Novel Chemoautotrophic Endosymbiosis between a Member of the Epsilonproteobacteria and the Hydrothermal-Vent Gastropod Alviniconcha aff. hessleri (Gastropoda: Provannidae) from the Indian Ocean

The hydrothermal-vent gastropod Alviniconcha aff. hessleri from the Kairei hydrothermal field on the Central Indian Ridge houses bacterium-like cells internally in its greatly enlarged gill. A single 16S rRNA gene sequence was obtained from the DNA extract of the gill, and phylogenetic analysis placed the source organism within a lineage of the epsilon subdivision of the Proteobacteria. Fluorescence in situ hybridization analysis with an oligonucleotide probe targeting the specific epsilonproteobacterial subgroup showed the bacterium densely colonizing the gill filaments. Carbon isotopic homogeneity among the gastropod tissue parts, regardless of the abundance of the endosymbiont cells, suggests that the carbon isotopic composition of the endosymbiont biomass is approximately the same as that of the gastropod. Compound-specific carbon isotopic analysis revealed that fatty acids from the gastropod tissues are all ¹³C enriched relative to the gastropod biomass and that the monounsaturated C₁₆ fatty acid that originates from the endosymbiont is as ¹³C enriched relative to the gastropod biomass as that of the epsilonproteobacterial cultures grown under chemoautotrophic conditions. This fractionation pattern is most likely due to chemoautotrophy based on the reductive tricarboxylic-acid (rTCA) cycle and subsequent fatty acid biosynthesis from ¹³C-enriched acetyl coenzyme A. Enzymatic characterization revealed evident activity of several key enzymes of the rTCA cycle, as well as the absence of ribulose-1,5-bisphosphate carboxylase/oxygenase activity in the gill tissue. The results from anatomic, molecular phylogenetic, bulk and compound-specific carbon isotopic, and enzymatic analyses all support the inference that a novel nutritional strategy relying on chemoautotrophy in the epsilonproteobacterial endosymbiont is utilized by the hydrothermal-vent gastropod from the Indian Ocean. The discrepancies between the data of the present study and those of previous ones for Alviniconcha gastropods from the Pacific Ocean imply that at least two lineages of chemoautotrophic bacteria, phylogenetically distinct at the subdivision level, occur as the primary endosymbiont in one host animal type.     

Molecular phylogenetic and isotopic evidence of two lineages of chemoautotrophic endosymbionts distinct at the subdivision level harbored in one host-animal type: the genus Alviniconcha (Gastropoda: Provannidae)

The hydrothermal-vent gastropod Alviniconcha hessleri from the Alice Springs deep-sea hydrothermal field in the Mariana Back-Arc Basin in the Western Pacific houses an intracellular bacterial endosymbiont in its gill. Although enzymatic analysis has revealed that the endosymbiont is a sulfur-oxidizing chemoautotroph using the Calvin-Benson cycle for the fixation of carbon dioxide, the phylogenetic affiliation of, and the trophic relationship of A. hessleri with, the chemoautotrophic endosymbiont remains undetermined. A single 16S rRNA gene sequence was obtained from the DNA extract of the gill, and phylogenetic analysis placed the source organism within the lineage of the gamma subdivision of the Proteobacteria that consists of many chemoautotrophic endosymbionts of marine invertebrates. Fluorescence in situ hybridization analysis showed the bacterium densely colonizing the gill filaments. The fatty acid profile of the symbiont-free mantle contains the high level of the 16:1 fatty acid originating from the endosymbiont, which indicates that the endosymbiont cells are digested by, and incorporated into, the host. Compound-specific carbon isotopic analysis revealed that fatty acids from the gastropod tissues are all (13)C-depleted relative to the gastropod biomass. This fractionation pattern is consistent with chemoautotrophy based on the Calvin-Benson cycle and subsequent fatty-acid biosynthesis from (13)C-depleted acetyl coenzyme A. The results from the present study are clearly different from those from our previous study for A. aff. hessleri from the Indian Ocean that harbors a chemoautotrophic endosymbiont belonging to the epsilon subdivision of the Proteobacteria, which mediates the reductive tricarboxylic acid cycle for carbon fixation. Thus, it is concluded here that two lineages of chemoautotrophic bacteria, phylogenetically distinct at the subdivision level, occur as the primary endosymbiont in one host-animal type, which is unknown for the other metazoans.     

