Introduction Host-parasite associations provide very useful models to study adaptive processes. We investigated the interaction between carnivorous marine gastropods, the Ovulidae or egg-cowries, and their cnidarian food targets. Ovulidae (Fleming, 1828), is a family of specialized carnivorous caenogastropods that feed by browsing on octocorals (Anthozoa: Octocorallia: Malacalcyonacea and Scleralcyonacea) or, to a much lesser degree, on antipatharians (Anthozoa: Hexacorallia: Antipatharia) and Stylasteridae (Hydrozoa: Hydroidolina: Anthoathecata). Very scanty information is available on the phylogenetic relationships and the degree of specificity of the relationship with the cnidarians of this corallivorous lineage, especially for deep-water taxa. Methods To assess taxonomic identifications and investigate cnidarian/ovulid relationships in the context of their evolution, we generated an extensive molecular dataset comprising two mitochondrial ( cox1 and 16S rDNA ) and one nuclear gene ( 28S rDNA ) from 524 specimens collected worldwide. The coral hosts of the ovulid species have been identified by integrating literature data with new records, employing morphological and/or molecular (the mitochondrial 16S rDNA and mtMSH , and the nuclear ITS2) markers. Results We obtained a molecular phylogenetic framework for the Ovulidae, time-calibrated with nine reliable fossil records. An ancestral state reconstruction allowed to identify Hexacorallia or Hydroidolina as the most likely ancestral cnidarian host for the Ovulidae. Discussion Our phylogenetic hypothesis revealed the existence of groups that do not completely correspond to the currently employed subfamilial arrangement. Concerning trophic ecology, while only pediculariines ( Pedicularia and allied) are associated with hydrozoans (Stylasteridae), our results suggest that some ovulid lineages shifted independently between octocorals and hexacorals.

ATIMO VATAE, AURORA 2007, BIOMAGLO, CORSICABENTHOS 1, CORSICABENTHOS 2, CORSICABENTHOS 3, EBISCO, EXBODI, KANACONO, KANADEEP, KARUBENTHOS 2, KAVIENG 2014, KOUMAC 2.3, MADEEP, MADIBENTHOS, MAINBAZA, MIRIKY, NORFOLK 2, NanHai 2014, PAKAIHI I TE MOANA, PANGLAO 2004, PAPUA NIUGINI, SALOMONBOA 3, SANTO 2006, SPANBIOS, ZhongSha 2015

Solenogastres and Caudofoveata (Aplacophora) remain some of the least known molluscs, despite ubiquity in the marine environment and importance in understanding molluscan evolution. The use of new morphological techniques and development of DNA barcode libraries have helped make specimen identification easier. However, for solenogasters, using histology for identification and adequate description of species remains necessary in most cases. This, together with the facts that knowledge about solenogaster species distributions is biased and that most species were described from one or very few individuals, explains why many open questions about the actual distribution, intra- and interspecific variability, etc., remain. We performed an integrative taxonomic study of eight specimens of solenogasters from the South China Sea (West Pacific Ocean) thatresulted in the identification of four new species of Proneomeniidae. Species identification and description following the established diagnostic characters were straightforward. However, phylogenetic analysis of molecular data obtained from these specimens and other members of Proneomeniidae indicate that the family is polyphyletic. We recovered representatives of two other families, Epimeniidae (Epimenia) and Strophomeniidae (Anamenia), nested within Proneomeniidae with strong support. Ancestral character state reconstruction indicates that characters commonly used in solenogaster taxonomy, such as the radula and foregut glands, may be more evolutionarily labile in this group than previously known. Therefore our work fills knowledge gaps regarding the diversity and distribution of members of this family but raises important questions about solenogaster taxonomy and systematics that should be further assessed with additional markers and broader taxon sampling.

