Lecithotrophic Larval Stage Research Articles

Morphological variation in first-formed shells of the Ordovician Paucicrura–Diceromyonia brachiopod lineage of North America

Abstract First-formed shells of several species of Dalmanellidae (Brachiopoda) from the Ordovician (Katian) of North America were measured and compared: Cincinnetina multisecta, Diceromyonia tersa, Diceromyonia storeya, Paucicrura corpulenta, Paucicrura rogata, and Paucicrura sillimani. Sizes and structures of the first-formed shells suggest that members of this family had planktotrophic subadults, with some species showing indications of only an unshelled larval stage and others showing both a larval stage and a shelled juvenile stage. This differs from modern rhynchonelliformean brachiopods, which all possess only a lecithotrophic larval stage. The range of sizes of first-formed shells of most studied species are similar, but P. sillimani of Baffin Island, Canada (middle Katian), has a significantly larger first-formed shell that formed during an extended juvenile stage. This may have enabled the species to colonize newly exploitable habitats during an interval of rapid sea level rise in Laurentia during the Katian. This plasticity of developmental modes in the Dalmanellidae shows not only that using distantly related modern brachiopods as an analog for extinct Paleozoic lineages may be misleading, but also that development can vary within a single lineage and that timing of developmental stages should not be considered a reliable character for use in phylogenetic studies of brachiopods.

The small polychaete Platynereis dumerilii revealed as a large species complex with fourteen MOTUs in European marine habitats

Recent studies reporting complexes of cryptic or pseudo-cryptic species with narrow geographic distributions have been challenging the cosmopolitan status of a fair number of marine benthic invertebrates. Morphologically similar species are often overlooked but molecular techniques have been extremely effective in signalling potential hidden diversity which, complemented with further detailed examination, might reveal unique morphological and ecological features. Evidence of morphological stasis, where no clear and stable morphological differences are apparent, can be exemplified by the annelids Platynereis dumerilii (Audouin & Milne Edwards, 1833) and Platynereis massiliensis (Moquin-Tandon, 1869). These sibling species, usually found among algae in marine intertidal and subtidal habitats, can only be distinguished by their different reproductive strategies and life histories. The former is gonochoric, with a single reproductive event in life (semelparous) transforming into a pelagic epitokous form called heteronereis, has free spawning synchronized by lunar periodicity and a larval stage with planktotrophic development; while the latter shows no epitokous transformation and is a protandrous hermaphrodite, characterized by egg brooding and lecithotrophic larval stages with a semi-direct development. In order to verify the possible existence of additional hidden Platynereis species within the P. dumerilii morphotype, we used a multi-locus approach to investigate 26 populations along Europe from the NE Atlantic and the Macaronesia islands (Azores, Madeira and Canaries) to the Western and Eastern Mediterranean Sea. We concatenated the mtDNA COI-5P, rDNA 16S and 28S-D2 sequences and performed a phylogenetic analysis through Bayesian inference (BI). To depict Molecular Operational Taxonomic Units (MOTUs), we applied three delimitation methods (ABGD, bPTP and GYMC) to the concatenated alignment, except for COI where we also applied the Barcode Index Number (BIN), implemented in BOLD, which is exclusive to this locus. Consensus MOTUs were defined based on the majority rule and, in case of draw, the most conservative MOTUs were chosen. We detected at least 14 MOTUs with 23.1% COI mean K2P distance (6.6 - 32.6%). The BI tree is split into three major clades (Clade A: MOTUs 1-3, Clade B: MOTUs 4-9 and Clade C: MOTUs 10-14), with MOTU 3 appearing to represent P. dumerilii sensu stricto and MOTU 9 P. massiliensis. This assumption is based on data from the Type locality and a previous study by Wage et al. (2017), which combined phylogeographic (COI barcode region), reproductive biology and life-history observations on some selected Platynereis populations thriving in the vent areas from the Italian islands of Ischia and Vulcano. Major Cade C seems to be pseudo-cryptic as some visible differences can be found in the specimens, as for example, the size of the tentacular cirri and lack of dorsal pigmentation, in contrast with Clades A and B, where only differences in pigmentation types were found so far. The Spanish archipelago of the Canary Islands and the whole Mediterranean Sea seem to be a cryptic hotspot. Five MOTUs are unique to the Macaronesia and five sympatric MOTUs are present in the Gran Canaria and La Palma islands alone. Additionally, three lineages were present exclusively in the Mediterranean with four sympatric MOTUs spotted in the southeast of Spain (Calpe) and the Greek island of Crete. Three out of four NE Atlantic MOTUs are shared with the Mediterranean with one exclusive to this part of the continent. Failure to recognise this hidden biodiversity may compromise the accuracy and the interpretation of biomonitoring data or other relevant ecological studies. Integrative taxonomy is thus essential to solve these uncertainties and to allow naming the involved undescribed species. Otherwise, most molecular data providing enough support for species hypothesis will continue to be unused, and large fractions of biodiversity will persist unnoticed.