Novel forms of structural integration between microbes and a hydrothermal vent gastropod from the Indian Ocean

Here we describe novel forms of structural integration between endo- and episymbiotic microbes and an unusual new species of snail from hydrothermal vents in the Indian Ocean. The snail houses a dense population of gamma-proteobacteria within the cells of its greatly enlarged esophageal gland. This tissue setting differs from that of all other vent mollusks, which harbor sulfur-oxidizing endosymbionts in their gills. The significantly reduced digestive tract, the isotopic signatures of the snail tissues, and the presence of internal bacteria suggest a dependence on chemoautotrophy for nutrition. Most notably, this snail is unique in having a dense coat of mineralized scales covering the sides of its foot, a feature seen in no other living metazoan. The scales are coated with iron sulfides (pyrite and greigite) and heavily colonized by epsilon- and delta-proteobacteria, likely participating in mineralization of the sclerites. This novel metazoan-microbial collaboration illustrates the great potential of organismal adaptation in chemically and physically challenging deep-sea environments.     

Aplysiid species from East Africa with notes on the Indian Ocean Aplysiomorpha (Gastropoda: Opisthobranchia)

Eleven species of the opisthobranch order Aplysiomorpha collected from East Africa and neighbouring parts of the Indian Ocean are described. Three of the species are new to science. The nineteen species of Aplysiomorpha which are reliably reported from the Indian Ocean are listed, together with a key for their identification.     

Molecular taxonomy and naming of five cryptic species of Alviniconcha snails (Gastropoda: Abyssochrysoidea) from hydrothermal vents

Large symbiont-hosting snails of the genus Alviniconcha (Gastropoda: Abyssochrysidae) are among the dominant inhabitants of hydrothermal vents in the Western Pacific and Indian oceans. The genus was originally described as monotypic, but unique DNA sequences for mitochondrial genes revealed six distinct evolutionary lineages that we could not distinguish based on external morphology. Subsumed under the name Alviniconcha hessleri Okutani & Ohta, the distinct allopatric and sympatric lineages have been assigned placeholder epithets that complicate scientific communications. Based on the present multi-gene sequence data, we hereby describe five Alviniconcha species (in the order of their discovery) – A. kojimai sp. nov., A. boucheti sp. nov., A. marisindica sp. nov., A. strummeri sp. nov. and A. adamantis sp. nov. Thus, we restrict application of the name A. hessleri to specimens that are genetically similar (≥95% for COI) to those found at localities in the Mariana Trough. Single distinct Alviniconcha species inhabit vent fields along the Central Indian Ridge, the Mariana volcanic arc, and the Mariana back-arc basin, whereas vents in the Manus, Fiji and Lau back-arc basins may host two or three additional species. Formal recognition of these species facilitates future attempts to assess their physiological differences and symbiont associations. Furthermore, their reported distributions have significant biogeographic implications, affecting estimates of the diversity within and overlap among Indo-Pacific vent localities.

http://zoobank.org/urn:lsid:zoobank.org:pub:1E4B2E71-9F1D-479E-9A9A-22A9E303AAE5     

Trochus kotschyi, the first Indian Ocean record of the genus Osilinus (Mollusca: Gastropoda: Trochidae)

The species Trochus kotschyi Philippi, 1849, and Priotrochus obscurus (Wood, 1828) have been much confused in the past. Consistent differences between these taxa in respect of shell morphology, external anatomy and radular and opercular form are demonstrated. The distribution of T. kotschyi and the extent to which it and P. obscurus are sympatric are discussed. T. korschyi is referred to the genus Osilinus Philippi, 1847, a new combination and the first record of the genus from the Indo-West Pacific. The species is thought to be a pre-Miocene relict reflecting a mid-Tethyan origin of the genus. A note on the type species designation of Osilinus is provided.     

