Identification of Phyllosoma Larvae of the Lobsters Acantharctus and Biarctus (Crustacea: Scyllaridae) from the SW Indian Ocean.

The accurate assignment of cryptic larvae to species-level is a key aspect of marine ecological research and can be achieved through integrated molecular and morphological studies. A combination of two mitochondrial markers (COI and 16S) and a detailed morphological analysis was used to identify phyllosoma larvae of slipper lobster (Scyllaridae) species collected during a survey in the SW Indian Ocean. Two morphotypes were tentatively assigned to Acantharctus ornatus and Biarctus pumilus, both genera for which the larval morphology was unknown. Morphological revision of an adult specimen used to generate the putative A. ornatus sequences in GenBank revealed that it was misidentified and corresponds to B. dubius. The final phyllosoma stage of B. pumilus and subfinal and final stages of A. ornatus were described, clarifying prior misidentifications in the literature. Scyllarid biodiversity in the SW Indian Ocean is underestimated and sampling of deeper water layers is recommended to complete current knowledge of species and larval stages present in the region.

Phylogeny of sea spiders (Arthropoda: Pycnogonida) inferred from mitochondrial genome and 18S ribosomal RNA gene sequences.

The phylogeny of sea spiders has been debated for more than a century. Despite several molecular studies in the last twenty years, interfamilial relationships remain uncertain. In the present study, relationships within Pycnogonida are examined in the light of a new dataset composed of 160 mitochondrial genomes (including 152 new sequences) and 130 18S rRNA gene sequences (including 120 new sequences), from 141 sea spider morphospecies representing 26 genera and 9 families. Node congruence between mitochondrial and nuclear markers was analysed to identify the most reliable relationships. We also reanalysed a multilocus dataset previously published and showed that the high percentages of missing data make phylogenetic conclusions difficult and uncertain. Our results support the monophyly of most families currently accepted, except Callipallenidae and Nymphonidae, the monophyly of the superfamilies Ammotheoidea (Ammotheidae + Pallenopsidae), Nymphonoidea (Nymphonidae + Callipallenidae), Phoxichilidioidea (Phoxichilidiidae + Endeidae) and Colossendeoidea (Colossendeidae + Pycnogonidae + Rhynchothoracidae), and the sister-group relationship between Ammotheoidea and Phoxichilidioidea. We discuss the morphological evolution of sea spiders, identifying homoplastic characters and possible synapomorphies. We also discuss the palaeontological and phylogenetic arguments supporting either a radiation of sea spiders prior to Jurassic or a progressive diversification from Ordovician or Cambrian.

A new deep-water palaemonid shrimp genus and species from the French Antilles, with a new record of Periclimenes milleri Bruce, 1986 (Decapoda: Caridea).

A new genus and species in the caridean shrimp family Palaemonidae is described based on three type specimens collected at a depth range of 208-385 m off Guadeloupe, French Antilles. Zoukaris festivus gen. et sp. nov. shares many characters with several western Atlantic deep-water species currently assigned to Periclimenes Costa, 1844, as well as with the monotypic western Atlantic genus Diapontonia Bruce, 1986 and the Indo-West Pacific genus Echinopericlimenes Marin Chan, 2014. Zoukaris gen. nov. can be separated from all of them by a unique combination of morphological features, especially the configuration of the dactylus of the ambulatory pereiopods. In addition, Periclimenes milleri Bruce, 1986 is recorded from the French Antilles based on a single specimen, also from Guadeloupe; its colour pattern is illustrated for the first time.

Dorotea gen. nov., a new bathyal genus (Amphipoda, Eusiridae) from the Solomon Sea (Papua New Guinea).

A new species ascribed to a new genus of Eusiridae, Dorotea papuana gen. nov., sp. nov. is described from bathyal bottoms of the Solomon Sea (Papua New Guinea). Closely related to the genus Cleonardo, this new genus can be distinguished from most other known eusirid genera by the presence of a telson distally cleft and distinctly bilobate, of a distal spiniform process on uropod 1 peduncle and of a simple, stout and medium length dactylus on pereopods 5-7. The combination of all these characters can be considered relevant for the affiliation of this species to a new genus within Eusiridae. Due to its very close morphological affinity to Dorotea gen. nov., the bathyal sub-Antarctic species Eusiroides aberrantis Bellan-Santini Ledoyer, 1987 cannot be maintained in the family Pontogeneiidae and it should be transferred to the   family Eusiridae.

A new species of the coral associated barnacle (Thoracica: Pyrgomatidae: Pyrgoma) from a deep-sea oculinid coral in New Caledonian waters.