Phylogeny, expression patterns and regulation of DNA Methyltransferases in early development of the flatfish, Solea senegalensis

BackgroundThe identification of DNA methyltransferases (Dnmt) expression patterns during development and their regulation is important to understand the epigenetic mechanisms that modulate larval plasticity in marine fish. In this study, dnmt1 and dnmt3 paralogs were identified in the flatfish Solea senegalensis and expression patterns in early developmental stages and juveniles were determined. Additionally, the regulation of Dnmt transcription by a specific inhibitor (5-aza-2′-deoxycytidine) and temperature was evaluated.ResultsFive paralog genes of dnmt3, namely dnmt3aa, dnmt3ab, dnmt3ba, dnmt3bb.1 and dnmt3bb.2 and one gene for dnmt1 were identified. Phylogenetic analysis revealed that the dnmt gene family was highly conserved in teleosts and three fish-specific genes, dnmt3aa, dnmt3ba and dnmt3bb.2 have evolved. The spatio-temporal expression patterns of four dnmts (dnmt1, dnmt3aa, dnmt3ab and dnmt3bb.1) were different in early larval stages although all of them reduced expression with the age and were detected in neural organs and dnmt3aa appeared specific to somites. In juveniles, the four dnmt genes were expressed in brain and hematopoietic tissues such as kidney, spleen and gills. Treatment of sole embryos with 5-aza-2′-deoxycytidine down-regulated dntm1 and up-regulated dntm3aa. Moreover, in lecithotrophic larval stages, dnmt3aa and dnmt3ab were temperature sensitive and their expression was higher in larvae incubated at 16 °C relative to 20 °C.ConclusionFive dnmt3 and one dnmt1 paralog were identified in sole and their distinct developmental and tissue-specific expression patterns indicate that they may have different roles during development. The inhibitor 5-aza-2′-deoxycytidine modified the transcript abundance of dntm1 and dntm3aa in embryos, which suggests that a regulatory feedback mechanism exists for these genes. The impact of thermal regime on expression levels of dnmt3aa and dnmt3ab in lecithotrophic larval stages suggests that these paralogs might be involved in thermal programing.

Abbreviated larval development of Macrobrachium inpa Kensley and Walker, 1982 (Crustacea: Decapoda: Palaemonidae) from an Amazon Basin forest stream, Brazil, reared in the laboratory

This paper brings the description and illustrations of the abbreviated larval development of the Amazonian freshwater palaemonid shrimp, Macrobrachium inpa Kensley and Walker, 1982. The study was based on ovigerous females (mean total body length of 27.0 ± 1.64 mm) collected in a small forest stream in the Reserva Florestal Ducke, near Manaus, Brazil, of which four released their larvae in the laboratory. The females carried 8 to 19 eliptical (2.39 ± 0.10 X 1.67 ± 0.08 mm), yolk-rich eggs. The larval period consists of three benthic, lecithotrophic larval stages, and lasts 10-11 days. The newly-hatched larvae bear very advanced morphological features such as antenna with several marginal plumose seta on scaphocerite and long, multi-articulated flagellum; fully developed, functional uniramous pereiopods 3-5 (walking legs) and biramous pleopods. The morphology of the carapace, all appendages of the cephalothorax and pleon, and the tail fan are described in detail and illustrated. The larval form was considered to be a decapodid because of the benthic behavior and due to the fact that functional walking legs and pleopods are the main structures for displacement and propulsion. The larval development of M. inpa is compared with those of the so-called continental group of the caridean shrimps from the Amazon River basin.

