Yu I Kantor

Returning to the roots: Morphology, molecular phylogeny and classification of the Olivoidea (Gastropoda: Neogastropoda)

Keywords:

The superfamily Olivoidea is broadly distributed in the World Ocean mostly in coastal waters at tropical and subtropical latitudes. It encompasses around 30 Recent genera and 460 species. Two families -Olividae and Olivellidaeare 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 5 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, that is partially returning to "the roots". to the classification of Thiele (1929).

INTRODUCTION

The evolutionary history of the predatory/scavenger Neogastropoda is a classic example of a successful, post-Cretaceous radiation of marine snails. They are the most prolific group of marine predatory gastropods, including no less than 12,000 species (WoRMS, 2016). At present Neogastropoda are classified in 40 Recent families (most represented in the fossil record) and seven exclusively fossil families [START_REF] Bouchet | Classification and nomenclator of gastropod families[END_REF]; with updates). Their phylogeny is far from resolved despite several attempts including the use of molecular characters (e.g. [START_REF] Cunha | Neogastropod phylogenetic relationships based on entire mitochondrial genomes[END_REF][START_REF] Oliverio | Relationships of the haematophagous marine snail Colubraria (Rachiglossa, Colubrariidae), within the neogastropod phylogenetic framework[END_REF][START_REF] Zou | Additional gene data and increased sampling give new insights into the phylogenetic relationships of Neogastropoda, within the caenogastropod phylogenetic framework[END_REF][START_REF] Osca | Caenogastropod mitogenomics[END_REF]. Nevertheless, significant progress has recently been achieved for individual neogastropod groups, and this has allowed the construction of robust, molecular-based phylogenies of selected clades at different ranks, from superfamilies (Conoidea - [START_REF] Puillandre | The dragon tamed? A molecular phylogeny of the Conoidea (Gastropoda)[END_REF] to families (eg. Muricidae - [START_REF] Barco | A molecular phylogenetic framework for the Muricidae, a diverse family of carnivorous gastropods[END_REF]Belomitridae -Kantor et al., 2012) and for non-monophyletic groups ("mitriform" neogastropods [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF]. Two new neogastropod families have already been established as a result of molecular studies combined with morphological and anatomical data: the above mentioned Belomitridae Kantor, Puillandre, Rivasseau & Bouchet, 2012and Bouchetispiridae Kantor, Strong & Puillandre, 2012(Kantor et al., 2012). The family Pyramimitridae Cossmann, 1901, "lost" in synonymy, was resurrected as valid and its position clarified (Kantor et al., 2014;[START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF]. It should be stressed that in none of these cases morphological data were sufficient for clarifying taxonomic position and relationships.

One of the reasons is that neogastropods are characterized by both high shell characters homoplasy and relative "uniformity" of anatomical characters traditionally used in systematics.

Correspondingly, the transfer of genera from one family to another is a frequent occurence. For instance, the mitriform genera Latiromitra Locard, 1897, Ceratoxancus Kuroda, 1952, and Exilioidea Grant & Gale, 1931, previously classified in the family Ptychatractidae on the basis of morphological characters (Kantor & Bouchet, 1997;[START_REF] Bouchet | The anatomy and systematics of Latiromitra, a genus of tropical deep-water Ptychatractinae (Gastropoda: Turbinellidae)[END_REF], were transferred respectively to the families Costellariidae (Latiromitra, Ceratoxancus) and Volutomitridae (Exilioidea) on the basis of a multilocus molecular phylogeny [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF], Fedosov et al., 2016).

By contrast, gastropods of the superfamily Olivoidea Latreille, 1825 (sensu [START_REF] Bouchet | Classification and nomenclator of gastropod families[END_REF] -the olive shells, or olividsis one of the most easily recognizable groups of tropical and subtropical Neogastropoda, encompassing around 30 Recent genera and 460 accepted Recent species (WoRMS, 2016). Olivoidea mostly live on soft bottoms from the intertidal to the bathyal, to depths of 1855 m ("Amalda" sibuetae Kantor & Bouchet, 1999).

Olivoidea are most speciose in the tropical realm, with few species penetrating mid latitude waters. Both planktotrophic and lecithotrophic larval development strategies are known in the superfamily.

The peculiar bullet-shaped glossy and smooth shell, together with the characteristic headfoot morphology (broad crescent-shaped propodium and parapodia, laterally embracing the shell), allow for the unambiguous allocation of most species to the Olivoidea. Cases where a species from a different family was mistakenly described as an olive, or vice versa, are rare but exist, e.g., Plicoliva Petuch, 1979, originally established as a subgenus of Oliva, was later reallocated to the Volutidae [START_REF] Petuch | Atlas of the living olive shells of the world[END_REF]Bouchet, 1990) . Remarkably, no olivid has ever been described in another family of Neogastropoda.

Despite their distinctiveness, Olivoidea have had a rather complicated taxonomic history.

The contents and taxonomic structure of the superfamily have been subjected to numerous alterations, sometimes rather profound. According to [START_REF] Bouchet | Classification and nomenclator of gastropod families[END_REF] seven available family group names has been proposed within olivids (in chronological order): Olividae proper; Ancillariinae [START_REF] Swainson | A treatise on malacology; on the natural classification of shells and shellfish[END_REF](= Ancillinae H. & A. Adams, 1853, raised to family rank by [START_REF] Iredale | A reference list of the marine Mollusca of New South Wales. The Australian Museum[END_REF]; Olivellinae Troschel, 1869 (raised to family rank by [START_REF] Golikov | Problems of phylogeny and system of the prosobranchiate gastropods[END_REF]; Agaroniinae [START_REF] Olsson | Studies of the genus Olivella[END_REF]Olivancillariidae Golikov & Starobogatov 1975;and Olivoidea was first ranked as a superfamily by [START_REF] Golikov | Systematics of prosobranch Gastropoda[END_REF], who included Harpidae. The same authors later [START_REF] Golikov | Problems of phylogeny and system of the prosobranchiate gastropods[END_REF]) erected a separate suborder Olivelloidei for Olivellidae, but did not discuss the position of other groups of Olivoidea, and this cannot be deduced from their publication. Neither Olivancillariidae nor Olivelloidei were considered as valid taxa by subsequent authors.

One more taxon sometimes associated with Olividae is Pseudolivinae de Gregorio, 1880 (raised to family by [START_REF] Delpey | Gastéropodes marins. Paléontologie, stratigraphie[END_REF]= Zemiridae Iredale, 1916), which has had a complicated taxonomic history. This small group includes nine Recent genera and slightly more than 20 species (WoRMS, 2016). Pseudolivinae was initially established as a subfamily of Buccinidae following the earlier opinion of [START_REF] Sowerby | Descriptions of Tertiary fossil shells from South America[END_REF] who believed that Pseudoliva [START_REF] Swainson | A treatise on malacology; on the natural classification of shells and shellfish[END_REF] and Buccinidae were closely related. This viewpoint was accepted by a number of authors (for a detailed account of the history of Pseudolividae, see [START_REF] Vermeij | Generic revision of the neogastropod family Pseudolividae[END_REF]) and it was either treated as a subfamily of Buccinidae (e. g. [START_REF] Cossmann | Essais de paléoconchologie comparée[END_REF] or as a separate family, but still closely related to Buccinidae (e. g., [START_REF] Golikov | Systematics of prosobranch Gastropoda[END_REF]. In fact, when establishing Pseudoliva, [START_REF] Swainson | A treatise on malacology; on the natural classification of shells and shellfish[END_REF] had already suggested affinities between his new genus and some olivids. This opinion was followed by many authors, including [START_REF] Thiele | What can we learn from confusing Olivella columellaris and O. semistriata (Olivellidae, Gastropoda), two key species in panamic sandy beach ecosystems?[END_REF], Wenz (1943[START_REF] Wenz | Gastropoda. Teil 1, Allgemeiner Teil und Prosobranchia[END_REF][START_REF] Wenz | Gastropoda. Teil 1, Allgemeiner Teil und Prosobranchia[END_REF]), [START_REF] Ponder | The genus Zemira H. and A. Adams (Mollusca: Neogastropoda)[END_REF], Ponder and[START_REF] Ponder | Classification of the Caenogastropoda and Heterostropha -A list of the family-group names and higher taxa[END_REF]Kilburn (1989), and pseudolivids were treated by them as a subfamily of Olividae. The position of Pseudolividae was re-assessed by [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF], who erected a new monotypical suborder Pseudolivoidei based on morphological and anatomical data. Although this was not followed in subsequent publications, Pseudolividae together with Ptychatractidae Stimpson, 1865 was placed in the superfamily Pseudolivoidea in the latest working classification of the Gastropoda [START_REF] Bouchet | Classification and nomenclator of gastropod families[END_REF].

Besides the controversial position of Pseudolivinae (-idae), the placement in Olivoidea of the remaining groups has never been questioned. The state of the art was reflected in [START_REF] Bouchet | Classification and nomenclator of gastropod families[END_REF], who attributed two families to Olivoidea -Olivellidae and Olividae (with the subfamilies Olivinae, Ancillariinae and †Vanpalmeriinae, the latter including the single species Vanpalmeria africana Agedoke, 1977 from the Paleocene of Nigeria).