The present study describes a new species of Pyrgoma Leach, 1817, a coral associated barnacle attached to Tubastrea, from the south of New Caledonia. Pyrgoma spurtruncata sp. nov. is morphologically close to P. cancellatum Leach, 1818, P. japonica Weltner, 1897 and P. kuri Hoek, 1913 in the absence of extended tergal muscle crests. Pyrgoma cancellatum and P. kuri have a shallow, fully open, medial furrow of the tergal spur, whereas in P. spurtruncata sp. nov. the medial furrow is deeper and closed. Pyrgoma spurtruncata sp. nov. differs from P. japonica Weltner, 1897 in the width of the tergal spur and the length of the rostral tooth of the scutum. Phylogenetic analyses based on two mitochondrial markers, 12S rDNA and COI, confirm a unique, distinct clade of P. spurtruncata sp. nov. among the current available molecular information regarding Pyrgoma species.

Gill chamber and gut microbial communities of the hydrothermal shrimp Rimicaris chacei Williams and Rona 1986: A possible symbiosis.

Rimicaris chacei Williams and Rona 1986, formerly named as Chorocaris chacei, is a caridean shrimp living in deep-sea hydrothermal ecosystems. This shrimp is endemic to the Mid Atlantic Ridge (MAR) and lives at the periphery of aggregates of its well-known congeneric R. exoculata Williams and Rona 1986. Contrasting with the very dense and mobile clusters formed by R. exoculata, R. chacei lives in small groups of several individuals that are not very mobile. Although devoid of the characteristic hypertrophied cephalothorax of R. exoculata, which harbors the ectosymbionts, a microbial community has also been reported in the cephalothorax of R. chacei. Previous data on morphology, behavior and isotopic values indicate a diet based on a combination of feeding on its epibiotic bacteria and scavenging or occasional predation. In this study, our objective was to describe, for the first time, the distribution, morphology and phylogeny of the microbial communities associated with R. chacei. This species is significantly less studied than R. exoculata, but nevertheless represents the only other known example of symbiosis in crustaceans of MAR hydrothermal vent sites. Microbial communities have been observed at the same locations as in R. exoculata (mouthparts, branchiostegites and digestive tract). However, in R. chacei, the surfaces occupied by the bacteria are smaller. The main lineages are affiliated to Epsilon and Gammaproteobacteria in the cephalothorax and to Deferribacteres, Mollicutes, Epsilon and Gammaproteobacteria in the digestive tract. Comparison with the well-described bacterial communities of R. exoculata and hypotheses about the role of these communities in R. chacei are discussed.

A genetic fingerprint of Amphipoda from Icelandic waters - the baseline for further biodiversity and biogeography studies.

Amphipods constitute an abundant part of Icelandic deep-sea zoobenthos yet knowledge of the diversity of this fauna, particularly at the molecular level, is scarce. The present work aims to use molecular methods to investigate genetic variation of the Amphipoda sampled during two IceAGE collecting expeditions. The mitochondrial cytochrome oxidase subunit 1 (COI) of 167 individuals originally assigned to 75 morphospecies was analysed. These targeted morhospecies were readily identifiable by experts using light microscopy and representative of families where there is current ongoing taxonomic research. The study resulted in 81 Barcode Identity Numbers (BINs) (of which >90% were published for the first time), while Automatic Barcode Gap Discovery revealed the existence of 78 to 83 Molecular Operational Taxonomic Units (MOTUs). Six nominal species (Rhachotropis helleri, Arrhis phyllonyx, Deflexilodes tenuirostratus, Paroediceros propinquus, Metopa boeckii, Astyra abyssi) appeared to have a molecular variation higher than the 0.03 threshold of both p-distance and K2P usually used for amphipod species delineation. Conversely, two Oedicerotidae regarded as separate morphospecies clustered together with divergences in the order of intraspecific variation. The incongruence between the BINs associated with presently identified species and the publicly available data of the same taxa was observed in case of Paramphithoe hystrix and Amphilochus manudens. The findings from this research project highlight the necessity of supporting molecular studies with thorough morphology species analyses.

First report on the complete mitochondrial genome of the deep-water scalpellid barnacle Arcoscalpellum epeeum (Cirripedia, Thoracica, Scalpellidae).

Scalpellids are one of the largest families of Scalpelliformes and reproduce either androdioeciously or dioeciously. Here, we characterized the first mitogenome of a scalpellid barnacle (Arcoscalpellum epeeum), which was 15,593 bp in length with a 71.5% AT content. In comparison with the pollicipedids Capitulum mitella and Pollicipes polymerus, the tRNA genes of A. epeeum were rearranged between ND3 and ND5, between CYTB and ND1, and between 12S rRNA and ND2. On the mitogenomic tree, the Scalpelliformes families Pollicipedidae and Scalpellidae were not monophyletic, which concurs with previous studies.