Morphology of First Zoeal Stage of Four Genera of Alvinocaridid Shrimps from Hydrothermal Vents and Cold Seeps: Implications for Ecology, Larval Biology and Phylogeny.

Alvinocaridid shrimps are endemic species inhabiting hydrothermal vents and/or cold seeps. Although indirect evidences (genetic and lipid markers) suggest that their larval stages disperse widely and support large scale connectivity, larval life and mechanisms underlying dispersal are unknown in alvinocaridids. Here we provide for the first time detailed descriptions of the first larval stage (zoea I) of four alvinocaridid species: Rimicaris exoculata and Mirocaris fortunata from the Mid-Atlantic Ridge, Alvinocaris muricola from the Congo Basin and Nautilocaris saintlaurentae from the Western Pacific. The larvae were obtained from onboard hatching of brooding females (either at atmospheric pressure or at habitat pressure in hyperbaric chambers) and from the water column near adult habitats, sampled with plankton pumps or sediment traps. Major characteristics of the alvinocaridid larvae include undeveloped mandible and almost complete absence of setation in the inner margin of the mouth parts and maxillipeds. Although the larvae are very similar between the four species studied, some morphological features could be used for species identification. In addition, undeveloped mouthparts and the large amount of lipid reserves strongly support the occurrence of primary lecithotrophy in the early stage of alvinocaridids. Although lecithotrophy in decapod crustaceans is usually associated with abbreviated larval development, as a mechanism of larval retention, morphological and physiological evidences suggest the occurrence of an extended and lecithotrophic larval stage in the Alvinocarididae. These traits permit the colonization of widely dispersed and fragmented environments of hydrothermal vents and cold seeps. Distribution of larval traits along the phylogenetic reconstruction of the Alvinocarididae and related families suggest that lecithotrophy/planktotrophy and extended/abbreviated development have evolved independently along related families in all potential combinations. However, the Alvinocarididae is the only taxa with a combination of lecithotrophy and extended larval development.

Influence of the larval phase on connectivity: strong differences in the genetic structure of brooders and broadcasters in the Ophioderma longicauda species complex.

Closely related species with divergent life history traits are excellent models to infer the role of such traits in genetic diversity and connectivity. Ophioderma longicauda is a brittle star species complex composed of different genetic clusters, including brooders and broadcasters. These species diverged very recently and some of them are sympatric and ecologically syntopic, making them particularly suitable to study the consequences of their trait differences. At the scale of the geographic distribution of the broadcasters (Mediterranean Sea and northeastern Atlantic), we sequenced the mitochondrial marker COI and genotyped an intron (i51) for 788 individuals. In addition, we sequenced 10 nuclear loci newly developed from transcriptome sequences, for six sympatric populations of brooders and broadcasters from Greece. At the large scale, we found a high genetic structure within the brooders (COI: 0.07 < F(ST) < 0.65) and no polymorphism at the nuclear locus i51. In contrast, the broadcasters displayed lower genetic structure (0 < F(ST) < 0.14) and were polymorphic at locus i51. At the regional scale, the multilocus analysis confirmed the contrasting genetic structure between species, with no structure in the broadcasters (global F(ST) < 0.001) and strong structure in the brooders (global F(ST) = 0.49), and revealed a higher genetic diversity in broadcasters. Our study showed that the lecithotrophic larval stage allows on average a 50-fold increase in migration rates, a 280-fold increase in effective size and a threefold to fourfold increase in genetic diversity. Our work, investigating complementary genetic markers on sympatric and syntopic taxa, highlights the strong impact of the larval phase on connectivity and genetic diversity.