CORSICABENTHOS 3, DongSha 2014, ZhongSha 2015

Species in the family Capulidae (Littorinimorpha: Capuloidea) display a wide range of shell morphologies. Several species are known to live in association with other benthic invertebrates—mostly bivalves and sabellid worms, but also other gastropods—and are believed to be kleptoparasitic filter feeders that take advantage of the water current produced by the host. This peculiar trophic ecology, implying a sedentary lifestyle, has resulted in highly convergent shell forms. This is particularly true for the genus Hyalorisia Dall, 1889, which occurs in deep water in the Caribbean and Indo-West Pacific provinces, with two nominal species recognized so far. Combining morphological, ecological and molecular data, we assessed the diversity of the genus, its phylogenetic position inside the family and its association with its bivalve host, the genus Propeamussium de Gregorio, 1884 (Pectinoidea), resulting in the description of nine new cryptic species. When sympatric, species of Hyalorisia are associated with different host species, but the same species of Propeamussium may be the host of several allopatric species of Hyalorisia.

AURORA 2007, CONCALIS, CORSICABENTHOS 1, EBISCO, KANACONO, KANADEEP, KARUBENTHOS 2, KAVIENG 2014, KOUMAC 2.3, MADEEP, MAINBAZA, MIRIKY, NanHai 2014, PANGLAO 2004, PANGLAO 2005, SALOMON 2, ZhongSha 2015

Abstract The systematics of the marine mollusc family Velutinidae has long been neglected by taxonomists, mainly because their often internal and fragile shells offer no morphological characters. Velutinids are usually undersampled owing to their cryptic mantle coloration on the solitary, social or colonial ascidians on which they feed and lay eggs. In this study, we address the worldwide diversity and phylogeny of Velutinidae based on the largest molecular dataset (313 specimens) to date, accounting for > 50% of the currently accepted genera, coupled with morphological and ecological data. Velutinids emerge as a diverse group, encompassing four independent subfamily-level lineages, two of which are newly described herein: Marseniopsinae subfam. nov. and Hainotinae subfam. nov. High diversity was found at genus and species levels, with two newly described genera (Variolipallium gen. nov. and Pacifica gen. nov.) and ≥ 86 species in the assayed dataset, 58 of which are new to science (67%). Velutinidae show a remarkable morphological plasticity in shell morphology, mantle extension and chromatic patterns. This variability is likely to be the result of different selective forces, including habitat, depth and trophic interactions.

ATIMO VATAE, BIOMAGLO, BIOPAPUA, CEAMARC-AA, CORSICABENTHOS 1, CORSICABENTHOS 2, CORSICABENTHOS 3, GUYANE 2014, ILES DU SALUT, KANACONO, KANADEEP 2, KARUBENTHOS 2, KAVIENG 2014, KOUMAC 2.1, KOUMAC 2.3, MADEEP, MADIBENTHOS, PANGLAO 2004, PAPUA NIUGINI, SAKIZAYA 2019, SANTO 2006, Tuhaa Pae 2013, ZhongSha 2015

Abstract The molluskan order Neogastropoda encompasses over 15,000 almost exclusively marine species playing important roles in benthic communities and in the economies of coastal countries. Neogastropoda underwent intensive cladogenesis in the early stages of diversification, generating a “bush” at the base of their evolutionary tree, which has been hard to resolve even with high throughput molecular data. In the present study to resolve the bush, we use a variety of phylogenetic inference methods and a comprehensive exon capture dataset of 1817 loci (79.6% data occupancy) comprising 112 taxa of 48 out of 60 Neogastropoda families. Our results show consistent topologies and high support in all analyses at (super)family level, supporting monophyly of Muricoidea, Mitroidea, Conoidea, and, with some reservations, Olivoidea and Buccinoidea. Volutoidea and Turbinelloidea as currently circumscribed are clearly paraphyletic. Despite our analyses consistently resolving most backbone nodes, 3 prove problematic: First, the uncertain placement of Cancellariidae, as the sister group to either a Ficoidea-Tonnoidea clade or to the rest of Neogastropoda, leaves monophyly of Neogastropoda unresolved. Second, relationships are contradictory at the base of the major “core Neogastropoda” grouping. Third, coalescence-based analyses reject monophyly of the Buccinoidea in relation to Vasidae. We analyzed phylogenetic signal of targeted loci in relation to potential biases, and we propose the most probable resolutions in the latter 2 recalcitrant nodes. The uncertain placement of Cancellariidae may be explained by orthology violations due to differential paralog loss shortly after the whole genome duplication, which should be resolved with a curated set of longer loci.