Olivoidea remain relatively poorly studied anatomically. Leaving aside radula descriptions, publications dealing with their anatomy include description of several species of Oliva, two species of Olivancillaria, two Amalda, two Olivella and several Belloliva [START_REF] Küttler | Die Anatomie von Oliva peruviana Lamarck[END_REF][START_REF] Marcus | Studies on "Olividae[END_REF][START_REF] Marcus | On the prosobranchs Ancilla dimidiata and Marginella fraterculus[END_REF][START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF]Kantor & Tursch, 2001a;Kantor & Bouchet, 2007). Even these rather fragmentary anatomical data demonstrate significant morphological and anatomical disparity within the group.

With regard to the anatomy of the Pseudolividae the data are available on Zemira H. Adams & A. Adams, 1853, two Pseudoliva, two Melapium, one Luizia and several Benthobia [START_REF] Ponder | The genus Zemira H. and A. Adams (Mollusca: Neogastropoda)[END_REF][START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF][START_REF] Simone | Revision of the genus Benthobia (Caenogastropoda, Pseudolividae)[END_REF][START_REF] Simone | Family Pseudolividae (Caenogastropoda, Muricoidea): a polyphyletic taxon[END_REF].

The feeding habits of Olivoidea are substantiated by some unique morphological and physiological adaptations and have attracted the attention of biologists. The large species (different species of Oliva and Agaronia) are predators and scavengers, preying on a wide range of invertebrates: polychaetes, crustaceans, bivalves and gastropods [START_REF] Tursch | Oliva shells. The genus Oliva and the species problem. L'Informatore Piceno[END_REF], sea cucumbers and echinoids [START_REF] Taylor | Diet of olives: Oliva tigridella Duclos, 1835 in Queensland[END_REF]. Prey is located and seized by large crescent-shaped propodium and rapidly transferred into a pouch formed by the folding of the posterior part of the metapodium (Kantor & Tursch, 2001b). At least partial digestion occurs inside the pouch and proteolytic ferments are produced by the epithelium of the sole (Kantor & Tursch, 2001b). Similar observations have been made for Agaronia propatula (Conrad, 1849) [START_REF] Rupert | Autotomy of the posterior foot in Agaronia (Caenogastropoda: Olividae) occurs in animals that are fully withdrawn into their shells[END_REF][START_REF] Cyrus | The behavioural and sensory ecology of Agaronia propatula (Caenogastropoda: Olividae), a swash-surfing predator on sandy beaches of the panamic faunal province[END_REF], and several short videos of Agaronia hunting its prey Olivella semistriata [START_REF] Gray | Molluscous animals, and their shells The zoology of Capt. Beechey's voyage[END_REF]) can be accessed online as supplementary data to above mentioned publications (doi: 10.1093/mollus/eyr019 and 10.1093/mollus/eys006 correspondingly). Agaronia is able to autotomize the posterior metapodium probably as a defense mechanism, but is not able to feed until the foot is totally regenerated.

For small olivoideans, limited data are available only for Olivella, which demonstrate a great disparity of both feeding behaviour and diet. For example, Olivella minuta (Link, 1807) feeds on the bivalve Donax hanleyanus Philippi, 1847, foraminiferans, different crustaceans and even scaphopods (Marcus & Marcus, 1959, as Olivella verreauxii). In the subgenus Olivella (Pachyoliva)which includes the two species O. columellaris (Sowerby, 1825) and O. semistriatathe snails use their foot appendages to deploy mucus nets between the large propodial and small metapodial appendages for suspension feeding in the backwash of sandy beaches [START_REF] Thiele | What can we learn from confusing Olivella columellaris and O. semistriata (Olivellidae, Gastropoda), two key species in panamic sandy beach ecosystems?[END_REF]. O. biplicata (Sowerby, 1825) feeds on Foraminifera [START_REF] Hickman | Foraminiferivory; selective ingestion of foraminifera and test alterations produced by the neogastropod Olivella[END_REF].

Up to now there is no molecular phylogeny of the Olivoidea. There are sequences for 10 species in GenBank (three Olivella, four Oliva, one each of Pseudoliva, Amalda, and Belloliva) and the taxonomy and relationships within the group are based on morphological data. The aim of the present work is to construct a robust molecular phylogeny of Olivoidea, review published data on the morphology and anatomy of the group and present original data, and revise the current classification on the basis of molecular and morphological data.

MATERIAL AND METHODS

TAXON SAMPLING

Material included in this paper was collected during several biodiversity surveys organized mainly by the Muséum National d'Histoire Naturelle (MNHN), Pro-Natura International (PNI) and the Institut de Recherche pour le Développement (IRD), as part of the Our Planet Reviewed and Tropical Deep-Sea Benthos programs (see [START_REF] Bouchet | A quarter-century of deep-sea malacological exploration in the South and West Pacific: Where do we stand? How far to go? In[END_REF] the BasExp database at http://expeditions.mnhn.fr/). In the field, living specimens were anaesthetized with MgCl2 or removed from their shells in a microwave oven [START_REF] Galindo | Using microwaves to prepare gastropods for DNA Barcoding[END_REF]. A piece of the head-foot was cut and preserved in 96-98% ethanol. Bodies were then separated from the shells, with shells kept dry to prevent deterioration by etching and carrying the same registration number as the corresponding body and tissue-clip in 96-98% ethanol. Most of the material (shell, tissue and DNA) is vouchered in MNHN (http://science.mnhn.fr), unless otherwise stated.

In total 124 specimens of Olividae and Pseudolividae (sensu [START_REF] Bouchet | Classification and nomenclator of gastropod families[END_REF], representing 60 species, were sequenced; additionally sequences of three species representing unique lineages absent in our material were taken from GenBank. Although we included representatives of as many genus-level taxa as possible, some genera recognized as valid are not covered in our dataset, e. g. Eburna Lamarck, 1801 andAnolacia Gray, 1857. Their taxonomic position will be discussed in the Taxonomy section. Several new species were discovered in the course of the project and they are described in a separate paper (Kantor et al., 2016).

A suite of 12 species from 12 other neogastropod families in the superfamilies Conoidea, Cancellarioidea, Buccinoidea, and Muricoidea was included in the analysis to ensure adequate representation of major evolutionary lineages of the Neogastropoda and likely detection of olivoidean gastropods relationships. Tonna galea (Caenogastropoda, Tonnoidea, Tonnidae) was used as a distal outgroup. Specimen data as well as repositories are presented in Table 1, and some shells of sequenced vouchers (or conspecific specimens) are illustrated in Figs 34.

PCR AMPLIFICATION AND DNA SEQUENCING Total genomic DNA was extracted from muscle tissue using NucleoSpinR 96 Tissues (Macherey-Nagel) and following the manufacturer's instructions. Fragments of the mitochondrial genes Cytochrome oxidase subunit I (COI), 16S rRNA and 12S rRNA as well as the nuclear gene histone 3 (H3) were sequenced (Table 1). All PCR reactions were performed following protocols described in detail earlier [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF]. All genes were sequenced for both directions to confirm accuracy of each sequence. The sequencing was performed by Eurofins. Chromatograms were edited using CodonCode Aligner version 3.7.1.1.

SEQUENCE ALIGNMENTS AND PHYLOGENETIC ANALYSIS

Sequences were aligned for each gene independently using MUSCLE [START_REF] Edgar | MUSCLE: multiple sequence alignment with high accuracy and high throughput[END_REF]. The accuracy of automatic alignments was confirmed by eye and where necessary edited using BioEdit version 7.0.9.0 [START_REF] Hall | BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT[END_REF]. No indels were detected in protein coding genes COI and H3.

To delimit species within the genus Ancilla, a first dataset including 36 COI sequences was of ABGD (Automatic Barcode Gap Discovery) [START_REF] Puillandre | ABGD, Automatic Barcode Gap Discovery for primary species delimitation[END_REF] was used to propose primary species hypotheses, with default paramters, and the simple-distance substitution model.

To reconstruct phylogenetic relationships of the entire Olivoidea, one specimen per species of Olivoidea was analyzed (Table 1). Best-fit substitution models were identified for each gene separately using ModelgeneratorV.85 (Keane et al., 2006) (Table 2). Each gene was first analyzed separately: they provided mostly unsupported trees, and will not be discussed further in this paper. Because no incongruence between each gene tree was detected, they were combined in a concatenated four-gene dataset. In both Maximum Likelihood (ML) and Bayesian analysis (BA), each codon position of the COI gene for Ancilla, Oliva and Olivoidea datasets and each gene (16, 12S, H3) for the Olivoidea dataset were considered as independent partitions, each following a GTR+G and a GTR+I+G model (for ML and BI, respectively). Best-scoring Maximum Likelihood (ML) trees were estimated using RAxML-HPC2 [START_REF] Stamatakis | RAxML-VI-HPC: maximum likelihoodbased phylogenetic analyses with thousands of taxa and mixed models[END_REF] on XSEDE, as implemented on the CIPRES Science Gateway v.3.1. Robustness of the nodes was assessed using 1,000 bootstraps. Bayesian Analyses (BAs) were performed running two parallel analyses in MrBayes 3.2.6 [START_REF] Huelsenbeck | MrBayes: Bayesian inference of phylogeny[END_REF] on XSEDE, as implemented on the CIPRES Science Gateway v.3.1. For the Ancilla and Oliva datasets, used to test the monophyly of the species hypotheses proposed by ABGD, as well as the single gene analyses of the Olivoidea dataset, each run consisted of six Markov chains and 10,000,000 generations with eight chains, five swaps, a sampling frequency of 1,000 and a temperature of 0.02. For the fourgene analyses of the Olivoidea dataset, the parameters were the same, except for the number of generations (50,000,000) and the sampling frequency (5,000). Convergence of each analysis was evaluated using Tracer1.4.1 [START_REF] Rambaut | Tracer v. 1.6[END_REF] to ensure that all ESS values exceeded 200.