Pseudobithynia guldeni sp. n., a new gastropod species from the Mediterranean region of Turkey (Gastropoda: Truncatelloidea)

A new Bithyniidae species Pseudobithynia guldeni sp.n. is described from the Mediter-ranean region of Turkey based on shell and male genitalia morphology. An identification key for the species of Pseudobithynia of Turkey is given.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:FA659D77-5B02-4C2F-9A6D-D61E676778E1     

Novel Forms of Structural Integration between Microbes and a Hydrothermal Vent Gastropod from the Indian Ocean

Here we describe novel forms of structural integration between endo- and episymbiotic microbes and an unusual new species of snail from hydrothermal vents in the Indian Ocean. The snail houses a dense population of γ-proteobacteria within the cells of its greatly enlarged esophageal gland. This tissue setting differs from that of all other vent mollusks, which harbor sulfur-oxidizing endosymbionts in their gills. The significantly reduced digestive tract, the isotopic signatures of the snail tissues, and the presence of internal bacteria suggest a dependence on chemoautotrophy for nutrition. Most notably, this snail is unique in having a dense coat of mineralized scales covering the sides of its foot, a feature seen in no other living metazoan. The scales are coated with iron sulfides (pyrite and greigite) and heavily colonized by - and δ-proteobacteria, likely participating in mineralization of the sclerites. This novel metazoan-microbial collaboration illustrates the great potential of organismal adaptation in chemically and physically challenging deep-sea environments.     

The circatidal rhythm of the estuarine gastropod Hydrobia ulvae (Gastropoda: Hydrobiidae)

Intertidal animals display a suite of cyclic behaviours that evolved as adaptations to the predictable cycle of inundation and exposure. In estuarine habitats, mud snails from the genus Hydrobia are among the most abundant grazers, and have received considerable attention with respect to the behavioural mechanisms mediating locomotion, dispersal, and feeding, although the nature of the control of these processes has remained elusive. In particular, it is not clear whether endogenous activity patterns are related to periodic changes of microphytobenthos biomass at the sediment surface, or whether they are timed to the tidal cycle at all. In the present study, we address the crawling activity of Hydrobia ulvae under constant conditions, as well as the effects of individual size and previous short‐term exposure to tides of different range, by recording immersed individual snails under constant dark conditions. We show that the species displays an overt circatidal pattern of crawling, with activity peaks around high water, and that the start of inundation may act as an entrainment agent of the rhythm. Moreover, the results obtained indicate that smaller snails display higher levels of activity, although neither the size nor previous in situ influence of tidal range has an effect on the period and on the amplitude of the rhythm. These findings suggest that fluctuations of microphytobenthos biomass are not a sufficiently strong selective pressure to have shaped locomotor activity in H. ulvae. Moreover, feeding of H. ulvae should take place mostly during high water and be independent of periodic fluctuations of microphytobenthos biomass at the surface of the sediment. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 439–450.     

Provannid and provannid‐like gastropods from the Late Cretaceous cold seeps of Hokkaido (Japan) and the fossil record of the Provannidae (Gastropoda: Abyssochrysoidea)

A fauna of provannid and provannid‐like shells is described from Upper Cretaceous seep carbonates in Hokkaido, Japan. We describe two new provannid species, Provanna tappuensis sp. nov. and Desbruyeresia kanajirisawensis sp. nov. , with preserved protoconchs of unquestionable provannid type with decollate apex. This material confirms the occurrence of Provannidae as early as the Middle Cenomanian. We also describe Hokkaidoconcha gen. nov. and a new family Hokkaidoconchidae fam. nov. , with two named species, H. hikidai sp. nov. and H. tanabei sp. nov . Hokkaidoconchidae are possibly related to the Provannidae, judging from a similar, but not decollate larval shell, although the juvenile teleoconch whorls differ in being of a general cerithimorph appearance and the details of the aperture are unknown. Furthermore, we review the published fossil record of Provannidae and Abyssochrysidae, and we consider that in those older than the Eocene, there is no evidence preserved that unequivocally supports a position there. The Jurassic Acanthostrophia acanthica from Italy seems to be the oldest known record of Abyssochrysidae, and the most reliable occurrence of the family, older than from the Miocene. Other fossil, pre‐Miocene species that have been classified in the Abyssochryssidae are provisionally referred to Hokkaidoconchidae. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 421–436.     