A preliminary assessment of the deep-sea Decapoda collected during the KARUBENTHOS 2015 Expedition to Guadeloupe Island.

A preliminary assessment of the deep-sea Decapoda is proposed for Guadeloupe Island based solely on high definition macro photographs taken during the KARUBENTHOS 2015 Expedition to the Island (R/V Antea, 7-29 June 2015). Overall, 190 species are recognized, several of which are depicted with their fresh color for the first time. Previous records in the Lesser Antilles are documented and the geographic distribution of the species in these Islands is given. The historical contribution of the steamer Blake (1878-1879) in the Lesser Antilles is emphasized. All species inventoried during KARUBENTHOS 2015 were already reported in the western Atlantic but 34 of them are new records for the Lesser Antilles and 116 are reported for the first time from Guadeloupe Island. This preliminary inventory is estimated to include about 38% of the deep-sea Decapoda potentially occurring around Guadeloupe Island.

A new bathyal mysid of the family Petalophthalmidae (Crustacea: Mysida) from the Bismarck Sea (Western Tropical Pacific Ocean).

A new species of the genus Petalophthalmus (Crustacea: Mysida: Petalophthalmidae) is described, based on one specimen collected from the Bismarck Sea (Papua New Guinea, Western Pacific Ocean). This species can be distinguished from the other species of the genus Petalophthalmus by the globular cornea and the armature of the telson. This new species lives between 800 and 1065 m depth. A discussion on the geographic and bathymetric distribution and an identification key to world species of Petalophthalmus are provided.

Papuadocus blodiwai gen. nov., sp. nov. (Crustacea: Amphipoda: Maeridae), a new bathyal species associated with sunken wood in the Bismarck Sea (Papua New Guinea).

A new species belonging to a new genus of Maeridae, Papuadocus blodiwai gen. nov., sp. nov., is described from bathyal bottoms of the Bismarck Sea (Papua New Guinea). This genus/species can be distinguished from most other known maerids by right and left maxillas 1 with asymmetrical palps and by gnathopod 2 not sexually dimorphic. Its closest relative is the genus Bathyceradocus also characterized by asymmetrical maxillas 1, but differing by the presence of gill on coxae 7. These observations lead to the conclusion that the diagnosis of the family Maeridae has to be amended to receive both Bathyceradocus and Papuadocus genera. All the collected specimens lived in association with sunken wood, at 500-580 m depth.

Two new deep-sea stalked barnacles, Arcoscalpellum epeeum sp. nov. and Gymnoscalpellum indopacificum sp. nov., from the Coral Sea, with descriptions of the penis in Gymnoscalpellum dwarf males.

The present study describes a new species of Arcoscalpellum Hoek, 1907, and a new species of Gymnoscalpellum Newman & Ross, 1971, collected by deep-sea expeditions led by the Muséum national d'Histoire naturelle (Paris) in the Coral Sea off New Caledonia, Papua New Guinea (PNG), the Solomon Islands and Vanuatu. Arcoscalpellum epeeum sp. nov. differs from all described species of Arcoscalpellum by the presence of a long, sharp, sword-shaped carina, which extends beyond the apices of the terga by 1/3 to 1/4 of their length. The species is dioecious, with large females and dwarf males that are sac-like, lack shell plates and are housed in paired receptacles at the inner edges of the scutal plates. Arcoscalpellum epeeum sp. nov. was collected in the waters of New Caledonia and Vanuatu. Gymnoscalpellum indopacificum sp. nov. differs from the six currently described species of Gymnoscalpellum by having a very small inframedian latus and a branched upper latus. The species is dioecious, with large females and dwarf males, the latter composed of 4 shell plates and housed in paired receptacles at the inner edges of the scutal plates. The penis of the dwarf males of G. indopacificum sp. nov. is about 0.8 of the total length of the male and has five side branches extending out along its length. Gymnoscalpellum indopacificum sp. nov. is distributed in the waters of Papua New Guinea, the Solomon Islands and Vanuatu, and represents the first record of this genus in the Indo-Pacific region.

Integrative biology of Idas iwaotakii (Habe, 1958), a 'model species' associated with sunken organic substrates.