Consensus tree were calculated after omitting the first 25% trees as burn-in.

ANATOMY

The anatomy of representatives of every genus confirmed as valid was examined, whenever possible from the sequenced specimens themselves. For some groups (Belloliva, Oliva, Pseudolividae) published data were available [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF]Kantor & Bouchet, 2007;Kantor & Tursch, 2001a). Special attention was paid to the mantle morphology, anterior foregut, and external stomach morphology. The radula was prepared for most species, following the standard protocol in Kantor & Puillandre (2012), and examined with scanning electron microscopes in MNHN and A.N.Severtzov Institute of Ecology and Evolution, Russian Academy of Sciences.

Abbreviations of depositories

NSMK -National

RESULTS

SPECIES DELIMITATION

ABGD delimited 11 and and between 23 and 26 species for the Ancilla and Oliva datasets correspondingly. In the phylogenetic tree of Ancilla (Fig. 12), 6 of the 11 ABGD groups are represented by several specimens, and they each correspond to a supported monophyletic group.

In the Oliva datasets, two partitions are proposed. In the first partitions (with the lowest number of groups), 10 groups out of the 23 are represented by several specimesn, and each of them correspond to a supported monophyletic group in the tree (Fig. 11). In the partition with the highest number of species, Oliva caroliniana is divided in two, but one of them is paraphyletic.

Furthermore, Oliva lacanientai is divided in four groups, three of them being represented by a single specimen each, the other corresponding to a non-supported clade (Fig. 11). In all cases, O. sericea and O. miniacea are grouped together.

PHYLOGENETIC ANALYSIS

The topologies originating from analyses of the concatenated dataset both from ML (Fig. 1) and BA (Fig. 2) are largely congruent and the supported clades were retrieved in both analyses.

Here we mainly discuss the topology of the tree obtained with BA (Fig. 2) and refer to ML bootstrap support values where applicable.

The analyzed ingroup taxa include the species and genera attributed to the superfamily Olivoidea by [START_REF] Bouchet | Classification and nomenclator of gastropod families[END_REF] -for convenience collectively referred to below as "oliviforms" -, as well as those attributed to the family Pseudolividae. Together they form a clade, although only weakly supported (posterior probability 0.81 in the BA tree and bootstrap support only 33 in the ML tree). A clade that includes Belloliva [START_REF] Peile | Some notes on radulae[END_REF]Olivellopsis Thiele, 1929 (the latter represented by 'Olivella (Janaoliva)' amoni [START_REF] Sterba | Olivella (Janaoliva) amoni, a new subgenus and species from the Bismark Archipelago and Indonesia (Mollusca: Gastropoda: Olivellidae)[END_REF], see below) is strongly supported molecularly (0.96 and 100, respectively for BA and ML analyses) and anatomically, and is sister-group to the rest of the ingroup.

The rest of the ingroup taxa form a clade supported by ML (84) and strongly supported by BA (0.99). Pseudolividae as currently construed is polyphyletic and includes two distant clades.

The first clade is formed by two species of Benthobia Dall, 1889 and Fusulculus crenatus [START_REF] Bouchet | Two new deep-water Pseudolividae (Neogastropoda) from the south-west Pacific[END_REF]. The second includes two species of Pseudoliva represented by P. ancilla (Hanley, 1859) and 'Fulmentum' sepimentum (Rang, 1832) [with regard to the generic position of this species, see below in the Taxonomy part]. Both clades are strongly supported, but the deeper nodes do not have significant support and their relationships to other ingroup taxa cannot be resolved. The name Pseudolividae can be applied to the second clade, which includes the representatives of the type genus. No name is available for the first clade.

The remaining oliviforms constitute two clades, well supported both in ML and BA, although relationships between them are not resolved. The first, referred to as Olividae on Figs 1, 2 and below, combines representatives of the genera Oliva [START_REF] Bruguière | Encyclopedie Methodique[END_REF], Agaronia Gray, 1839, Calyptoliva Kantor & Bouchet, 2007and Olivella Swainson, 1831. This clade encompasses two currently recognized families -Olividae and Olivellidae, and we retain for it the senior name Olividae Latreille, 1825. The species of Olivella and Agaronia (both monophyletic and well supported) cluster together in ML and BA, although this grouping is not supported (only by ML analysis bootstrap support = 81). The single species of Calyptoliva included in the analysis is sister to Olivella + Agaronia, albeit again without support. The remaining species in Olividae are representatives of Oliva and also form a highly supported clade.

The second large, well-supported, clade corresponds to the generally accepted subfamily Ancillariinae (formerly included in Olividae), and representatives of the five genera are included in the analysis. Entomoliva [START_REF] Bouchet | A new genus of Ancillinae from New Caledonia, with the description of two new species[END_REF] is represented by a single species, but each of the four remaining genera Ancilla, Turrancilla, Ancillina and Amalda corresponds to a highlysupported clade in both analyses. On the contrary, relationships between genera remain unresolved.

Conversion of the molecular tree into a classification is straightforward, taking into account the anatomical data, which are described and discussed below. All the terminal suprageneric taxa are highly supported both in Bayesian and ML analyses (the posterior probabilities are 1, while the bootstrap supports range from 81 for Ancillariinae to 96-100 for all the others).

Conchologically and anatomically, Belloliva and Olivellopsis are rather similar to Olividae, sharing a number of common characters, including the mantle morphology (which has a mantle filament, and correspondingly a filament channel is present on the shell), similar radular morphology and presence of the anterior shell band (the character that are simingly apomophic for Olivoidea). Conversely, members of Pseudolividae and Benthobiidae are notably different in shell morphology and anatomy, lacking the above mentioned characters. Thus the topology of our molecular trees contradicts the parsimonious scheme of relationships that could be suggested based on morphology.

The ranks of suprageneric taxa are always subjective, even if all the data are taken into consideration. Olivoidea is a remarkable example of this. Olivella is morphologically very distinct from other oliviforms, having a very unusual radula with five teeth per transverse row (while all other ingroup genera have only three), and lacking a gland and valve of Leiblein.

Nevertheless, the molecular analysis confidently places Olivella in the same clade as Agaronia and Oliva, whereas the morphologically more similar Belloliva forms a totally separate branch.

Correspondingly, we rank as family the clade combining Olivella, Agaronia, Oliva, and Calyptoliva and, to accentuate the high morphological disparity, we recognize within Olividae the four subfamilies Olivinae, Agaroniinae Olsson, 1956, Olivellinae Olsson, 1956, and Calyptolivinae subfam. nov. For consistency, we rank Ancillariidae Swainson, 1840, as a family.

MORPHOLOGY

Shell and anatomy are rather variable in Olivoidea and have previously -and in the current paper -been used for classification at different taxonomic levels. After giving consideration to the subject for many years, we offer hypotheses of homologies of the different shell structures and correlate them with the morphology of the mantle. A bold typeface is used for the terms accepted by us, and italics are used for terms proposed earlier that we regard as synonyms (see Table 3 for shell characters).

Shell

The shells of Olivoidea as defined in the current paper can be divided into two types. The first is found in Olividae, Ancillariidae and Bellolividae and is characterized by a glossy surface without periostracum and the presence of the so-called anterior band and fasciole on the anterior part of the last whorl (Fig. 3 A-C; F-L; 4). In Benthobiidae and Pseudolividae, the shell is normally covered with a thin to thick periostracum, and the anterior band is not pronounced (Fig. 3 D-E, M). While refraining from implying any taxonomic value to these terms, we will for convenience refer to these two shell types as the "oliviform" and "pseudoliviform" respectively, and describe them below.

The most complex shell morphology is probably found in species of Oliva (Olivinae). It was treated in detail by Tursch and Greifineder (2001), who suggested homologies with other groups of the superfamily (op. cit.: fig. 7.21). By giving closer attention to other representatives of the superfamily, we have had to reconsider Tursch & Greifeneder's scheme (which was focused on Oliva) and this leads us to a revised terminology based on revised suggested homologies. Because shell characters have commonly been used in olivoidean taxonomy, many terms have been proposed, usually based on a particular taxon.

The oliviform shell

The oliviform shell is glossy, with the spire, columella and parietal wall of aperture callused to a different degree; the modified anterior part of the last whorl and fasciole are also callused. The last whorl is separated in three main regions: the last whorl cloak, the anterior band and the plication plate. The last whorl cloak is the surface of the adapical part of the last whorl (Fig. 5 -CL), it is glossy, enameled and, except for a few Ancillariidae (e.g. Entomoliva), devoid of sculpture. Abapically from the cloak (or from the olivoid band, when present) there is a sharp border (Fig. 5, ) that delimits the callused band, which is called here the anterior band following [START_REF] Tursch | Oliva shells. The genus Oliva and the species problem. L'Informatore Piceno[END_REF] (Fig. 5 -AB). As was pointed out by these authors, this border is represented by a sharp step (which can be easily felt by the finger), although it is often referred to as a groove. In some oliviforms, there is also a spiral groove (Fig. 5, ) situated at some distance adapically from the rear edge (Fig. 5, ) of the anterior band. In most cases it is a narrow and deep groove (rarely it looks like a very low ridge) that delimits one more band, the surface of which is similar to the rest of the cloak, although it can differ in colour sometimes rather contrasted, as in, e. g., Amalda contusa (Reeve, 1864). The groove is often referred to as ancillid groove and the band delimited by it as ancillid band (Kilburn, 1977 and1981). To avoid reference to a particular taxon (in this case, the genus Ancilla), we suggest here the terms olivoid groove and olivoid band for the band of the cloak that is delimited by the groove. It is important to note that the surface of the olivoid band is not callused and is similar to the rest of the last whorl cloak. An olivoid groove is found in a number of unrelated olivoideans. It is present in Ancillariidae (in some Ancilla spp. and Turrancilla spp., and always present in Ancillina, Amalda and Eburna) and Agaroniinae. In species with a well pronounced olivoid groove there is usually a more or less developed labral denticle (Fig. 5, Ancillald), terminating the groove on the lip.