Siphonostomes (Copepoda -- Cyclopoida) from the Indian Ocean

Siphonostomes (Copepoda — Cyclopoida) from the InternationalIndian Ocean Expedition collections were studied. The groupwas represented by Pontoeciella abyssicola of family Pon-toeciellidaeand Ratania flava of family Rataniidae. This study has extendedthe distribution of both the species considerably, and includesthe first record of the male of P. abyssicola from the IndianOcean and of both species from the Bay of Bengal.     

A new type of gastropod proboscis: the foregut of Hastula bacillus (Gastropoda: Terebridae)

Hastula bacillus (Deshayes) is a small terebrid gastropod which inhabits sandy surf beaches in southern Thailand, where it feeds upon spionid polychaetes. It possesses a foregut anatomy unlike that of any other gastropod. An elongate arborescent muscular organ, known as the accessory proboscis structure, is extended through the mouth during foraging. When retracted, it is folded into an 's' shape in the permanent rhynchodeum. The accessory proboscis structure bears numerous tufts of short, stiff cilia which are associated with pairs or triplets of dome-like structures. It is suggested that the structures may be chemosensory and concerned with prey location. Hastula bacillus also possesses a retractable labial tube, a long proboscis and buccal tube, dart-shaped radular teeth, an odontophore, an accessory salivary gland, a pair of salivary glands and a well-developed venom gland with muscular bulb. A comparison with other terebrid species suggests that H. bacillus is the most plesiomorphic taxon yet described from the family.     

Structure of shell muscles in the Cambrian gastropod genus Bemella (Gastropoda: Archaeobranchia: Helcionellidae)

Zones with peculiar microornamentation interpreted as muscle scars were found on the internal molds of the Early Cambrian gastropod genus Bemella Missarzhevsky, 1969 (family Helcionellidae). Shell muscles of helcionellids are reconstructed based on the topographic pattern of muscle scars, i.e., the pedal, cephalic, and mantle retractors are recognized. The reconstruction proposed here of the shell musculature corroborates affinity between ancient gastropod and helcionelloid mollusks.     

New species, anatomy, and systematic position of the hydrothermal vent and hydrocarbon seep gastropod family Provannidae fam.n. (Caenogastropoda)

The anatomy of the hydrothermal vent gastropod Provanna is described. It is shown to be a primitive neotaenioglossan with mainly plesiomorphic features in all systems, including open pallial gonoducts, epiathroid nervous system with accessory pedal ganglia and a circulatory system where a considerable portion of the venous blood from the coelomic spaces enters the pallial skirt. Apomorphic features include the possession of an anterior notch, or short canal in the shell (and mantle edge), an annulated sensory pallial tentacle, paraspermatozoa modificd as nurse cells. multiple seminal receptacles, a posterior bursa copulatrix, and a renal oviduct with three distinct regions. the middle one expanded. Comparisons are madc with Abyssochrysos, a genus assumed to be a relict member of the Loxonematoidea, which appears to be the closest known relative. A new family. Provannidae fam.n., is created and tentatively included in the Loxonematoidea. The following new specics are described: Provanna segonzaci sp.n. (from hydrothermal vents at the Fiji Back Arc), P. laevis sp.n. (from hydrothermal seeps in the Gulf of California), P. sculpta sp.n. and P. admetoides sp.n. (from hydrocarbon seeps in the Gulf of Mexico). New records are added for previously described species of Provanna.     