Earliest ontogeny of Early Palaeozoic Craniiformea: compelling evidence for lecithotrophy

Popov, L.E., Bassett, M.G. & Holmer, L.E. 2012: Earliest ontogeny of Early Palaeozoic Craniiformea: compelling evidence for lecithotrophy. Lethaia, Vol. 45, pp. 566–573. The early ontogeny of Palaeozoic Craniiformea (Brachiopoda) remains controversial, with conflicting reports of evidence indicating lecithotrophic versus planktotrophic larval stages. Further compelling evidence for lecithotrophy in Palaeozoic craniiforms is described here. Newly obtained, well-preserved Silurian specimens of craniiforms, including Craniops (Craniopsida), and Lepidocrania? and Orthisocrania (Craniida) from Gotland and the St. Petersburg region, form the basis for this study. The new material demonstrates that the characters of shell structure and shell formation provide evidence of early differentiation of an adult dorsal mantle, and the presence of a distinctive primary layer with a characteristic lath-like pattern indicates that these craniiforms underwent a lecithotrophic larval stage, more or less identical to that of living. □Novocrania. Brachiopoda, Craniiformea, ontogeny, phylogeny, Early Palaeozoic.

Contrasting population makeup of two intertidal gastropod species that differ in dispersal opportunities

This study of population structure of two intertidal monodontine species: Diloma nigerrima and Diloma subrostrata, revealed the level of genetic connectedness among populations. Despite their markedly different geographic ranges (D. nigerrima is a geographically widespread species, inhabiting both Chile and New Zealand, including its subantarctic islands, whereas D. subrostrata is endemic to New Zealand), both species are believed to possess short-lived lecithotrophic larval stages. Polymorphic DNA microsatellite sequences were used to reveal the level of genetic connectedness among populations, thus inferring the two species' relative effective dispersal abilities. For each species F statistics, AMOVA values and the strength of the relationship between geographic and genetic distance were calculated. We observed a higher within-species level of genetic variation (ΦST=0.099 vs. ΦST=0.016) and a higher proportion of variance (11.15% vs. 0%) among populations of D. nigerrima than of D. subrostrata. A larger fraction of significant FST values was observed among D. nigerrima population pairs (65%) than among D. subrostrata population pairs (33%). Significant correlation between genetic and geographic distance was observed for D. nigerrima but not for D. subrostrata, but this relationship was not consistent among pairwise D. nigerrima population comparisons and PCA analysis confirms that, for each species, population structure does not follow a consistent pattern of increasing with geographic distance. The lack of population structure among D. subrostrata populations is probably due to its ubiquitous distribution, meaning little opportunity exists for genetic structure. D. nigerrima, by contrast has a patchier distribution, which allows for greater opportunities for genetic differentiation to occur. We argue that, despite the probable short larval stage in this species, the lack of geographical pattern in the genetic structure found in D. nigerrima is best explained by occasional dispersal over relatively short distances around the coast of New Zealand, over longer distances from New Zealand to the subantarctic islands and even across the Pacific Ocean from New Zealand to Chile.

Earliest ontogeny of Early Palaeozoic Craniiformea: implications for brachiopod phylogeny