The anterior band of the Ancillariidae has been called the fasciolar band by Kilburn (1977 and1981). However, we avoid using this term, since the band extends adapically beyond the limit of the fasciole. This anterior band is callused, usually shagreened to some extent. It can be uniform and rather thin (e.g. in some Olivella and Belloliva), or it can be very thick, distinctly subdivided into an upper (Fig. 5 uab) and a lower anterior band (Fig. 5 lab), sometimes distinguished by a slight difference in colour (in some Oliva and Olivella), sometimes separated by an obtuse or very sharp step (in most Ancillariidae, except Ancilla). For the border between the upper and lower parts of the anterior band we introduce here the term fasciolar ridge (Fig. 5,). It was referred to as the posterior fasciolar groove in Amalda and Eburna (Kilburn, 1977;[START_REF] Voskuil | The recent species of the genus Eburna Lamarck, 1801[END_REF] and as the submedian line for some Olivella [START_REF] Olsson | Studies of the genus Olivella[END_REF], although it is not a groove. An important observation in recognizing the homology of the anterior band is that it abuts to the rest of the last whorl, i.e. the cloak.

The rear edge of the anterior band sometimes reaches the outer apertural lip (in, e. g., Turrancilla, Eburna, and Entomoliva -Fig. 6 C-D), or comes in very close proximity to the lip in Amalda (Fig. 6 B) but, in Oliva and Ancilla, this sharp step is smoothened and merged with the anterior notch callus and disappears at some distance from the lip (Fig. 6 A).

The anteriormost part of the shell is most highly raised above the surface of the other part of the posterior whorl and in most cases is off-white and enameled. It is referred to here as the plication plate (Fig. 5 -PPfollowing [START_REF] Tursch | Oliva shells. The genus Oliva and the species problem. L'Informatore Piceno[END_REF]= columella pillar of Kilburn, 1977and 1981). We do not follow Kilburn's terminology, since this structure is not really a columella (this is very convincingly demonstrated by the illustration of the partial section of the shell of Oliva (Tursch & Greifeneder, 2001: fig. 7.20) and it also extends significantly on the side of the shell, as well as to the parietal wall of aperture. In most cases it bears variously pronounced spiral grooves, defining several differently pronounced spiral ridges, but can also be completely smooth. The ridges are especially well pronounced in Oliva, Agaronia, and Entomoliva. Further subdivision of the plication plate in the forner genus is described in detail by Tursch & Greifeneder's (2001: fig. 7.15). The border between the plication plate and anterior band is also represented by a step or ridge, which is referred to here as the rear edge of the plication plate (Fig. 5 ). The plication plate can be pronounced on the anterior part of the shell only, or it can extend significantly along the inner lip and parietal part of the aperture. It is adjoining or sometimes fused with the primary spire callus.

In some Olivella (e. g. in the subgenus Niteoliva [START_REF] Olsson | Studies of the genus Olivella[END_REF]) the primary spire callus extends anteriorly nearly along the entire plication plate, in fact becoming part of it, although Olsson (1956: fig. 1, 5) gave it a separate name (parietal callus).

The shell of Eburna, being generally rather typically ancillariid, has two unique characters, the homology of which is not obvious. First, it has two closely spaced olivoid grooves and therefore it is not clear which one is homologous with the olivoid groove of other Olivoidea.

Second, the plication plate is separated in its upper part from the parietal wall of the last whorl and forms a false umbilicus (Fig. 5, Eburnafu).

The shell of Olivoidea is callused to different degrees, the functional significance of which remains unclear. Sometimes (Belloliva, Calyptoliva) the callus is limited to a thin glaze on the parietal wall of the aperture, but more often it extends adapically, sometimes nearly reaching the upper spire whorls and effectively covering the entire spire, only leaving exposed the protoconch and a part of the cloak of the last whorl. In Ancillariidae, the callus nearly always overlays the suture, which cannot be seen externally. The callus can be coloured contrastingly to the shell cloak, or can be of the same colour. In some cases the callus seems to be uniform, but in others it obviously consists of two different layers, the primary (Fig. 6 I,Mpc) and the secondary spire callus (Fig. 6 I,Msc) (following the terminology of Kilburn, 1977). The primary spire callus extends spirally, while the secondary callus rather extends axially, covering the parietal part of the last whorl, sometimes also the columella. When both calluses are present, sometimes they fuse without distinct border in the upper part of the aperture but, at least in Entomoliva, it is obvious that the primary spire callus, developed as a band over the suture (and practically leaving the suture free), is slightly overlaid by the secondary callus above the aperture in side view of the shell (Fig. 6 H,J). It also should be mentioned that the secondary callus is in many cases positioned on top of the primary callus, deposited on the previous whorl and thus forming a complex multilayered structure.

A different pattern is observed in Olividae and Bellolividae. In Olivinae, the primary callus extends to the penultimate and upper spire whorls, but not to the last whorl. On the penultimate whorl, it does not reach the suture, leaving it open (for the definition of suture see below) (Fig. 6 E, F), but on the upper spire whorls it can overlay the suture. For the primary callus of Oliva, [START_REF] Tursch | Oliva shells. The genus Oliva and the species problem. L'Informatore Piceno[END_REF] used the term posterior callus. In Agaroniinae, the callus does not overlay the suture on spire whorls, but is otherwise similar to that in Oliva. In Olivellinae, the callus extends abapically on the parietal wall and fuses with the plication plate, sometimes extending as far as the anterior end of the shell. On the spire it can cover all whorls, but above the suture, leaving it open.

The suture on the shells of Olividae is unique and requires special attention. [START_REF] Sterba | Bemerkungen zur Schale der Olividae[END_REF] was the first to point out that the narrow channel seen between the whorls of Oliva does not correspond to the true suture. The whorls in Oliva do not come into direct contact with each other, but the subsequent whorl is connected to the spire callus of the previous one and, in addition, the thin layer is embedded between the whorl and spire callus of the previous whorl. Tursch & Greifeneder (2001: figs 7.04, 7.09) described the suture area of the shell in detail and came to the conclusion that the narrow channel does not correspond to a canaliculate suture and therefore proposed the term filament channel. This terminology is followed herein. A filament channel, which opens into the posterior edge of the aperture (Fig. 6 E-Ffc), is present in Olividae (except Calyptoliva) and Bellolividae, but is absent in other members of the superfamily.

The pseudoliviform shell

In the pseudolivifrom shell, which is much more "typical" of standard Neogastropoda (Fig. 3,D*,E,M), the callus is restricted to the columella and extends slightly onto the parietal wall of the aperture, but never onto the spire. The anterior band is absent. The shell is normally covered by periostracum, which can be very thick (e. g. in Pseudoliva). An olivoid groove is always present, usually terminating with a weakly pronounced labral denticle.

Soft body morphology and its correlation with shell

The external morphology of most Olivoidea (except Benthobiidae and Pseudolividae) is rather characteristic. The foot is broad, usually thin and clearly subdivided by a transversal furrow into the anterior propodium and posterior metapodium. The propodium is crescent-shaped and, on the dorsal side, separated into symmetrical halves by a longitudinal furrow (Figs 7; 8 A-B, E, F). In some Olivellinae [subgenus Olivella (Pachyoliva)], the lateral tips of the propodium are thickened and elongated (Fig. 7 F, arrow), and play an important role in feeding, producing the mucus nets in which the snails collect suspended detritus in the backwash [START_REF] Thiele | What can we learn from confusing Olivella columellaris and O. semistriata (Olivellidae, Gastropoda), two key species in panamic sandy beach ecosystems?[END_REF].

The metapodium has large paired lateral lobes, the parapodia, which laterally embrace the shell of the crawling snail. In Ancilla, they nearly completely cover the shell (Fig. 6 K), leaving free only the apical part of the spire. In Oliva (Kantor & Tursch, 2001) and Agaronia [START_REF] Rupert | Autotomy of the posterior foot in Agaronia (Caenogastropoda: Olividae) occurs in animals that are fully withdrawn into their shells[END_REF], the posterior part of the foot can be folded transversely to form a tightly sealed ventral pouch, in which the prey is kept after capture and during feeding (see details in Kantor & Tursch, 2001). The epithelium of this portion of the foot is different in colour and structure from the rest of the sole (Fig. 7 Dfp).