The giant bathymodioline mussels from vents have been studied as models to understand the adaptation of organisms to deep-sea chemosynthetic environments. These mussels are closely related to minute mussels associated to organic remains decaying on the deep-sea floor. Whereas biological data accumulate for the giant mussels, the small mussels remain poorly studied. Despite this lack of data for species living on organic remains it has been hypothesized that during evolution, contrary to their relatives from vents or seeps, they did not acquire highly specialized biological features. We aim at testing this hypothesis by providing new biological data for species associated with organic falls. Within Bathymodiolinae a close phylogenetic relationship was revealed between the Bathymodiolus sensu stricto lineage (i.e. "thermophilus" lineage) which includes exclusively vent and seep species, and a diversified lineage of small mussels, attributed to the genus Idas, that includes mostly species from organic falls. We selected Idas iwaotakii (Habe, 1958) from this latter lineage to analyse population structure and to document biological features. Mitochondrial and nuclear markers reveal a north-south genetic structure at an oceanic scale in the Western Pacific but no structure was revealed at a regional scale or as correlated with the kind of substrate or depth. The morphology of larval shells suggests substantial dispersal abilities. Nutritional features were assessed by examining bacterial diversity coupled by a microscopic analysis of the digestive tract. Molecular data demonstrated the presence of sulphur-oxidizing bacteria resembling those identified in other Bathymodiolinae. In contrast with most Bathymodiolus s.s. species the digestive tract of I. iwaotakii is not reduced. Combining data from literature with the present data shows that most of the important biological features are shared between Bathymodiolus s.s. species and its sister-lineage. However Bathymodiolus s.s. species are ecologically more restricted and also display a lower species richness than Idas species.

Inorganic carbon fixation by chemosynthetic ectosymbionts and nutritional transfers to the hydrothermal vent host-shrimp Rimicaris exoculata.

The shrimp Rimicaris exoculata dominates several hydrothermal vent ecosystems of the Mid-Atlantic Ridge and is thought to be a primary consumer harbouring a chemoautotrophic bacterial community in its gill chamber. The aim of the present study was to test current hypotheses concerning the epibiont's chemoautotrophy, and the mutualistic character of this association. In-vivo experiments were carried out in a pressurised aquarium with isotope-labelled inorganic carbon (NaH(13)CO(3) and NaH(14)CO(3)) in the presence of two different electron donors (Na(2)S(2)O(3) and Fe(2+)) and with radiolabelled organic compounds ((14)C-acetate and (3)H-lysine) chosen as potential bacterial substrates and/or metabolic by-products in experiments mimicking transfer of small biomolecules from epibionts to host. The bacterial epibionts were found to assimilate inorganic carbon by chemoautotrophy, but many of them (thick filaments of epsilonproteobacteria) appeared versatile and able to switch between electron donors, including organic compounds (heterotrophic acetate and lysine uptake). At least some of them (thin filamentous gammaproteobacteria) also seem capable of internal energy storage that could supply chemosynthetic metabolism for hours under conditions of electron donor deprivation. As direct nutritional transfer from bacteria to host was detected, the association appears as true mutualism. Import of soluble bacterial products occurs by permeation across the gill chamber integument, rather than via the digestive tract. This first demonstration of such capabilities in a decapod crustacean supports the previously discarded hypothesis of transtegumental absorption of dissolved organic matter or carbon as a common nutritional pathway.

New digestive symbiosis in the hydrothermal vent amphipoda Ventiella sulfuris.

Ventiella sulfuris Barnard and Ingram, 1990 is the most abundant amphipod species inhabiting the Eastern Pacific Rise (EPR 9°N) vent fields. This vent-endemic species is frequently encountered near colonies of Pompeii worms Alvinella pompejana. V. sulfuris specimens were collected during the oceanographic cruise LADDER II at the Bio9 (9°50.3'N, 2508m depth) hydrothermal vent site. Main objectives were to highlight the occurrence of bacterial symbiosis in V. sulfuris and to hypothesise their implications in nutrition. Observations in light and electron microscopy (SEM, TEM) showed that the outer body surface and appendages are free of microorganisms. In contrast, the digestive system revealed two major microbial communities settled in the midgut and in the hindgut. Gut contents showed bacterial traces together with abundant fragments of Alvinellid cuticle and setae, from A. pompejana, suggesting that V. sulfuris could directly feed on Alvinellids and/or on their bacterial epibionts. Molecular analyses based on the 16S rRNA genes revealed the diversity of bacterial communities in the digestive system, of which, the Epsilonproteobacteria phylum, could be considered as one of the major bacterial group. Hypotheses were proposed on their symbiotic features and their implications in V. sulfuris nutrition.

Oxygen as a driver of early arthropod micro-benthos evolution.

