metamorphosis were higher in animals emerging from the pheromone exposure treatment than from the control treatment. Nonetheless, body size differentials established at metamorphosis persisted through the first 8 days of post-metamorphic life. We will need substantial additional research before evaluating whether the alarm pheromone provides a way to reduce cane toad recruitment in nature, but our field trials are encouraging in this respect.     

Combined effects of temperature and salinity on the survival and growth of the larvae of Pandalus jordani (Decapoda: Pandalidae)

Survival and growth over an environmental range of temperature and salinities were examined in order to help assess the importance of these environmental factors in affecting the distribution, abundance and survival of larvae and provide greater understanding of factors affecting fluctuations in adult Pandalus jordani Rathbun population sizes. Larvae were shown to have a wide tolerance to salinity, especially in the early stages, but a relatively narrow tolerance to temperature. The optimal temperatures for survival, 8° to 11°C, were also optimal for growth as reflected by maximal growth increments and body size. It is therefore felt that fluctuations in temperature as seen within and between successive larval seasons would have profound effects on larval survival, growth rates and size at metamorphosis to the benthic juvenile phase.     

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.     

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.     

Relationship between larval and juvenile growth rates in two marine gastropods,Crepidula plana and C. fornicata

Previous studies on various marine mollusc species have shown that both larval and juvenile growth rates are substantially heritable, but few workers have examined the extent to which larval and juvenile growth rates covary. We examined the relationship between larval and juvenile growth rates in seven laboratory experiments conducted between 1986 and 1993, using the prosobranch gastropods Crepidula plana Say and C. fornicata (L.). In most experiments larvae were reared individually, measured twice nondestructively to determine larval grwoth rate, allowed or stimulated (daily 5-h exposure to 20 mM excess K⁺ in seawater) to metamophose, and then measured at least twice after metamorphosis to determine juvenile growth rates. Generally, there was no significant (p >0.10) relationship between larval and juvenile growth rates, suggesting that in these two species selection can act independently on the two stages of development. A positive correlation (p=0.007) between larval and juvenile growth rates was observed for C. fornicata in one experiment, but only for offspring from females maturing the most rapidly in laboratory culture. Even for these larvae, however, variation in larval growth rate explained<2% of the variation in juvenile growth rate, so that larval and juvenile growth rates are at most only weakly associated in this species.     

Halogenated metabolites in two marine polychaetes and their planktotrophic and lecithotrophic larvae

This study investigated the occurrence and ontogenetic changes of halogenated secondary metabolites in planktotrophic and lecithotrophic larvae and adults of two common, infaunal polychaetes, Streblospio benedicti (Spionidae) and Capitella sp. I (Capitellidae), with different life-history traits. S. benedicti contains at least 11 chlorinated and brominated hydrocarbons (alkyl halides) while Capitella sp. I contains 3 brominated aromatic compounds. These halogenated metabolites are potential defense compounds benefiting both larvae and adults. We hypothesized that: (1) planktotrophic larvae contain halogenated metabolites because they are not protected by adult defenses, (2) quantitative and qualitative variation of planktotrophic larval halogenated metabolites parallels that of adults, and (3) brooded lecithotrophic larvae initiate the production of halogenated metabolites only after metamorphosis. To address these hypotheses, volatile halogenated compounds from polychaete extracts were separated using capillary gas chromatography and identified and quantified using mass spectrometry with selected ion monitoring. All four life stages (pre- and post-release larvae, new recruits, adults) of both S. benedicti and Capitella sp. I contained the halogenated metabolites previously identified from adults. This is the first report of halocompounds identified and quantified in polychaete larvae. Allocation of potential defense compounds to offspring varied as a function of species, feeding type and developmental stage. Pre-release larvae of S. benedicti with planktotrophic development contained the lowest concentration of total halogenated metabolites (1.75 ± 0.65 ng mm⁻³), post-release and new recruits contained intermediate concentrations (8.29 ± 1.72 and 4.73 ± 2.63 ng mm⁻³, respectively), and planktotrophic adults contained significantly greater amounts (28.9 ± 9.7 ng mm⁻³). This pattern of increasing concentrations with increasing stage of development suggests synthesis of metabolites during development. Lecithotrophic S. benedicti post-release larvae contained the greatest concentrations of halometabolites (71.1 ± 10.6 ng mm⁻³) of all S. benedicti life stages and developmental types examined, while the amount was significantly lower in new recruits (34.0 ± 15.4 ng mm⁻³). This pattern is consistent with a previously proposed hypothesis suggesting a strategy of reducing potential autotoxicity during developmental transitions. Pre-release lecithotrophic larvae of Capitella sp. I contained the highest concentration of total halogenated metabolites (1150 ± 681 ng mm⁻³), whereas the adults contained significantly lower total amounts (126 ± 68 ng mm⁻³). All concentrations of these haloaromatics are above those known to deter predation in previously conducted laboratory and field trials. As a means of conferring higher larval survivorship, lecithotrophic females of both species examined may be expending more energy on chemical defenses than their planktotrophic counterparts by supplying their lecithotrophic embryos with more of these compounds, their precursors, or with energy for their synthesis. This strategy appears common among marine lecithotrophic larval forms. Received: 14 July 1999 / Accepted: 20 January 2000