The head of Olivoidea is very variable, with a clear tendency to asymmetry. In Benthobiidae, the head morphology is rather typical of Neogastropoda, with more or less long cylindrical tentacles with closely spaced bases, with (Fusulculus) or without (Benthobia) eyes (Kantor, 1991, fig. 14 D;Simone, 2003: figs 7A, 9 A). In Pseudolividae, the cephalic tentacles are rather short, but with broad to very broad basal flaps bearing eyes. In Pseudoliva sepimenta (formerly Fulmentum), the head bears very short tentacles without basal flaps and large eyes (Fig. 8 C), it is slightly asymmetrical, with the rhynchostome, i.e. the opening through which the proboscis is everting, shifted to the base of the right tentacle.

In Olivinae, the head is externally symmetrical and is formed by paired, vertical, closely spaced, flaps which bear tentacles with eyes (Fig. 8 Afl, cept), however, a closer examination shows that the rhynchostome lies under the right tentacle. In all other groups (except Benthobiidae and Pseudolividae), the head is strongly asymmetrical and formed by dorsoventrally flattened cephalic flaps (Fig. 8 B,E,F). Sometimes the flaps are very broadly spaced and the left one is shifted to the base of the siphon.

The shell morphology of Olivoidea is correlated with mantle morphology, particularly with the presence of several appendages that are found in different combinations in different families.

It should be noted that in many cases it is impossible to examine the mantle in living crawling snails, since the parapodia completely cover the shell and, when disturbed, the animal withdraws immediately. Nevertheless, in Olividae -especially in Oliva, Agaronia and Olivella -the parapodia are less developed and cover the shell only laterally, thus allowing direct observations of the mantle appendages.

The presence of a filament channel on the shell is correlated with the presence of a posterior mantle filament (or posterior mantle tentacle) (Figs 7 A,8 Imf). This is a muscular agile rod, originating at the right anterior corner of the mantle, passing through the upper part of the aperture and positioned in the filament channel of the shell. The filament can move in the channel and can be completely withdrawn into the shell when the snail is disturbed. Its histology appears similar in the several studied species, Olivancillaria auricularia, Oliva sayana, Olivella verreauxii [START_REF] Marcus | Studies on "Olividae[END_REF] and in Olivella borealis [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF]. The filament consists of a layer of separate bundles of longitudinal muscles and contains numerous secretory cells and, at least in some species, a well developed nerve. The filament extends along all the length of the open part of the channel, which in Oliva is usually covered in its upper part by the callus, but can be free in Olivella and Agaronia, although often occluded with debris in upper whorls, thus indicating that the filament does not extend to the tip of the shell.

In all Olivoidea except Bellolividae, the right anterior corner of the mantle is transformed into a more or less extended posterior mantle lobe. In live Oliva animals, this very thin lobe is nearly transparent and highly extensible and covers the area of the primary callus, extending adapically beyond the limits of the aperture and posterior to the outer lip. In preserved specimens, the lobe is strongly contracted, even when the snails were relaxed in magnesium chloride prior to fixation (Fig. 8 Jml). As in Bellolividae neither the primary nor the secondary callus is pronounced and a mantle lobe is not visible (Kantor & Bouchet, 2007), we hypothesize that the primary spire callus is deposited by the posterior mantle lobe. In Pseudolividae and Benthobiidae a posterior mantle lobe is also present (Fig. 8 Dml), but there is no primary spire callus and the mantle lobe probably is responsible for depositing the thick parietal callus. In Pseudolividae it forms the plate-like parietal ridge that separates a narrow adapical extension from the rest of the aperture (Fig. 3 M,arrow).

In Ancillariidae, in addition to the primary, there can be a pronounced secondary spire callus. The distinction between the two is sometimes conventional, because in, e. g., Ancilla the seemingly uniform callus is covering the entire shell surface and it is not clear whether it can be considered a primary or a secondary one (Kilburn, 1981 considered it primary). Nevertheless in Amalda the distinction is rather clear on the ventral shell surface, where both calluses are adjoining each other and sometimes differ markedly in colour. In this case the posterior mantle lobe is hypertrophied and clearly subdivided in two partitions, or lobes (Fig. 8 G-Hml).

Therefore we hypothesize that both calluses are deposited by the mantle lobe, the primary callus is deposted by a smaller lobe, that is more ventral (Fig. 8 Hmlp) while the secondary callus is deposited by a broader lobe, situated more dorsally (Fig. 8Hmls).

The columella of Olivoidea can be smooth and callused or, as in Oliva and Agaronia, it can bear strong oblique plicae. Judging from their position callus and plicae should be deposited by the ventral side of the mantle adjoining the columellar muscle. This part of the mantle edge is relatively broad in Oliva, although very thin and, in preserved specimens, smooth, without any fold corresponding to the plicae. It is impossible to observe the ventral mantle side in crawling animals.

The left outer corner of the mantle edge surrounding the base of the siphon is free. In living snails, it is reflected outwards and forms an additional structure for which we herein suggest the term anterior mantle lobe (Fig. 7 Baml). From observations of live Oliva, Agaronia and Olivella, it extends through the siphonal notch as well as around the anteriormost part of the lip and covers the anterior (closest to the lip) region of the anterior band. When touched, it briefly retracts, but then extends again to the same position. Based on its exact correspondence with the position of the anterior band, there is little doubt that this is the structure responsible for it. The anterior band is present in most olivoideans (except Pseudolividae and Benthobiidae), although it can be very weak (e. g., in Bellolividae). In Olivinae, Olivellinae and Agaroniinae the lobe ends in a narrowed mobile partthe anterior mantle tentacle (Fig. 7 A,B,E,Gamt). It appears to be an autapomorphy of Olividae.

Digestive system

Anterior foregut. In general, the anterior foregut is typically neogastropod.

The proboscis is rather variable in morphology, from medium-long, cylindrical, to short or very short and conical in the contracted state. In Oliva the proboscis is medium-long, with the buccal mass situated anteriorly [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF]Kantor & Tursch, 2001). As observed during feeding observations on several species of Oliva, the proboscis is very extensible and mobile, and can reach the posterior foot poach, where the prey is kept (Kantor & Tursch, 2001). An anterior position of the odontophore in the retracted proboscis has also been recorded in Agaronia. In other Olivoidea, the proboscis is relatively shorter and the buccal mass is shifted away from the proboscis tip and often protrudes beyond the rear end of the retracted proboscis.

The proboscis is particularly short in Benthobiidae and Pseudolividae where the major part of the odontophore protrudes backwards [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF][START_REF] Simone | Family Pseudolividae (Caenogastropoda, Muricoidea): a polyphyletic taxon[END_REF]. In Olivellidae, the proboscis is very short and the odontophore lies nearly completely outside the proboscis, but when the proboscis is protracted the odontophore comes close to the mouth opening (Kantor, unpublished observations on Olivella olssoni van Regteren Altena, 1971). It should be noted that it is nearly impossible to estimate the length of the extended proboscis in live specimens based on its contracted state in preserved specimens, since it is capable of extreme contraction. The arrangement of the odontophoral retractor(s) differs between taxa. In Benthobiidae and Pseudolividae they are paired and pass through the nerve ring to fuse with the bottom of the body haemocoel [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF][START_REF] Simone | Family Pseudolividae (Caenogastropoda, Muricoidea): a polyphyletic taxon[END_REF]. In Bellolividae, Olivinae and some Ancillariidae the odontophoral retractor bypasses the nerve ring and is attached to the bottom of the haemocoel. In Olivellidae it is branched with some branches passing through the nerve ring, and others bypassing it (unpublished observations). The proboscis retractors are thin, few and attached laterally to the middle or posterior part of the rhynchodaeum.

In Benthobiidae and Pseudolividae, the salivary glands are compact and acinous, while in Bellolividae and Olividae they are rather loose, composed of narrow branching tubes (ramifiedtubular). In Ancillariidae three types of salivary glands are present, ramified tubular in Amalda, Eburna, and Turrancilla, acinous in some Ancilla, or broad and tubular in other Ancilla, Ancillina and in Entomoliva. Accessory salivary glands were observed in most Olivoidea. In most cases they are tubular, which is standard in Neogastropoda, but in Benthobiidae the single accessory salivary gland ends in a very large muscular sac [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF][START_REF] Simone | Revision of the genus Benthobia (Caenogastropoda, Pseudolividae)[END_REF].

The radula of Olivoidea is rather variable and is generally genus specific (although considerable within-genus disparity can be found in Ancilla -Figs 13 and 17 J-L). In most cases a radula row consists of three teeth, but in Olivellinae it contains five (Fig. 9 G-H). Judging from the tree topology, this is an autapomorphy of the subfamily. In addition to the multicuspid rachidian with convex base, there are simple, short, curved lateral teeth. In dorsal view these seem cylindrical, but in some projections it is obvious that they are flat plates, convex on the dorsal side and concave ventrally. Interestingly, and contrary to what is seen in other olivoideans, they are directed outwards, not towards the center of the membrane. Laterally, there are additional flat rectangular cuspless plates with slightly thickened edges that are very slightly lifted above the membrane. Only in Pseudoliva are the laterals bicuspid (Fig. 10 A,B); in other genera of Olivoidea the lateral teeth are unicuspid, rather uniformly hook-shaped with a more or less broad base. In many cases the teeth are convex on the anterior (dorsal) side and concave on the posterior side. In Olivinae, Agaroniinae, Calyptoliva and Bellolividae the tip of the lateral tooth is characteristically attenuated and is additionally bent (Fig. 9 A,C,D,F). The teeth can be bifurcated on the tip (e. g., in Amalda cf. hilgendorfi richeri -Fig. 10 E,F, and particularly in Ancilla atimovatae -Fig. 17 J-L). The central tooth displays more disparity, from tricuspid (e. g. All Olivinae have tricuspid central teeth with a shallowly notched anterior edge and more or less extended lateral flaps of the central tooth (Fig. 9 F; Kantor & Tursch, 2001: figs 6.20-6.27). In Turrancilla (Ancillariidae), all species have very similar central tooth with three large pointed cusps and a very deeply notched anterior profile (Fig. 10 H). The posterior edge of the base of the central tooth forms a distinct, long and posteriorly rounded projection which corresponds in shape to the notch of the adjoining row. Rather similar central teeth are found in the two known species of Entomoliva (Fig. 10 C).