ATIMO VATAE, AURORA 2007, BIOMAGLO, BIOPAPUA, CORSICABENTHOS 1, DongSha 2014, EXBODI, ILES DU SALUT, INHACA 2011, KANACONO, KANADEEP, KARUBENTHOS 2, KAVIENG 2014, KOUMAC 2.1, KOUMAC 2.3, MADEEP, MADIBENTHOS, MAINBAZA, NanHai 2014, PAPUA NIUGINI, TAIWAN 2013, Tuhaa Pae 2013, ZhongSha 2015

The four recent species of the genus Leufroyia Monterosato, 1884 are revised based on an integrative taxonomy approach: L. leufroyi (Michaud, 1828), L. concinna (Scacchi, 1836), L. erronea Monterosato, 1884 and L. villaria (Pusateri & Giannuzzi-Savelli, 2008).

CORSICABENTHOS 1

AbstractTarget‐capture approaches have facilitated a rapid growth in the field of phylogenomics but few probe sets exist for molluscs, an exceptionally rich phylum with unparalleled ecological and morphological diversity. We designed and tested the first universal probe set using Phyluce to capture ultraconserved elements (UCEs) and exon loci from the Subclass Caenogastropoda ‐ one of six major lineages of gastropods. The probe set consists of 29,441 probes designed to target 1142 UCE loci and 1933 exon loci (3075 total). In silico analyses of our probe set yielded an average of 2110 loci from genomes and 1389 loci from transcriptomes of diverse caenogastropods, from which an average of 1669 and 849 loci were retained respectively after screening to remove those that matched multiple contigs. Phylogenetic analyses of the loci extracted from transcriptomes produced well‐supported trees very similar to those published based on transcriptomic analyses. Phylogenetic relationships estimated from loci extracted from genomes recover similar phylogenetic relationships, and indicate that the loci targeted with this probe set are informative for resolving deep phylogenetic relationships. An in vitro analysis of the probe set with the Epitoniidae, a diverse caenogastropod family of uncertain affinity and with poorly resolved evolutionary relationships, recovered a total of 2850 loci. Although preliminary, the analysis of loci captured by our probe set for a small number of epitoniid taxa produced a well‐resolved tree indicating that this probe set is also able to resolve relationships at shallower hierarchical scales. Together, the in silico and in vitro analyses indicate that target‐capture enrichment with this probe set is a useful tool for reconstructing phylogenetic relationships across taxonomic levels and evolutionary time scales.

ATIMO VATAE, CORSICABENTHOS 2, KANACONO, KANADEEP 2, KARUBENTHOS 2, KAVIENG 2014, KOUMAC 2.1, KOUMAC 2.3, MADEEP, MADIBENTHOS, SPANBIOS, ZhongSha 2015

