Species delimitation in poorly known and diverse taxa is usually performed based on monolocus, DNA-­barcoding-­like approaches, while multilocus data are often used to test alternative species hypotheses in well-­studied groups. We combined both approaches to delimit species in the Xenuroturris/Iotyrris complex, a group of venomous marine gastropods from the Indo-P­ acific. First, COI sequences were analysed using three methods of species delimitation to propose primary species hypotheses. Second, RAD sequencing data were also obtained and a maximum-l­ikelihood phylogenetic tree produced. We tested the impact of the level of missing data on the robustness of the phylogenetic tree obtained with the RAD-­seq data. Alternative species partitions revealed with the COI data set were also tested using the RAD-­seq data and the Bayes factor species delimitation method. The congruence between the species hypotheses proposed with the mitochondrial nuclear data sets, together with the morphological variability of the shell and the radula and the distribution pattern, was used to turn the primary species hypotheses into secondary species hypotheses. Allopatric primary species hypotheses defined with the COI gene were interpreted to correspond to intraspecific structure. Most of the species are found sympatrically in the Philippines, and only one is confidently identified as a new species and described as Iotyrris conotaxis n. sp. The results obtained demonstrate the efficiency of the combined monolocus/multilocus approach to delimit species.

ATIMO VATAE, INHACA 2011, MAINBAZA, Restricted, PAPUA NIUGINI, SANTO 2006, TERRASSES

Transcriptome-based exon capture methods provide an approach to recover several hundred markers from genomic DNA, allowing for robust phylogenetic estimation at deep timescales. We applied this method to a highly diverse group of venomous marine snails, Conoidea, for which published phylogenetic trees remain mostly unresolved for the deeper nodes. We targeted 850 protein coding genes (678,322 bp) in ca. 120 samples, spanning all (except one) known families of Conoidea and a broad selection of non-Conoidea neogastropods. The capture was successful for most samples, although capture efficiency decreased when DNA libraries were of insufficient quality and/or quantity (dried samples or low starting DNA concentration) and when targeting the most divergent lineages. An average of 75.4% of proteins was recovered, and the resulting tree, reconstructed using both supermatrix (IQ-tree) and supertree (Astral-II, combined with the Weighted Statistical Binning method) approaches, are almost fully supported. A reconstructed fossil-calibrated tree dates the origin of Conoidea to the Lower Cretaceous. We provide descriptions for two new families. The phylogeny revealed in this study provides a robust framework to reinterpret changes in Conoidea anatomy through time. Finally, we used the phylogeny to test the impact of the venom gland and radular type on diversification rates. Our analyses revealed that repeated losses of the venom gland had no effect on diversification rates, while families with a breadth of radula types showed increases in diversification rates, thus suggesting that trophic ecology may have an impact on the evolution of Conoidea.

ATIMO VATAE, AURORA 2007, BIOPAPUA, CEAMARC-AA, CONCALIS, Restricted, DongSha 2014, EXBODI, GUYANE 2014, ILES DU SALUT, INHACA 2011, KARUBENTHOS 2012, KAVIENG 2014, MAINBAZA, NORFOLK 2, NanHai 2014, PANGLAO 2005, PAPUA NIUGINI, Restricted, SALOMONBOA 3, TAIWAN 2013, TERRASSES, Restricted

BIOPAPUA

With about 800 Recent species, ‘miters’ are a widely distributed group of tropical and subtropical gastropods that are most diverse in the Indo-West Pacific. They include the two families Mitridae and Costellariidae, similar in shell morphology and traditionally treated as close relatives. Some genera of deep-water Ptychatractidae and Volutomitridae are close to miters in shell morphology, and the term ‘mitriform gastropods’ has been introduced to refer to Mitridae, Costellariidae, and this assortment of convergent forms. The present study aimed at the reconstruction of phylogenetic relationships of mitriform gastropods based on representative taxon sampling. Four genetic markers [cytochrome c oxidase subunit I (COI), 16S and 12S rRNA mitochondrial genes, and H3 (Histone 3) nuclear gene] were sequenced for over 90 species in 20 genera, and the molecular data set was supplemented by studies of radula morphology. Our analysis recovered Mitridae as a monophyletic group, whereas the genus Mitra was found to be polyphyletic. Of 42 mitrid species included in the analysis, 37 formed a well-supported ‘core Mitridae’ consisting of four major clades, three of them consistent with the subfamilies Cylindromitrinae, Imbricariinae, and Mitrinae, and Strigatella paupercula standing out by itself. Basal to the ‘core Mitridae’ are four minor lineages, with the genus Charitodoron recognized as sister group to all other Mitridae. The deepwater family Pyramimitridae shows a sister relationship to the Mitridae, with high support for a Pyramimitridae + Mitridae clade. Our results recover the monophyly of the Costellariidae, which form a wellsupported clade that also includes Ptychatractidae, Columbariinae, and Volutomitridae, but not Mitridae. Most derived and diverse amongst Costellariidae are species of Vexillum, characterized by a bow-shaped, multicuspidate rachidian tooth. Several previously unrecognized deep-water costellariid lineages are revealed. Their members retain some plesiomorphies – in particular a tricuspidate rachidian tooth – that makes them morphologically intermediate between ptychatractids and Vexillum. The taxa of Ptychatractidae included in the analysis are not monophyletic, but form three well-supported, unrelated groupings, corresponding respectively to Ceratoxancus + Latiromitra, Exilia, and Exiliodea. None of them shows an affinity to Pseudolividae.