Most Olivoidea have a well-developed pyriform valve of Leiblein situated just in front of the circumoesophageal nerve ring. The gland of Leiblein is bulky or tubular; in Ancilla it is very tightly coiled and its posterior part can have muscular walls; the duct of the gland is always markedly constricted. The mid-oesophagus (between the valve of Leiblein and the duct of the gland) is lined with a glandular epithelium. A remarkable exception is Olivella which altogether lacks a gland of Leiblein, a valve of Leiblein, as well as a glandular mid-oesophagus.

In all olivoideans, the stomach is usually sac-like, with a long to very long and narrow posterior mixing area. In Olivella the entrance of the posterior oesophagus and the intestine opening are very close together and the stomach has a thick muscular ring in the middle of the posterior mixing area [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF] fig. 6 A-F). In Amalda the very different stomach is tubular, U-shaped, without any posterior mixing area (original data).

TAXONOMY

In the following section we provide brief diagnoses for all recognized suprageneric taxa of Olivoidea and list the valid genera attributed to the recognized families. Genera are marked in

Diagnosis

Shell variable, from broadly oval to narrowly fusiform, with high last whorl and diameter of the spire whorls expanding slowly or rapidly. Siphonal canal either absent (in which case the shell is strongly notched) or very short, unnotched. Shell sometimes covered with thick periostracum, but in most cases not and then shell surface glossy. Protoconch multispiral or paucispiral. In all families except Pseudolividae and Benthobiidae, anterior end of the shell strongly callused and bearing an anterior band. Callus differently pronounced, sometimes limited to inner apertural lip, but at other times extending on significant part of the last whorl and spire, or even completely covering the shell (some Ancillidae). Radula with 5 to 3 teeth per transverse row.

FAMILY OLIVIDAE LATREILLE, 1825

Type genus: Oliva Bruguière, 1789.

Diagnosis

Shell glossy, lacking periostracum, broadly to narrowly fusiform, with high to very high last whorl and narrow aperture tapering adapically. Siphonal canal absent, anterior end of shell distinctly notched. Anterior part of the shell forming a well defined anterior band, raised above the shell cloak. Plication plate nearly always with more or less defined spiral plicae. Primary spire callus present, covering at least partially the spire whorls. Narrow filament channel present, but in Calyptoliva overlaid by narrow primary callus.

Foot with well developed crescent-shaped propodium, with median longitudinal cleft on dorsal side and parapodia, partially embracing the shell. Radula with 3 or 5 (Olivella) teeth per transverse row. Mantle with mantle filament (except in Calyptoliva), with posterior and anterior mantle lobes, and with anterior mantle tentacle (except in Calyptoliva and Olivancillaria).

SUBFAMILY OLIVINAE LATREILLE, 1825

Type genus: Oliva Bruguière, 1789.

Diagnosis

Plication plate subdivided into parietal plate, shoe and belt. Filament channel well defined, eventually overlaid by primary spire callus on upper spire whorls, but free at least on last whorl.

Operculum absent. Anterior mantle tentacle well defined. Head with two vertical flaps with narrow tentacles bearing eyes. Radula with three teeth per row, central tooth with short or long lateral flaps and three large cusps. Stomach with narrow and rather long posterior mixing area.

Genus Oliva Bruguière, 1789 (1). Type species: Voluta oliva Linnaeus, 1758; by subsequent monotypy by Lamarck, 1799.

Remarks

The genus Oliva is rather heterogeneous and a number of subgenera have been proposed, some of them even raised to full genus [START_REF] Petuch | Molluscan communities of the Florida Keys and adjacent areas: their ecology and biodiversity[END_REF]. [START_REF] Petuch | Atlas of the living olive shells of the world[END_REF] recognized 19 subgenera, of which five were monotypic. [START_REF] Tursch | Oliva shells. The genus Oliva and the species problem. L'Informatore Piceno[END_REF] discussed briefly their status but abstained from the use of subgenera, since in many cases they did not find clear conchological distinctions between them. On the contrary, [START_REF] Hunon | Olividae (Mollusca, Gastropoda). A complete survey of Recent species of the genus Oliva. Xenophora, Conchbooks[END_REF] used as valid most of the subgenera that had been proposed earlier, while still leaving some species in Oliva (e.g. O. lacanientai Greifeneder & Blöcher, 1985). Our COI molecular analysis demonstrates that at least some of the subgenera in their current usage are paraphyletic (Fig. 11 abbreviated subgenera indicated in bold after species names). Nevertheless, we foresee that, with increased taxon sampling in molecular phylogenies, a number of (sub)genera will be found to be justified. However, with the molecular data currently available, we simply list below the accepted subgenera, but abstain from using them.

The species delimitation approach would suggest that O. sericea and O. miniacea would be a single species. However, because they are represented by a single specimen each in our analysis and because the two species are distinct conchologically, we remained conservative and did not change the status of the species. Furthermore, Oliva lacanientai shows great within species variability…

List of nominal subgenera (in alphabetic order)

Acutoliva [START_REF] Petuch | Atlas of the living olive shells of the world[END_REF] [START_REF] Petuch | Atlas of the living olive shells of the world[END_REF] where it is attributed to [START_REF] Mörch | Catalogus conchyliorum quae reliquit D. Alpfionso d'Aguirra et Gadea Comes de Yoldi[END_REF].

Hoewever, earlier than Mörch, Gray (1847: 140) had already made the name available (Petit, 2012)

SUBFAMILY OLIVELLINAE TROSCHEL, 1869

Type genus: Olivella Swainson, 1831.

Diagnosis

Plication plate not distinctly subdivided, with defined spiral plicae that can extend to columella. Internal shell walls partially resorbed. Primary callus extending up to anterior end of plication plate. Filament channel well defined, free on most spire whorls. Anterior mantle tentacle well defined. Head with small, broadly spaced, dorso-ventrally compressed flaps; eyes absent. Operculum present or absent. Radula with 5 teeth per transverse row; central tooth broad, with numerous short cusps and convex anterior edge. Lateral teeth hook-shaped; additional rectangular weak plates exterior to lateral teeth.

Genus Olivella Swainson, 1831 (1). Type species: Olivella purpurata Swainson, 1831 [= Voluta dama Mawe, 1828]; SD, [START_REF] Dall | Contributions to the Tertiary paleontology of the Pacific coast. I. The Miocene of Astoria and Coos Bay, Oregon[END_REF].

Remarks

Olivella is conchologically rather variable, and a number of (sub)genera have been proposed. Three species only were included in our molecular analysis and, based on this limited dataset, we cannot make any conclusion on the possible subgeneric division of the genus. Below we follow the classification proposed by [START_REF] Olsson | Studies of the genus Olivella[END_REF] with some additions and corrections.

List of nominal subgenera (in alphabetic order)

Anasser [START_REF] Absalão | A new subgenus and three new species of Brazilian deep waters Olivella (Mollusca, Gastropoda, Olivellidae) collected by the RV Marion Dufresne in 1987[END_REF] [START_REF] Olsson | Studies of the genus Olivella[END_REF] as genus of uncertain status, pending data on radular morphology, although Peile (1922: fig. 6) Genus Calyptoliva Kantor & Bouchet, 2007 (1). Type species: Calyptoliva bolis Kantor & Bouchet, 2007; OD. Kantor et al., 2016 is the only sequenced species and forms a long branch and the sister group cannot be recognized in our analysis. However, the genus is confidently placed in Olividae and differs from all other representatives of the family by the absence of a channeled suture and, correspondingly, by the absence of the mantle filament. The suture is overlaid by a very narrow primary spire callus. [ Gray (1858: 48) listed Oliva claneophila in the synonymy of Claneophila gibbosa.

Remarks C. bbugeae

Insertae sedis

Genus

Although it is not clear which species he meant under the name gibbosa, since no authorship or date was provided, the remaining species attributed by Gray to Claneophila, beside O. claneophila itself, are species of Olivancillaria]

Remarks

The position of Olivancillaria remains unclear. We did not have access to molecular material and therefore the genus is not included in our analysis. In their revision of Olivancillaria in the southwestern Atlantic, [START_REF] Teso | A revision of the genus Olivancillaria (Mollusca: Olividae) from the southwestern Atlantic[END_REF] briefly considered similarities and differences with other genera of Olivoidea and suggested affinities to Agaronia. [START_REF] Marcus | Studies on "Olividae[END_REF] described the anatomy of two species, O. vesica (as Lintricula auricularia) and O. urceus (as O. brasiliensis). The radular characters of all examined species are similar to that of Agaronia in having rachidian teeth with three large cusps and smaller additional cusp exterior to the main lateral cusp on each side of the tooth (Fig. 9 E), a character not found in Olivinae. Besides, the head has flaps without eyes and tentacles, similar to the situation in Agaronia. Based on the description of [START_REF] Marcus | Studies on "Olividae[END_REF], [START_REF] Golikov | Systematics of prosobranch Gastropoda[END_REF] erected the new family Olivancillariidae, remarking the differences in shell and radulae as well as in some details of the female reproductive system, which however are not very clearly described by Marcus & Marcus. Despite a radula similar to that in Agaronia, Olivancillaria lacks the olivoid groove and anterior mantle tentacle. Molecular data will be crucial to clarify the position of Olivancillaria and, although morphologically and conchologically it is close to Olivinae, it may prove to form a distinct lineage, consistent with the recognition of a separate subfamily.