Monognathus berteli sp. nov. from the Indian Ocean (Pisces,Monognathidae)

A new deepsea monognathid species,Monognathus berteli, is described based on one specimen collected pelagically in the northwestern Indian Ocean at 1440–1018 meters of depth. LikeM. taningi andM. bertini, it belongs to the short-skulled species-group and has very long pectoral fins, but differs from these species by several meristic characters. The described specimen has a very long caudal filament (53% SL) which is not known from otherMonognathus. However, the fragile filament is not used as a diagnostic character since it could easily be broken.     

Acantholaimus (chromadoridae:nematoda) from the Indian Ocean: description of seven species

Seven species of the Acantholaimus aredescribed. Acantholaimus vermeuleni sp.n. ischaracterised by labial and cephalic sensilla thatare located at the same level, two post amphidialsetae on the dorso-lateral side and a poorlydeveloped stoma without distinct teeth. Acantholaimus verscheldi sp.n. is characterised bya narrow elongate pharyngeal region, a long stomawith distinct teeth and short (4–7 µm) cephalicsensilla. Acantholaimus heipi sp.n. ischaracterised by a narrow elongate pharyngealregion, well developed teeth in the stoma and longcephalic sensilla (11–13 µm). Acantholaimuselegans Jensen 1988 has a narrow anteriorpharyngeal region that increases in lengthposteriorly, it has setae before and after theamphids on both sides and stoma with well developedteeth. Acantholaimus gathumai sp.n. ischaracterised by long cephalic (10–15 µm) andsomatic (8–10 µm) setae and lateraldifferentiation with fine dots (5–7 µm inwidth). Acantholaimus geraerti sp.n. has longcephalic sensilla (15–19 µm) and narrow (4–6 µm)distinct lateral differentiation. Acantholaimusinvaginatum sp.n. is characterised byvery long cephalic setae (16–21 µm), severalsetae at the pharyngeal region and wide lateraldifferentiation with fine dots, often the stoma isinvaginated. Distribution of the different species of Acantholaimus encoutered in the four transectsstudied is also described.     

Egg capsules, eggs and embryos of the southwestern Atlantic gastropod Coronium coronatum (Gastropoda: Muricidae)

The egg capsules, eggs and embryos of the muricid gastropodCoronium coronatum are described for the first time. Capsulesare sessile, bulliform, semi-circular, with a plug in the dorsalcenter. Sutures split the capsule into two asymmetrical halves.Recently laid capsules showed the presence of 3639 (n = 2) uncleavednurse eggs with a diameter of 180–210 µm (mean= 197.4 ± 8.9). The number of early embryos was 9–11.The size of the embryos was 320 x 320 to 820–880 µm.Nine pre-hatching embryos of 3.94 mm (n = 8, SD = 0.32)were found inside the older capsule. SEM illustrations of embryosand radulae are provided. Comparison of shell and radula ofembryos with the protoconch and radulae of adults of C. coronatumrevealed that the capsule belongs to this species. (Received 18 March 2006; accepted 10 October 2006)     

Gonad characterization and reproductive seasonality in Siphonaria lessonii (Gastropoda: Heterobranchia) from the southwestern Atlantic Ocean

Histological characterization of the hermaphroditic gonad (HG) and seasonality of gametogenesis were investigated in a population of Siphonaria lessonii from the coast of Buenos Aires Province (37°16′S, 56°58′W). Monthly analysis of the frequency of gametogenic stages, as well as the number and mean size of oocytes, were used to determine reproductive events over a 2‐year period (June 2012–May 2014). Female and male gametes were observed simultaneously within acini of adult individuals and continuously throughout the period studied. Oogenesis commenced in the beginning of austral autumn, with gonads characterized mainly by proliferation of female cells. From this moment, oocytes gradually increased in number and area until spring, when a large number of individuals were found in the evacuation stage. The same trend was observed from early gonad maturation to advanced stages, indicating that gonad development was closely related to the frequency of oocyte stages and to the area (size) of oocytes. Spermatogenesis was also observed as a continuous process throughout the year, although spent acini were more frequent from November until February. Reproductive seasonality and gametogenesis were associated with changes in temperature and day length.