ATIMO VATAE, AURORA 2007, BIOPAPUA, CONCALIS, EBISCO, EXBODI, INHACA 2011, MAINBAZA, MIRIKY, NORFOLK 2, PANGLAO 2004, PANGLAO 2005, PAPUA NIUGINI, Restricted, SALOMON 2, SALOMONBOA 3, SANTO 2006, TARASOC, TERRASSES, Tuhaa Pae 2013, Restricted

Alongside confirmation of the monophyly of the gastropod family Mitridae, a recent molecular phylogenetic analysis disclosed multiple inconsistencies with the existing taxonomic framework. In the present study, we expanded the molecular sampling to 103 species, representing 26% of the 402 extant species currently accepted in the family and 16 of the 19 currently accepted extant genera; 83 species were sequenced for four molecular markers [cytochrome c oxidase subunit I (COI), 16S and 12S rRNA, and H3 (Histone 3)]. Molecular analyses were supplemented by morphological studies, focused on characters of the radula and, in a more restricted data set, proboscis anatomy. These data form the basis for a revised classification of the Mitridae. A first dichotomy divides mitrids into two unequal clades, Charitodoron and the Mitridae s.s. Species of Charitodoron show profound differences to all other Mitridae in foregut anatomy (lacking an epiproboscis) and shell morphology (smooth columella, bulbous protoconch of non-planktotrophic type), which leads to the erection of the separate family Charitodoronidae fam. nov. Three traditional subfamilies (Mitrinae, Cylindromitrinae and Imbricariinae) correspond to three of the inferred phylogenetic lineages of Mitridae s.s.; we redefine their contents, reinstate Strigatellinae Troschel, 1869 as valid and establish the new subfamily Isarinae. In the absence of molecular material, a sixth subfamily, Pleioptygmatinae, is included in Mitridae based on morphological considerations only. To resolve the polyphyly of Mitra and Cancilla in their current taxonomic extension, we reinstate the genera Episcomitra Monterosato, 1917, Isara H. & A. Adams, 1853 and Probata Sarasúa, 1989 and establish 11 new genera: Quasimitra, Roseomitra, Fusidomiporta, Profundimitra, Cancillopsis, Pseudonebularia, Gemmulimitra and Neotiara in Mitrinae; Imbricariopsis in Imbricariinae; Carinomitra and Condylomitra are left unassigned to a subfamily. Altogether 32 genera are recognized within the family. Their diversity and distribution are discussed, along with general trends in morphological evolution of the family.

ATIMO VATAE, AURORA 2007, BIOPAPUA, CONCALIS, EBISCO, EXBODI, GUYANE 2014, INHACA 2011, KARUBENTHOS 2, KARUBENTHOS 2012, KAVIENG 2014, MAINBAZA, MIRIKY, PANGLAO 2004, PANGLAO 2005, PAPUA NIUGINI, SALOMONBOA 3, SANTO 2006, TARASOC, Tuhaa Pae 2013