Remarks

When describing Janaoliva, [START_REF] Sterba | Olivella (Janaoliva) amoni, a new subgenus and species from the Bismark Archipelago and Indonesia (Mollusca: Gastropoda: Olivellidae)[END_REF] compared it with Olivella, correctly pointing out the very simplified structure of the plication plate, which differs from other subgenera of Olivella. Kantor & Bouchet (2007) overlooked the description of Janaoliva and thus did not compare Olivella amoni with the very similar species Olivellopsis simplex, which they included in Belloliva. The very long branches of both Belloliva and Olivellopsis suggest that they are two separate genera. In the vicinity of Kavieng (New Ireland), the type locality of O. amoni, another similar, although molecularly different species was recently found. Their radulae are similar (Fig. 9 C) and are close to the radula of O. simplex from New Caledonia (Kantor & Bouchet, 2007: fig. 12 A-D). In the light of the new data indicating the presence of a species complex, O. simplex from New Caledonia may be specifically different from O. simplex, the type locality of which is the "Paumotus Islands" [Tuamotu Archipelago], French Polynesia. We also found very similar shells off Madagascar. It is possible that a whole radiation of Olivellopis will be uncovered by molecular phylogenetics.

?Genus Jaspidella Olsson, 1956 (2). Type species: Voluta jaspidea Gmelin, 1791; OD

Remarks

The genus was established in the subfamily Olivinae on the basis of the tricuspidate rachidian tooth. The type species from the Caribbean has a shell similar to that of Belloliva, as well as the operculum, also characteristic for this family. No material was available to us and we attribute Jaspidella conditionally to Bellolividae on the basis of shell and radula similarities.

FAMILY ANCILLARIIDAE SWAINSON, 1840

Synonym: Ancillinae H.

Remarks

The monotypic family Vanpalmeriidae is characterized by a heavily callused spire and few weak plications on the columella. It was originally classified in the Volutacea and was reduced to a subfamily of Olividae by [START_REF] Bouchet | Classification and nomenclator of gastropod families[END_REF] together with Olivinae and Ancillariinae.

Although the position of this enigmatic species remains unclear, it is transferred here conditionally to Ancillariidae based on the presence of the strong primary spire callus that is overlaying the suture.

Genus Ancilla Lamarck, 1799 (1) (1981) tentatively used Sparella, Sparellina, Chilotygma and Hesperancilla as subgenera of Ancilla. The obtained sequences show considerable genetic distances between the 11 groups of Ancilla delimited with ABGD (Fig. 12), and the radulae also show considerable disparity (Figs 13,17,19). Therefore the presence of discrete (sub)genera is possible, although the currently accepted system is hardly plausible. Kilburn's recognition of Sparella and Sparellina was based mainly on radular characters, Sparella generally having a tricuspid rachidian teeth, often with intermediate denticles between main cusps, while Sparellina has multicuspid rachidian teeth. We examined the radula of the type species Ancilla cinamomea (Fig. 13 J), and it proved to have a tricuspid rachidian. Thus Sparella in the definition of Kilburn becomes a synonym of Ancilla (Ancilla), and most studied species should be attributed to the nominative subgenus. The topology of the COI-based tree of Ancilla (Fig. 12) is nearly identical to that of the reduced dataset based on four genes (Figs 1, 2). A COI sequence was not available for Ancilla atimovatae Kantor et al., 2016, but in the four-gene tree it is sister to the clade including A. ventricosa (Lamarck, 1811), A. adelphe Kilburn, 1982 andA. lhaumeti Kantor et al., 2016. The radula of A. atimovatae is very distinctive (especially in the shape of the bifurcating lateral teeth), but is more similar to the radula of Sparellina (as defined by Kilburn, 1981). Thus Ancilla s.s. (= Sparella) becomes paraphyletic. Moreover, Ancillista aureocallosa Kilburn & Jenner, 1977, the only species of Ancillista that was available to us, was confidently placed in Ancilla. Its radula (Fig. 13 A) is similar to some other Ancilla spp. (e.g., Ancilla boschi Kilburn, 1980;Kilburn 1981: fig. 83). In the four-gene analysis, Ancillista aureocallosa is sister to Ancilla sp. (MNHN IM-2007-36606), while in the COI analysis it is basal to all other Ancilla.

Remarks

Kilburn

The radula of the type species of Ancillista remains unknown, but that of Ancillista muscae (unpublished) is different from A. aureocallosa in the shape of the central tooth, although it is also tricuspid. Therefore we conditionally synonymize Ancillista with Ancilla, pending molecular data on the type species.

We did not have specimens of the type species of Chilotygma, and Hesperancilla and cannot draw any conclusions on them. Both were established based on the shell characters, and we conditionally treat them as separate entities.

Genus Ancillina Bellardi, 1882 (1). Type species: † Ancillaria pusilla Fuchs, 1877; M (Austria, Miocene) Synonym: Ancilla (Gracilancilla) Thiele, 1925. Type species: Ancilla sumatrana Thiele, 1925; OD Genus Anolacia Gray, 1857 (3). Type species: Ancilla mauritiana G.B. Sowerby I, 1830; M.

Remarks

Anatomy and radular characters remain unknown, therefore the validity of the genus is uncertain.

Genus Amalda H. Adams & A. Adams, 1853 (1). Type species: Ancillaria tankervillii Swainson, 1825;SD, Cossmann (1899).

Remarks

The species of Amalda together present a wide range of shell forms and it is not surprising that several genus group names have been proposed. Our analysis covers very little of the general diversity of the group, but at least three subgenera are present in our dataset: Amalda s.s., Baryspira [represented in our analysis by Amalda contusa (Reeve, 1864)] and Alocospira [represented by Amalda bellonarum Kilburn & Bouchet, 1988]. Although the support of the genus itself is significant (1 in BA), the genetic distances within its constituants are rather small, compared to, e.g., the genus Ancilla. Nevertheless, at least Alocospira constitutes a rather long branch, and it is possible that future molecular studies will reveal a more complex and robust structuring necessitating the recognition of (sub)genera. At present we abstain from using any subgenera and provide a list of names with Recent type species for future reference.

List of nominal subgenera (in alphabetic order

Remarks

We examined the anatomy of Eburna lienardi (unpublished). It is rather typical ancillariid and the genus is attributed to the family on the basis of morphology and radula (Fig. 10 I). Bouchet & Kilburn, 1991 (1). Type species: Entomoliva incisa [START_REF] Bouchet | A new genus of Ancillinae from New Caledonia, with the description of two new species[END_REF]; OD.

Genus Entomoliva

Genus Micrancilla Maxwell, 1992 (3). Type species: † Amalda granum Maxwell, 1992 (New Zealand, Eocene); OD.

Remarks

In addition to several fossil species from New Zealand and recently described ones from the Paris and Aquitanian basins, the Recent species Ancillaria longispira Strebel, 1908, from the Falkland Islands, was attributed to Micrancilla by [START_REF] Pacaud | Nouvelles espèces d'Ancillariinae (Mollusca: Gastropoda: Olividae) du Paléogène des bassins de Paris et d'Aquitaine[END_REF]. Neither the anatomy nor the radula of M. longispira have been examined so far.

Genus Turrancilla Martens, 1904 (1). Type species: Ancillaria lanceolata Martens, 1901 [a primary homonym of Ancillaria lanceolata Tate, 1889; has been renamed Ancillus (Turrancilla) akontistes Kilburn, 1980]; SD, Wenz (1943).

FAMILY BENTHOBIIDAE, fam. nov.

Type genus: Benthobia Dall, 1889.

Diagnosis

Shell covered by periostracum, ovoid to biconical, with high or very high last whorl and medium-broad to broadly ovate aperture. Siphonal canal very short and broad, not notched.

Anterior band not pronounced. Olivoid groove present, terminating by a short denticle on the lip.

Plication plate either weakly pronounced, limited to columella, smooth (Benthobia) or absent (Fusulculus). Primary spire callus limited to inner lip, extending up to parietal wall. Spiral sculpture either absent (Benthobia) or consisting of rather distinct spiral cords (Fusulculus).

Suture simple, adpressed.

Foot with narrow propodium, lacking parapodia. Operculum large, with terminal nucleus.

Radula with 3 teeth per transverse row. Central tooth either tricuspid (Fusulculus; [START_REF] Bouchet | Two new deep-water Pseudolividae (Neogastropoda) from the south-west Pacific[END_REF] or multicuspid (Benthobia; [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF][START_REF] Simone | Revision of the genus Benthobia (Caenogastropoda, Pseudolividae)[END_REF]. Lateral teeth unicuspid. Mantle without mantle filament, with weak to moderately pronounced posterior mantle lobe. Head with long cylindrical tentacles, no eyes in all studied species. Rhynchostome shifted to the base of right tentacle. Proboscis very short when retracted, with basal buccal mass and radular odontophoral retractor passing through the nerve ring. Salivary glands acinous.