BACKGROUND: We examine the physiological and lifestyle adaptations which facilitated the emergence of ostracods as the numerically dominant Phanerozoic bivalve arthropod micro-benthos. METHODOLOGY/PRINCIPAL FINDINGS: The PO(2) of modern normoxic seawater is 21 kPa (air-equilibrated water), a level that would cause cellular damage if found in the tissues of ostracods and much other marine fauna. The PO(2) of most aquatic breathers at the cellular level is much lower, between 1 and 3 kPa. Ostracods avoid oxygen toxicity by migrating to waters which are hypoxic, or by developing metabolisms which generate high consumption of O(2). Interrogation of the Cambrian record of bivalve arthropod micro-benthos suggests a strong control on ecosystem evolution exerted by changing seawater O(2) levels. The PO(2) of air-equilibrated Cambrian-seawater is predicted to have varied between 10 and 30 kPa. Three groups of marine shelf-dwelling bivalve arthropods adopted different responses to Cambrian seawater O(2). Bradoriida evolved cardiovascular systems that favoured colonization of oxygenated marine waters. Their biodiversity declined during intervals associated with black shale deposition and marine shelf anoxia and their diversity may also have been curtailed by elevated late Cambrian (Furongian) oxygen-levels that increased the PO(2) gradient between seawater and bradoriid tissues. Phosphatocopida responded to Cambrian anoxia differently, reaching their peak during widespread seabed dysoxia of the SPICE event. They lacked a cardiovascular system and appear to have been adapted to seawater hypoxia. As latest Cambrian marine shelf waters became well oxygenated, phosphatocopids went extinct. Changing seawater oxygen-levels and the demise of much of the seabed bradoriid micro-benthos favoured a third group of arthropod micro-benthos, the ostracods. These animals adopted lifestyles that made them tolerant of changes in seawater O(2). Ostracods became the numerically dominant arthropod micro-benthos of the Phanerozoic. CONCLUSIONS/SIGNIFICANCE: Our work has implications from an evolutionary context for understanding how oxygen-level in marine ecosystems drives behaviour.

The early life history of tissue oxygenation in crustaceans: the strategy of the myodocopid ostracod Cylindroleberis mariae.

We studied basic principles of respiratory physiology in Cylindroleberididae, Cylindroleberis mariae Baird 1850, which are millimetre-sized crustaceans (myodocop ostracod) having a fossil record dating back to about 425 millions years ago. Facing experimental changes of O2 partial pressures in the range 2-40 kPa (normoxia is 21 kPa), C. mariae lack any regulatory mechanism to adapt their ventilatory and circulatory activity. Thus, the oxygenation status of their internal milieu must follow, as a dependent variable, the ambient oxygenation. Freely behaving C. mariae exhibit a marked diurnal activity rhythm. They are actively swimming in the water column during night, where they inspire in normoxic-normocapnic water. They are resting in self-made nests during daytime, where they are rebreathing in a confined and hypoxic environment. By analogy to extensive previous literature data, we suggest that these changes of respiratory gas content, and the associated tissue gas status, participate to the shaping of their metabolic activity and behaviour. To conclude, as Cylindroleberididae are early crustaceans exhibiting a remarkable stasis since the Palaeozoic, present data illustrates how principles of tissue oxygenation strategy can cover an impressive time scale.

How a low tissue O2 strategy could be conserved in early crustaceans: the example of the podocopid ostracods.

An adaptation strategy whereby O(2) partial pressure, P(O(2)), in the tissues is maintained within a low, narrow range of 1-3 kPa, largely independent of the inspired P(O(2)), has been reported in water- and air-breathing poikilotherms and in homeotherms. Based on the postulate that this basic cellular mechanism has been established since the early stages of evolution, it has been hypothesized that it could be the consequence of an early adaptation strategy to maintain cellular oxygenation within the same low and primitive range. To test this hypothesis we studied the basic mechanisms of oxygen regulation in podocopid ostracods, minute crustaceans that have existed on earth for at least 500 million years. Podocopids lack any regulatory mechanism for adapting their ventilation to cope with changes in water oxygenation, and instead adjust their tissue oxygenation status by migrating through the O(2) gradient to sediment layers where the P(O(2)) of the water is 3-5 kPa. Experimental manipulation of the O(2) profile induced their vertical migration to follow this precise water P(O(2)) and demonstrates the existence of a regulation strategy. This strategy must be associated with the lower P(O(2)) values within the animal's carapace valves, showing that podocopids can actively regulate their tissue P(O(2)) at constant but even lower values than the water. In conclusion, the low tissue P(O(2)) strategy could have existed in early crustaceans and, by extension, in early animals.