Abstract The superfamily Stromboidea is a clade of morphologically distinctive gastropods which include the iconic Strombidae, or ‘true conchs’. In this study, we present the most taxonomically extensive phylogeny of the superfamily to date, using fossil calibrations to produce a chronogram and extant geographical distributions to reconstruct ancestral ranges. From these results, we confirm the monophyly of all stromboidean families; however, six genera are not monophyletic using current generic assignments (Strombidae: Lentigo , Canarium , Dolomena , Doxander ; Xenophoridae: Onustus , Xenophora ). Within Strombidae, analyses resolve an Indo‐West Pacific (IWP) clade sister to an East Pacific/Atlantic clade, together sister to a second, larger IWP clade. Our results also indicate two pulses of strombid diversification within the Miocene, and a Tethyan/IWP origin for Strombidae—both supported by the fossil record. However, conflicts between divergence time estimates and the fossil record warrant further exploration. Species delimitation analyses using the COI barcoding gene support several taxonomic changes. We synonymise Euprotomus aurora with Euprotomus bulla , Strombus alatus with Strombus pugilis , Dolomena abbotti with Dolomena labiosa , and Dolomena operosa with Dolomena vittata . We identified cryptic species complexes within Terebellum terebellum , Lambis lambis , “Canarium” wilsonorum , Dolomena turturella and Maculastrombus mutabilis . We reinstate Rimellopsis laurenti as a species (previously synonymised with R. powisii ) and recognise Harpago chiragra rugosus and Lambis truncata sowerbyi valid at the rank of species. Finally, we establish several new combinations to render Lentigo , Dolomena , and Canarium monophyletic: Lentigo thersites , Dolomena robusta , Dolomena epidromis , Dolomena turturella , Dolomena taeniata , Dolomena vanikorensis , D. vittata , “Canarium” wilsonorum , Hawaiistrombus scalariformis , Maculastrombus mutabilis , Maculastrombus microurceus .

ATIMO VATAE, AURORA 2007, CONCALIS, CORSICABENTHOS 3, Restricted, EXBODI, GUYANE 2014, ILES DU SALUT, INHACA 2011, KANACONO, KARUBENTHOS 2, KARUBENTHOS 2012, KAVIENG 2014, KOUMAC 2.1, MADEEP, MADIBENTHOS, MIRIKY, PANGLAO 2004, PAPUA NIUGINI, SANTO 2006, Tuhaa Pae 2013

Abstract The superfamily Buccinoidea is distributed across the oceans of the world from the Arctic Ocean to the Antarctic and from intertidal to abyssal depths. It encompasses 3351 recent species in 337 genera. The latest taxonomic account recognized eight full families. For the first time, the monophyly of the superfamily and the relationships among the families are tested with molecular data supplemented by anatomical and radula data. Five genetic markers were used: fragments of mitochondrial COI, 16S rRNA, 12S rRNA and nuclear Histone 3 (H3) and 28S rRNA genes (for 225 species of 117 genera). Our analysis recovered Buccinoidea monophyletic in Bayesian analyses. The relationships between the formerly recognized families and subfamilies are drastically revised and a new classification of the superfamily is here proposed, now including 20 taxa of family rank and 23 subfamilies. Five new families (Chauvetiidae, Dolicholatiridae, Eosiphonidae, Prodotiidae and Retimohniidae) and one subfamily of Nassariidae (Tomliniinae) are described. Austrosiphonidae and Tudiclidae are resurrected from synonymy and employed in a new taxonomical extension. All but 40 recent genera are reclassified. Our results demonstrate that anatomy is rather uniform within the superfamily. With exceptions, the rather uniform radular morphology alone does not allow the allocation of genera to a particular family without additional molecular data.

ATIMO VATAE, AURORA 2007, BIOPAPUA, BOA1, CEAMARC-AA, CHALCAL 2, CONCALIS, CORSICABENTHOS 1, Restricted, Restricted, DongSha 2014, EBISCO, GUYANE 2014, ILES DU SALUT, INHACA 2011, KANACONO, KARUBENTHOS 2, KARUBENTHOS 2012, KAVALAN 2018, KOUMAC 2.1, KOUMAC 2.3, MADIBENTHOS, MAINBAZA, MIRIKY, MUSORSTOM 4, Restricted, NORFOLK 2, NanHai 2014, PANGLAO 2004, PANGLAO 2005, PAPUA NIUGINI, Restricted, SALOMON 2, SALOMONBOA 3, SANTO 2006, TAIWAN 2000, TAIWAN 2004, TARASOC, TERRASSES, Tuhaa Pae 2013, Restricted, ZhongSha 2015