Popov, L.E., Bassett, M.G., Holmer, L.E., Skovsted, C.B. & Zuykov, M.A. 2010: Earliest ontogeny of Early Palaeozoic Craniiformea: implications for brachiopod phylogeny. Lethaia, Vol. 43, pp. 323–333. Well preserved specimens of the Early Palaeozoic craniiform brachiopods Orthisocrania and Craniops retain clear evidence of a lecithotrophic larval stage, indicating the loss of planktotrophy early in their phylogeny. The size of the earliest mineralized dorsal shell was <100 μm across, and the well preserved shell structure in these fossil craniiforms allows their earliest ontogeny to be compared directly with that of living Novocrania, in which the first mineralized dorsal shell (metamorphic shell) is secreted only after settlement of the lecithotrophic larvae. Immediately outside this earliest shell (early post-metamorphic or brephic shell) and in the rest of the dorsal valve the primary layer in both fossil and living craniiforms has characteristic radially arranged laths, which are invariably lacking in the earliest dorsal shell. The ventral valve of the fossil specimens commonly preserves traces of an early attachment scar (cicatrix), which is equal in size to the dorsal metamorphic shell, and the brephic post-metamorphic ventral valve also has a primary shell with radially arranged laths. However, a primary shell with radial laths is completely lacking in the ventral valve of living Novocrania, indicating that heterochrony may have been involved in the origin of the encrusting mode of life in living craniids; the entire ventral valve of Recent craniids (with the possible exception of Neoancistrocrania) may correspond to the earliest attachment scar of some fossil taxa such as Orthisocrania. It is also probable that the unique absence of an inner mantle lobe as well as the absence of lobate cells in Novocrania could be the result of heterochronic changes. The dorsal valve of both fossil and living craniiforms has a marked outer growth ring, around 500 μm across, marking the transition to the adult, and a significant change in regime of shell secretion. The earliest craniiform attachment is considered to be homologous to the unique attachment structures described recently in polytoechioids (e.g. Antigonambonites) and other members of the strophomenate clade. However, unlike the craniiforms, polytoechioids and strophomenates all have planktotrophic larvae, and planktotrophy is most probably a plesiomorphic character for all Brachiopoda. □Brachiopoda, Craniiformea, Early Palaeozoic, ontogeny, phylogeny.

Reproduction among protobranch bivalves of the family Nuculidae from sublittoral, bathyal, and abyssal depths off the New England coast of North America

Abstract Protobranch bivalve species of the family Nuculidae pass through either a planktonic lecithotrophic larval stage or a direct non-planktonic development. Oogenesis of the three sublittoral species examined is synchronous. Deposition of egg masses by Nucula delphinodonta and spawning by Nucula annulata and Nucula proxima occur only during summer months. Among the four bathyal and abyssal species, Ennucula similis , Ennucula granulosa , Deminucula atacellana , and Brevinucula verrilli , oogenesis is asynchronous and there is no discernable pattern of periodicity of spawning. Absence of periodicity in reproduction in these deep-sea species is confirmed by examination of individuals from dredge samples taken at different times of the year. The median apparent fecundity among both sublittoral and deep-sea species is directly related to size (i.e. shell length) and age. Among the Nuculidae the median apparent fecundity is greater among sublittoral than bathyal and abyssal species. The geographic distribution of a species depends on its capacity to disperse. The dispersal of the planktonic lecithotrophic larvae of the sublittoral species N. annulata and N. proxima is limited to the continental shelf of the northwestern Atlantic by inshore bottom circulation and because these very small planktonic larvae ( D. atacellana .

Protobranch bivalves

The subclass Protobranchia comprises more than 600 species of bivalves that occur throughout the world ocean. Mostly deposit feeders in soft sediments, they are abundant in the deep sea. Apomorphies that unite them as a group include gill structure, hinge conformation, shell microstructure, larval development, foot morphology, respiratory pigments, trophic mode and digestion. They are relatively small and highly conserved in form, originating in the Cambrian era. They may represent an ancestral, derived or paraphylectic group of the Bivalvia. The protobranchs include two orders, the Nuculoida and Solemyoida, which previously were classified separately in the subclasses Paleotaxodonta and Cryptodonta, respectively. They are of ecological interest and have a unique functional morphology. They feed mostly under the surface of the sediment with highly modified labial palps, but the degree to which they are selective in diet remains difficult to determine. They are important bioturbators in many soft-sediment assemblages; their feeding and locomotion affects sediment structure and community development. Solemyoids are unusual in inhabiting reducing environments and hydrocarbon seeps and in deriving their nutrition from endosymbiotic chemosynthetic bacteria. A variety of species of protobranchs are found in oceanic trenches, near hydrothermal vents, and in submarine caves. Protobranchs produce a lecithotrophic larval stage, the pericalymma, making their development unique among bivalves. The pericalymma remains in the plankton for a short time and presumably has low dispersal ability. Recruitment may be intermittent. Growth is rapid in post-larvae but decreases with age, though rates may not necessarily be slow, especially in continental shelf species. Life spans are commonly 1 to 2 decades, but deep-sea representatives may grow more slowly and live longer. Bottom fish, seastars and gastropods are their major predators and a few parasites and commensals have been documented. The predominance of protobranchs in deep-sea sediments may be a result of deep-sea origin or displacement from shallow waters by lamellibranchs. Their ability to deposit-feed, digest food extracellularly, and develop by means of lecithotrophic larvae make them particularly well adapted to cold and oligotrophic habitats.