Marginelliform gastropods are a heterogeneous and diverse group of molluscs encompassing over 1,600 living species, among which are the smallest known neogastropods. The relationships of marginelliform gastropods within the order Neogastropoda are controversial, and the monophyly of the two marginelliform families the Marginellidae J. Fleming, 1828 and the Cystiscidae Stimpson, 1865, remains unconfirmed. DNA sequence data have never been used to assess the relationships of the marginelliform gastropods, making this group the only major branch missing in our current understanding of the neogastropod tree of life. Here we report results of the first multilocus phylogenetic analysis of marginelliform gastropods, which is based on a dataset comprising 63 species (20 genera) of Marginellidae and Cystiscidae, and a wide range of neogastropod lineages. The Marginellidae and Cystiscidae form a moderately supported clade that is sister to the family Volutidae. Marginellona gigas appears to be sister to all other marginelliforms. The subfamily Marginellinae was recovered as a well-supported clade, and good resolution of this part of the tree makes it possible to propose amendments to the family-level classification of the group. The relationship between Granulina and other marginelliforms could not be resolved and requires further study. Due to poor resolution of basal relationships within the Marginellidae–Cystiscidae clade, the monophyly of the Cystiscidae was neither confirmed nor convincingly rejected. The shell morphology of most marginellid and cystiscid genera is taxonomically not very informative but, nevertheless, of the traditionally recognized genera only Gibberula and Dentimargo were shown to be polyphyletic. Although a comprehensive systematic revision of the group requires more extensive taxonomic sampling (e.g. with better representation of the type species of nominal genus-group names), our results support the superfamily Volutoidea, comprising four families (Volutidae, Cystiscidae, Marginellidae and Marginellonidae), with the placement of the Granulinidae uncertain for the time being.

ATIMO VATAE, Restricted, DongSha 2014, EXBODI, GUYANE 2014, ILES DU SALUT, INHACA 2011, KANACONO, KARUBENTHOS 2, KAVIENG 2014, MADEEP, MADIBENTHOS, MAINBAZA, PAPUA NIUGINI, Restricted

LIFOU 2000, SANTO 2006

A new genus and species of Fasciolariidae, Angulofusus nedae n. gen. n. sp. conchologically superficially resembling some Conoidea is described. Radula, anatomy and coloration of the body of the new species suggest that it belongs to subfamily Fusininae, being very similar in anatomy to species of the genus Amiantofusus. The species is broadly distributed in the Indo-Pacific, from the Philippines, through Vanuatu and New Caledonia to Wallis and Futuna in 40 to 105 m.

BATHUS 1, LAGON, MUSORSTOM 10, MUSORSTOM 7, PANGLAO 2004, SANTO 2006

Conoidean gastropods of the genera Kermia, Oliver, and Pseudodaphnella Boettger, (Raphitomidae) are common in shallow-water habitats of the tropical Indo-Pacific. They form a distinct morphologically homogeneous complex, easily recognizable by sculpture and colour pattern, encompassing around 80 described species. Examination of a vast material accumulated during recent expeditions in various regions of the Indo-Pacific revealed a number of undescribed species of this complex. Our material included 32 morphospecies available for molecular phylogenetic study; phylogenetic reconstruction based on the COI gene confirmed the species hypotheses based on morphological characters. A total of 18 terminal taxa were attributed to known species and 14 were identified as new species. Of these, 12 species, for which sufficient material was available, are described. Phylogenetic analysis indicated close relationships of the genera Kermia and Pseudodaphnella with members of some other conoidean genera (specifically Exomilus Hedley, , Paramontana Laseron, and Thetidos Hedley, ) and taxonomic implications of the data obtained are discussed. To test the taxonomic value of protoconch and review its wide use in classification of Conoidea, the evolution of the protoconch morphology was reconstructed using a phylogenetic tree. It has revealed that protoconchs of different types may appear in closely related species, sometimes hardly distinguishable by teleoconch morphology. A switch from planctotrophic to non-planctotrophic mode of development occurred at least four times in the evolutionary history of the Kermia Pseudodaphnella complex, indicating high developmental plasticity of the group. Its role in radiation of the Kermia Pseudodaphnella complex and applications for use of protoconch morphology in the classification of Conoidea are discussed.

ATIMO VATAE, EBISCO, LIFOU 2000, MONTROUZIER, PANGLAO 2004, PANGLAO 2005, SALOMONBOA 3, SANTO 2006

The genus Casmaria H. Adams & A. Adams, 1853 (family Cassidae) is widespread in the tropical Indo-Pacific and has been documented from some Atlantic localities as well. Two Casmaria species, C. erinaceus (Linnaeus, 1758) and C. ponderosa (Gmelin, 1791), are common in Indo-Pacific shallow-water sandy bottom communities and are characterized by high morphological variability; both species encompass multiple, often sympatric forms of uncertain status. In the present study we carry out a phylogenetic analysis of some Philippine Casmaria morphs and demonstrate that one of the distinctive morphs earlier assigned to Casmaria ponderosa is in fact a different species, which we describe as Casmaria boblehmani sp. nov. The smooth form of Casmaria ponderosa, C. ponderosa ponderosa, and the solid nodulose form, widely called “form nodulosa” despite being strikingly different in shell morphology, are shown to be conspecific. Studied specimens of these two morphs even from different localities share the same haplotype of the CO1 gene. In light of these new data on the morphological variability of Casmaria species, we discuss criteria of species delimitation in the genus Casmaria and possible affinities of Casmaria boblehmani sp. nov. within the genus.