The exon capture approach allows for sequencing a large number of loci to reconstruct phylogenetic relationships at varying taxonomic levels. In order to efficiently recover the targeted loci, the probes designed to capture the exons need to be genetically sufficiently similar to bind to the DNA, with a proposed limit of 10% of divergence. However, this threshold has never been tested with a specific protocol. We have designed a set of probes to capture 1125 exons in the Neogastropoda (Mollusca, Gastropoda), processed with the same protocol from the field to the DNA sequencing to control for potential bias in DNA quantity and quality. We sequenced 30 different species, including 14 species of Neogastropoda and 16 species of Caenogastropoda non-­Neogastropoda. Each species includes five specimens, for a total of 150 specimens, and for four specimens among the 150, DNA extracts were aliquoted in four samples, sequenced separately, to estimate the intraspecific and intraspecimen variability in capture success. Our results confirm the impact of genetic distance on the success of exon capture with a negative linear correlation between the genetic distance and the number of exons captured for each sample. Consequently, designing new capture probes would allow for capturing exons in genetically more distant groups without the need to redesign a new set of exons.

CORSICABENTHOS 3

In this study, we delved into the interaction between corallivorous marine gastropods, the muricid Coralliophilinae Chenu, 1859, and their cnidarian food targets. Coralliophilinae is a subfamily of specialised corallivorous caenogastropods that feed by browsing on octocorals or hexacorals. Only sparse information is available on the phylogenetic relationships and the degree of specificity of the trophic relationships within this corallivorous lineage. To address these gaps, we generated the largest molecular dataset to date, comprising two mitochondrial (cox1 and 16S rDNA) and one nuclear gene (ITS2 rDNA) from 586 specimens collected worldwide. The coral hosts of coralliophilines were identified through an integrative approach, combining literature data with new records, employing morphological and/or molecular markers, and incorporating data from DNA barcoding of the snail stomach content. Our comprehensive approach unveiled the existence of numerous cryptic species in Coralliophilinae, while the phylogeny showed that most of the currently accepted genera are not monophyletic. The molecular dating confirmed the origin of the Coralliophilinae in Middle Eocene, with diversification of most lineages during the Miocene. Our results indicate that the subfamily’s ancestor evolved in shallow waters in association with Scleractinia. Through the evolutionary history of Coralliophilinae, multiple host shifts to other cnidarian orders were observed, not correlated with changes in the depth range. The results of diversification analyses within the subfamily further suggest that the association with the host has influenced the evolutionary patterns of Coralliophilinae, but not vice versa.

ATIMO VATAE, BIOMAGLO, CONCALIS, CORSICABENTHOS 2, CORSICABENTHOS 3, EBISCO, EXBODI, GUYANE 2014, INHACA 2011, KARUBENTHOS 2, KAVIENG 2014, LIFOU 2000, MADIBENTHOS, MAINBAZA, MIRIKY, NORFOLK 2, NanHai 2014, PAKAIHI I TE MOANA, PANGLAO 2004, PANGLAO 2005, PAPUA NIUGINI, SANTO 2006, TARASOC, TERRASSES, Tuhaa Pae 2013, Walters Shoal, ZhongSha 2015