The genetic structure of intertidal populations of two species of nudibranch molluscs with planktotrophic and pelagic lecithotrophic larval stages: are pelagic larvae “for” dispersal?

Planktonic larvae of benthic marine invertebrates may confer larval dispersal and colonization potential on the reproductive adult. Analyses of genetic heterogeneity of local populations provide an indirect means of assessing gene flow and the scale of realized dispersal of larval migrants. Here we compared populations of two species of nudibranchs with long term planktotrophic ( Goniodoris nodosa) and shorter term pelagic lecithotrophic ( Adalaria proxima) veliger stages. Both species have annual life cycles, population turnover is absolute and free-swimming larvae hatch from benthic spawn masses. G. nodosa larvae are pelagic for up to ∼3 months, whereas those of A. proxima can metamorphose within 1–2 d of release although, at least in the laboratory, they may delay metamorphosis for several weeks. The broad ranges of pelagic larval transport potential were estimated at 10 2–10 3 km for G. nodosa and 10 1–10 2 km for A. proxima: significant differentiation of populations of G. nodosa was predicted to occur on a scale of 10 3–10 4 km, whereas for A. proxima differentiation was expected on a scale of perhaps only 10 2–10 3 km. For both species, analyses were undertaken of allozyme variation among sites throughout the 1600 km British Isles range of A. proxima. F-statistic analyses showed no evidence of population differentiation for G. nodosa (11 populations). By contrast, comparable data for 19 populations of A. proxima showed very high levels of differentiation, but no obvious pattern of variation or significant isolation by distance. More detailed sampling of nine populations of A. proxima over a 26.18 km range in W Scotland – in a location characterised by highly dispersive tidal currents – revealed (1) similar levels of differentiation to the 1600 km range, (2) clear distance-related clinal patterns of variation, and (3) a significant inverse relationship between genetic differentiation and geographic distance. For Adalaria proxima there was evidence of significant differentiation of populations on a scale of ≤10 1 km. This was counter to our expectations of larval transport potential for A. proxima and implies that these larvae are behaviourally adapted to avoid dispersal in the water column and thereby recruit locally. Whilst the planktotrophic species, G. nodosa, conformed to our expectations, the clear implication of the data for A. proxima is that possession of a pelagic larval stage does not necessarily facilitate dispersal and colonisation potential for this sedentary invertebrate. Extrapolations and predictions of dispersal potential, and gene flow in natural populations, based on laboratory observations of the larval phase duration may be markedly error-prone.

Carrying on regardless: the echinoid genus Cyclaster at the Cretaceous–Tertiary boundary

A taxonomic revision and subsequent cladistic analysis show that Maastrichtian and Palaeocene species of the genus Cyclaster are split into three clades. Members of all three are present before and after the Cretaceous-Tertiary boundary, demonstrating that the genus Cyclaster as a whole was unaffected by the crisis at the end of the Cretaceous. Unlike most spatangoids, Cyclaster has the characteristic crystallographic signature of a lecithotrophic larval stage. Further work on other groups will determine whether larval strategy affected survivorship at this time.

Head Kidneys in Hatchlings of Scoloplos Armiger (Annelida: Orbiniida): Implications for the Occurrence of Protonephridia in Lecithotrophic Larvae