PANGLAO 2004, SANTO 2006

LIFOU 2000, MONTROUZIER, PANGLAO 2004, PAPUA NIUGINI, RAPA 2002, SANTO 2006

KAVIENG 2014, LAGON, LIFOU 2000, MONTROUZIER, MUSORSTOM 10, PANGLAO 2004, PAPUA NIUGINI, SANTO 2006

The superfamily Olivoidea is broadly distributed in the world’s oceans mostly in coastal waters at tropical and subtropical latitudes. It encompasses around 30 Recent genera and 460 species. Two families – Olividae and Olivellidae – are classically recognized within the superfamily. Their shell is very characteristic due to the presence of a modified callused anterior end and a fasciole. Prior to the present work, neither the monophyly of the superfamily nor the relationships among its genera had been tested with molecular phylogenetics. Four genetic markers [cytochrome c oxidase subunit I (COI), 16S and 12S rRNA mitochondrial genes, and Histone 3 (H3) nuclear gene] were sequenced for 42 species in 14 genera. Additionally, 18 species were sequenced for COI only. The molecular dataset was supplemented by anatomical and radula data. Our analysis recovered, albeit with weak support, a monophyletic Olivoidea, which in turn includes with 100% support, in addition to traditional olivoideans, representatives of a paraphyletic Pseudolividae. The relationships between the former families and subfamilies are drastically revised and a new classification of the superfamily is here proposed, now including five families: Bellolividae fam. nov., Benthobiidae fam. nov., Olividae, Pseudolividae and Ancillariidae. Within Olividae four subfamilies are recognized, reflecting the high morphological disparity within the family: Olivinae, Olivellinae, Agaroniinae and Calyptolivinae subfam. nov. All the recent genera are discussed and reclassified based on molecular phylogeny and/or morphology and anatomy. The homology of different features of the shells is established for the first time throughout the superfamily, and a refined terminology is proposed. Based on a correlation between anatomical characteristics and shell features and observations of live animals, we make hypotheses on which part of the mantle is responsible for depositing which callused feature of the shell. Our results demonstrate that morphological data alone should be used with caution for phylogenetic reconstructions. For instance, the radula – that is otherwise considered to be of fundamental importance in the taxonomy of Neogastropoda – is extremely variable within the single family Olividae, with a range of variation larger than within the rest of the entire superfamily. In the refined classification, Pseudolividae are nested within Olivoidea, which is partially returning to ‘the roots’, that is to the classification of Thiele (1929).

ATIMO VATAE, AURORA 2007, BIOPAPUA, CONCALIS, Restricted, EBISCO, INHACA 2011, KARUBENTHOS 2012, KAVIENG 2014, MAINBAZA, MIRIKY, NORFOLK 2, PANGLAO 2004, PANGLAO 2005, PAPUA NIUGINI, Restricted, SALOMON 2, SALOMONBOA 3, SANTO 2006, TARASOC, TERRASSES

Buccinidae—like other canivorous and predatory molluscs—are generally considered to be occasional visitors or rare colonizers in deep-sea biogenic habitats. However, casual observations during tropical deep-sea cruises suggest that associations between buccinids and sunken wood, in particular, are not fortuitous. Enigmatocolus monnieri has been found to co-occur in Madagascar with bathymodiolines, vesicomyids and solemyids, indicating the presence of seeps, and species of Thermosipho gen. Nov. Have been sampled by submersibles and remotely operated vehicles, exclusively from hydrothermal vents. A molecular phylogeny (based on CO1, 12S and 28S genes) reveals that buccinid genera potentially associated with sunken wood (Eosipho, Gaillea gen. Nov., Calagrassor gen. Nov., and Manaria) are closely related to taxa from vents (Thermosipho gen. Nov.) and seeps (Enigmaticolus). The anatomy of several dissected species did not reveal any special trait that could be interpreted as a special adaptation to biogenic substrates. Buccinids from sunken wood are most diverse in the Indo-Pacific centre of marine biodiversity, the ‘Coral Triangle’, at depths between 100 and 1000 m, with numerous species still undescribed.