Introduction Host-parasite associations provide very useful models to study adaptive processes. We investigated the interaction between carnivorous marine gastropods, the Ovulidae or egg-cowries, and their cnidarian food targets. Ovulidae (Fleming, 1828), is a family of specialized carnivorous caenogastropods that feed by browsing on octocorals (Anthozoa: Octocorallia: Malacalcyonacea and Scleralcyonacea) or, to a much lesser degree, on antipatharians (Anthozoa: Hexacorallia: Antipatharia) and Stylasteridae (Hydrozoa: Hydroidolina: Anthoathecata). Very scanty information is available on the phylogenetic relationships and the degree of specificity of the relationship with the cnidarians of this corallivorous lineage, especially for deep-water taxa. Methods To assess taxonomic identifications and investigate cnidarian/ovulid relationships in the context of their evolution, we generated an extensive molecular dataset comprising two mitochondrial ( cox1 and 16S rDNA ) and one nuclear gene ( 28S rDNA ) from 524 specimens collected worldwide. The coral hosts of the ovulid species have been identified by integrating literature data with new records, employing morphological and/or molecular (the mitochondrial 16S rDNA and mtMSH , and the nuclear ITS2) markers. Results We obtained a molecular phylogenetic framework for the Ovulidae, time-calibrated with nine reliable fossil records. An ancestral state reconstruction allowed to identify Hexacorallia or Hydroidolina as the most likely ancestral cnidarian host for the Ovulidae. Discussion Our phylogenetic hypothesis revealed the existence of groups that do not completely correspond to the currently employed subfamilial arrangement. Concerning trophic ecology, while only pediculariines ( Pedicularia and allied) are associated with hydrozoans (Stylasteridae), our results suggest that some ovulid lineages shifted independently between octocorals and hexacorals.

ATIMO VATAE, AURORA 2007, BIOMAGLO, CORSICABENTHOS 1, CORSICABENTHOS 2, CORSICABENTHOS 3, EBISCO, EXBODI, KANACONO, KANADEEP, KARUBENTHOS 2, KAVIENG 2014, KOUMAC 2.3, MADEEP, MADIBENTHOS, MAINBAZA, MIRIKY, NORFOLK 2, NanHai 2014, PAKAIHI I TE MOANA, PANGLAO 2004, PAPUA NIUGINI, SALOMONBOA 3, SANTO 2006, SPANBIOS, ZhongSha 2015

ATIMO VATAE, AURORA 2007, BIOMAGLO, BIOPAPUA, BOA1, CONCALIS, CORSICABENTHOS 1, DongSha 2014, EBISCO, EXBODI, GUYANE 2014, INHACA 2011, KANACONO, KANADEEP, KARUBENTHOS 2, KARUBENTHOS 2012, KAVIENG 2014, KOUMAC 2.1, KOUMAC 2.3, MADEEP, MADIBENTHOS, MAINBAZA, MIRIKY, MUSORSTOM 9, NORFOLK 2, NanHai 2014, PAKAIHI I TE MOANA, PANGLAO 2004, PANGLAO 2005, PAPUA NIUGINI, SALOMON 2, SALOMONBOA 3, SANTO 2006, TAIWAN 2004, TAIWAN 2013, TARASOC, TERRASSES, ZhongSha 2015

Studies of non-indigenous species (NIS) often tend to focus on medium and large-sized taxa with potential for remarkable ecological and/or economic impact, whereas the early detection of small invertebrates is often delayed due to taxonomic challenge, lack of consistent, standardised monitoring efforts and limited funding. This study represents the first records of the marine amphipod Laticorophium baconi (Shoemaker, 1934) in Morocco, Tunisia, Corsica (France), Italy, Greece, Egypt, Saudi Arabia and New Caledonia (France). Furthermore, it reports an expansion of its known distribution in Portugal (earliest and northernmost record for the country and first record for Macaronesia), Spain (earliest record for Atlantic and Mediterranean European waters), and Australia (first record for Indian Ocean). Recreational boating and commercial shipping, mainly through hull fouling and secondarily ballast waters, are proposed as vectors for introduction and secondary spread of L. baconi. The following traits, analysed during the present study, could contribute to its invasive potential: (i) quick and extensive spread of the species worldwide, (ii) high densities in marinas, harbours, hull fouling and other artificial habitats, including aquaculture facilities and floating debris, (iii) high ability for short-term colonisation of empty artificial niches, (iv) diet based on detritus suggesting an opportunistic feeding behaviour, and (v) population survival during seasonal fluctuations in different regions. Taxonomic expertise and scientific collaboration, based on multidisciplinary networks of experts, are crucial for the early detection, distribution updates, and risk assessment of small and overlooked stowaways in marine environments.

CORSICABENTHOS 1, CORSICABENTHOS 2, CORSICABENTHOS 3, KOUMAC 2.3