It is generally believed that lecithotrophic larvae of annelids do not possess functional excretory organs. However, as in certain annelids the planktotrophic trochophora larva has been secondarily modified into a lecithotrophic developmental stage and because protonephridia are characteristic for the trochophora, lecithotrophic developmental stages should also possess such organs. To test this assumption hatchlings of the orbiniidan Scoloplos armiger, which develops directly without a free-living larval stage, were investigated ultrastrucrurally. Each hatchling possesses a pair of protonephridia which lie caudal to the eyes and almost level with the frontal margin of the foregut. Each organ consists of three multiciliated cells, a terminal cell, a duct cell and a nephropore cell. The terminal cell bears a distally oriented hollow cytoplasmic cylinder, which surrounds the cilia. Adherens junctions connect this structure to the duct cell. Several clefts and pores perforate the wall of the hollow cylinder. Extracellular material covers the pores and clefts and thus may function as a molecular sieve during filtration. A comparison with the protonephridia of other annelid larvae reveals: (1) that one pair of protonephridial head kidneys consisting of a terminal cell, a duct cell and a nephropore cell must be assumed for the trochophore in the ground pattern of annelids and (2) that these organs are preserved when lecithotrophic larval stages evolved within the Annelida

Population genetics of Trochus niloticus and Tectus coerulescens, topshells with short-lived larvae

Trochus niloticus L. and Tectus coerulescens Lmk., two coral reef trochid gastropods that have similar life-histories including a lecithotrophic larval stage, were sampled from reefs in the northern, central and southern sections of the Great Barrier Reef (GBR) in 1991. Significant sex-ratio biases were noted, and these varied among reefs, apparently with latitude. Demographic data suggested that highly discontinuous and localized recruitment occurs. Surveys of allozyme frequencies at 12 loci revealed no significant genetic differences among populations of Trochus niloticus at any geographical scale. High gene flow between zones of the GBR was inferred, with the number of migrants per generation (Nm) of the order of 100. Directional selection was thought to occur at one locus (GDH*). Genetic variability in T. niloticus (H=0.069 to 0.110, and only three loci polymorphic) was low compared with other trochids. This was thought to be due to smaller effective population size, resulting from an unbalanced sexratio, aggregative spatial distribution of adults, high variance in reproductive success, and/or the occurrence of population extinctions and recolonisations. In contrast, Tectus coerulescens exhibited significant genetic differences between zones, indicating a much lower rate of migration between populations (Nm∼1 to 10), and displayed high genetic diversity (H=0.225 to 0.279). A trend for increasing genetic diversity from the northern to southern GBR was found in both species. It is not clear whether the occurrence of two contrasted population genetic structures in species with apparently similar life-histories is due to ecological or historical factors.

Evolutionary Conservation of the Larval Serotonergic Nervous System in a Direct Developing Sea Urchin: (sea urchin development/larval nervous systems/heterochrony/direct development/Heliocidaris erythrogramma).

Development of the larval serotonergic nervous system is examined by indirect immunofluorescence in two congeneric species of sea urchins that exhibit divergent embryonic and larval development. Heliocidar is tuberculata undergoes indirect planktotrophic development via a pluteus larva, whereas Heliocidaris erythrogramma develops directly, passing through a brief, highly derived lecithotrophic larval stage. We have cleared the opaque embryos of H. erythrogramma and discuss internal features of its development. The serotonergic nervous system of H. tuberculata arises in the apical plate at the end of gastrulation and develops into a bilaterally symmetric ganglion lying between the anterolateral arms in the preoral hood. Putatively homologous neurons appear at the apical end of the modified larva of H. erythrogramma well after the completion of gastrulation, coincident with development of the primary podia of the adult rudiment. The neurons form a bilaterally symmetric ganglion whose orientation relative to the vestibule is conserved with respect to that found in planktotrophic larvae. This allows us to define a left and right side for this larva which lacks external points of asymmetry such as a larval mouth. The alteration in the time of nervous system development in H. erythrogramma relative to that of H. tuberculata, and other indirect developers, implicates heterochronies in cellular differentiation as an important component of the evolution of direct development.

Spirorbidae (Polychaeta) collected by R. V. “Vitjas” from abyssal depths of the Pacific Ocean

Pileolaria levensteinae sp. nov. was represented by juvenile and presumably adult stages, attached to ferromanganese substrata. None bore embryos, but the larger individuals had deep opercular cups and the species is closely related to certain shallow water forms which incubate in such cups. A second species was dextrally coiling but represented by a single fragment too small to describe. Spirorbids would appear to be preadapted for abyssal life, by having a brief, lecithotrophic larval stage.