AURORA 2007, MIRIKY, MUSORSTOM 2, PANGLAO 2005, SALOMON 2, SANTO 2006

Five new species of Olivoidea are described based on molecular and morphological evidence: four shallow subtidal Ancilla from Madagascar and Papua New Guinea, and one deep water (500-600 m) Calyptoliva from the Tuamotus. The sympatric – but not syntopic - Ancilla morrisoni and A. kaviengensis, from New Ireland province, are morphologically cryptic, differing mostly in shell colour, but are molecularly distinct. The sympatric – and possibly syntopic – Ancilla atimovatae and A. lhaumeti, belong to a species flock from southernmost Madagascar; A. atimovatae is conchologically nearly indistinguishable from A. ventricosa, but differs markedly in radular morphology. Calyptoliva was previously known only from the Coral Sea; C. bbugae is the first representative of the genus to yield molecular data. The new Ancilla are described based on sequenced holotypes; the type material of the new Calyptoliva includes a sequenced paratype.

ATIMO VATAE, CONCALIS, EXBODI, KAVIENG 2014, MAINBAZA, MIRIKY, PANGLAO 2005, PAPUA NIUGINI, TARASOC

The genus Pseudolatirus Bellardi, 1884, with the Miocene type species Fusus bilineatus Hörnes, 1853, has been used for 13 Miocene to Early Pleistocene fossil species and eight Recent species and has traditionally been placed in the fasciolariid subfamily Peristerniinae Tryon, 1880. Although the fossil species are apparently peristerniines, the Recent species were in their majority suspected to be most closely related to Granulifusus Kuroda & Habe, 1954 in the subfamily Fusininae Wrigley, 1927. Their close affinity was confirmed by the molecular phylogenetic analysis of Couto et al. (2016). In the molecular phylogenetic section we present a more detailed analysis of the relationships of 10 Recent Pseudolatirus-like species, erect two new fusinine genera, Okutanius gen. nov. (type species Fusolatirus kuroseanus Okutani, 1975) and Vermeijius gen. nov. (type species Pseudolatirus pallidus Kuroda & Habe, 1961). Five species are described as new for science, three of them are based on sequenced specimens (Granulifusus annae sp. nov., G. norfolkensis sp. nov., Okutanius ellenae gen. et sp. nov.) and two (G. tatianae sp. nov., G. guidoi sp. nov.) are attributed to Granulifusus on the basis of conchological similarities to sequenced species. New data on radular morphology is presented for examined species.

ATIMO VATAE, AURORA 2007, CONCALIS, DongSha 2014, EBISCO, GUYANE 2014, KANACONO, KARUBENTHOS 2012, KAVIENG 2014, MADEEP, MIRIKY, NanHai 2014, Restricted, PANGLAO 2004, PANGLAO 2005, PAPUA NIUGINI, SALOMON 2, SANTO 2006, TARASOC, TERRASSES

Species of the genus Clavus of the conoidean family Drilliidae that occur in the littoral and shallow waters of New Caledonia are here revised. This study is based primarily on recent expedition material from the Institut de Recherche pour le Développement (New Caledonia) and Muséum National d’Histoire Naturelle (France). A total of 22 species is recorded, of which eight are described as new. New species: Clavus boucheti, Clavus delphineae, Clavus virginieae, Clavus picoides, Clavus squamiferus, Clavus devexistriatus, Clavus hylikos, Clavus maestratii; New synonyms: Tylotiella Habe, 1958 = Clavus; Clavus leforestieri Hervier, 1896 = Pleurotoma obliquicostata Reeve, 1845; Pleurotoma mariei Crosse, 1869 = Pleurotoma lamberti Montrouzier, 1860; Clavus mighelsi Kay, 1979, new name for Pleurotoma acuminata Mighels, 1845, non J. Sowerby, 1816, was misidentified by Kay 1979; the lectotype of P. acuminata Mighels, 1845, is mangeliine. Clavus mighelsi sensu Kay 1979, is a synonym of Pleurotoma humilis E. A. Smith, 1879. It is suggested that Pleurotoma pulchella Reeve, 1845, sometimes treated as an Indo-Pacific species, may be a senior synonym of Fenimorea halidorema Schwengel, 1940, from the tropical western Atlantic. Nomen dubium: Pleurotoma mediocris Deshayes, 1863.

CHALCAL 1, CORAIL 2, LAGON, MONTROUZIER, MUSORSTOM 5, MUSORSTOM 6, PALEO-SURPRISE, PANGLAO 2004, Restricted, SANTO 2006

AURORA 2007, INHACA 2011, KAVIENG 2014, PANGLAO 2004, PAPUA NIUGINI, SANTO 2006