Accessory salivary gland terminating with very large muscular bulb.

Genus Benthobia Dall, 1889 (1). Type species: Benthobia tryonii Dall, 1889; OD. Synonym: Nux [START_REF] Barnard | New species of South African marine gastropods[END_REF]. Type species: Nux alabaster [START_REF] Barnard | New species of South African marine gastropods[END_REF]; OD.

Remarks

In the absence of anatomical data, [START_REF] Simone | Revision of the genus Benthobia (Caenogastropoda, Pseudolividae)[END_REF] considered Nux a separate genus, pointing out its close affinity to Benthobia. The radula of N. alabaster is nearly identical to that of Benthobia [START_REF] Barnard | New species of South African marine gastropods[END_REF] and there are no good reasons to consider it a separate genus. The mantle lobe (called anal siphon by Simone 2003) is moderately pronounced, simple.

Genus Fusulculus Bouchet & Vermeij, 1998 (1). Type species: Fusulculus crenatus [START_REF] Bouchet | Two new deep-water Pseudolividae (Neogastropoda) from the south-west Pacific[END_REF]; OD.

Remarks

The anatomy of Fusulculus crenatus (own observations) is similar to that of Benthobia spp., especially the anterior foregut. The mid-oesophagus is glandular, as in Benthobia, although the glandular epithelium is more pronounced in the former. The synapomorphy is the presence of a single accessory salivary gland, terminating in a very large muscular bulb. Similarly, in both genera, the anteriormost part of the osphradium lacks lamellae on the left side.

FAMILY PSEUDOLIVIDAE de GREGORIO, 1880

Synonym: Zemiridae [START_REF] Iredale | Results from Roy Bell's molluscan collections[END_REF] Type genus: Pseudoliva Swainson, 1840.

Diagnosis

Shell covered by periostracum, ovoid to fusiform, with high or very high last whorl and medium broad to broad ovate aperture. Siphonal canal short to very short, moderately broad, shallowly to deeply notched. Anterior band not pronounced. Olivoid groove present, terminating by short denticle on outer aperture lip, sometimes denticle poorly pronounced. Plication plate not pronounced. Primary spire callus limited to inner lip, at most slightly extending to parietal wall.

Parietal wall sometimes forming well defined fold, partially separating the upper part of aperture (Pseudoliva). Spiral sculpture absent (Pseudoliva) or present. Suture simple, adpressed, eventually broadly channeled (Zemira).

Foot with narrow propodium, lacking parapodia. Operculum large, with terminal nucleus.

Radula with 3 teeth per transverse row. Central tooth tricuspid. Lateral teeth unicuspid or bicuspid (Pseudoliva, Fig. 10 A-B). Mantle without mantle filament, with weak to moderately pronounced posterior mantle lobe. Head with short flaps with eyes, sometimes flaps producing short tentacles. Rhynchostome slightly shifted to the base of right tentacle. Proboscis very short when retracted, with basal buccal mass and radular odontophoral retractor passing through the nerve ring. Salivary glands acinous. Accessory salivary gland tubular.

Remarks

The anatomy of various Pseudolividae has been reasonably well studied, for some species in detail [START_REF] Ponder | The genus Zemira H. and A. Adams (Mollusca: Neogastropoda)[END_REF][START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF][START_REF] Simone | Family Pseudolividae (Caenogastropoda, Muricoidea): a polyphyletic taxon[END_REF].

The genera Macron H. Adams & A. Adams, 1853 (type species: Buccinum aethiops Reeve, 1847; M) and Triumphis Gray, 1856 (type species: Buccinum distortum W. Wood, 1828; M). were attributed to Pseudolividae by [START_REF] Vermeij | Generic revision of the neogastropod family Pseudolividae[END_REF]. [START_REF] Thiele | What can we learn from confusing Olivella columellaris and O. semistriata (Olivellidae, Gastropoda), two key species in panamic sandy beach ecosystems?[END_REF] illustrated the radula of the former and described that of the latter. The tricuspid lateral teeth (not found in any other Pseudolividae) and multicuspid central ones are more similar to Buccinoidea. [START_REF] Landau | Systematics and palaeobiogeography of the gastropods of the middle Miocene (Serravallian) Karaman Basin, Turkey[END_REF] suggested a placement of Macron in Nassariidae, a position that was not rejected by [START_REF] Galindo | The phylogeny and systematics of the Nassariidae revisited (Gastropoda, Buccinoidea)[END_REF] but requires molecular confirmation. The position of Triumphis is even more uncertain, but it may be tentatively assigned to the Nassariidae as well.

Genus

Remarks

Besides size, the distinction between Pseudoliva and Fulmentum is in the better development of the plate-like parietal ridge (Fig. 3 M), which divides the aperture of Fulmentum in two compartments; however, a similar, although weaker, plate is also present in the type species of Pseudoliva. Pseudoliva sepimenta and P. crassa have rather similar and characteristic bicuspid lateral teeth, differing in this from all other pseudolivids that have unicuspid lateral teeth. We consider the conchological differences insufficient for separation of two genera. Kilburn, 1989 (2). Type species: Naudoliva caitlinae Kilburn, 1989; OD. Genus Luizia Douvillé, 1933 (2). Type species: Buccinum costae Douvillé, 1933; M.

Genus Naudoliva

Genus

Remarks

The only Recent species, L. zebrina (A. Adams, 1855) was examined anatomically by [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF] and proved to be similar to Pseudoliva ancilla.

Doubtful taxonomic position

Genus Ramoliva [START_REF] Cotton | South Australian shells (including descriptions of new genera and species) Part III[END_REF]. Type species: Olivella adiorygma Verco, 1909; OD.

Remarks

The position of this monotypical genus is unclear. Overall, it has an olivoid shell with nonchanelled sutures, a very simplified pication plate and, according to the original illustration of the type species (Verco, 1909: pl. 25, figs. 3-4), it lacks an anterior band. The last whorl is lifting towards the aperture, a character not known in any other Olivoidea. The species is known only from empty shells. [START_REF] Lee | What's a rice olive, anyway?[END_REF] has suggested that this may be a columbellid, a hypothesis that we find convincing.

DISCUSSION

PHYLOGENY Of THE OLIVOIDEA

One of the most unexpected -based on anatomical characters -results of the phylogenetic analysis is the close relationship between Pseudolividae (in their previously accepted taxonomic extension, e. g. by [START_REF] Vermeij | Generic revision of the neogastropod family Pseudolividae[END_REF] and the remaining Olivoidea. Based on their anatomy [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF] had rejected the similarity between two groups and even suggested the new suborder Pseudolivoidei. This view was partially accepted later and, in the latest working classification of the Gastropoda [START_REF] Bouchet | Classification and nomenclator of gastropod families[END_REF], the Pseudolividae were attributed together with the Ptychatractidae to the superfamily Pseudolivoidea. However, recent phylogenetic analyses of "mitriform" gastropods [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF] have already demonstrated that Ptychatractidae as previously recognized are paraphyletic, with some genera (Latiromitra Locard, 1897 andCeratoxancus Kuroda, 1952) included in the family Costellariidae, and others representing independent lineages, none of them showing close affinity to the Pseudolividae, and there is currently no support for classifying them in the same superfamily. It should be remembered that [START_REF] Thiele | What can we learn from confusing Olivella columellaris and O. semistriata (Olivellidae, Gastropoda), two key species in panamic sandy beach ecosystems?[END_REF] classified Pseudolivinae as a subfamily of Olividae, a position followed by [START_REF] Wenz | Gastropoda. Teil 1, Allgemeiner Teil und Prosobranchia[END_REF], with all other olivoideans united in the subfamily Olivinae. Thus, in some respects, the new classification proposed herein is a return to the roots.

Another unexpected result is the non-monophyly of Pseudolividae, which is split here in two families, Pseudolividae and Benthobiidae fam. nov. The former inhabit shallow water (subtidal to 150 m) off western and southern Africa (Pseudoliva, Luizia, Naudoliva) and Australia (Zemira). The new family Benthobiidae currently includes only the two genera Benthobia and Fusulculus, both from bathyal and abyssal depths of the Atlantic and Indo-Pacific regions. The paraphyly of Pseudolividae has already been discussed by [START_REF] Simone | Family Pseudolividae (Caenogastropoda, Muricoidea): a polyphyletic taxon[END_REF] on the basis of a cladistic analysis of morphological characters. His analysis however failed to recognize Pseudolividae as a distinct family, which was recovered as a grade, with Nassodonta H. Adams, 1867 (Nassariidae) emerging as a sister group. Thus, although Simone's paraphyly of the Pseudolividae is congruent with our own results, its relationship to other Neogastropoda as hypothesized here is very different from that suggested by [START_REF] Simone | Family Pseudolividae (Caenogastropoda, Muricoidea): a polyphyletic taxon[END_REF].

Conchologically and anatomically Pseudolividae and Benthobiidae are different from other

Olivoidea, although there are not many autapomorphies that can characterize them.

Conchologically the two families differ from all other olivoideans in the absence of the characteristic anterior band, the presence of periostracum, and the very limited development of the primary callus that does not cover the spire. Pseudolividae and Benthobiidae share the presence of the olivoid groove, which is present in at least some Olividae (Agaroniinae) and many Ancillariidae. A very similar groove can however also be found in other unrelated neogastropods, e. g. Ceratoxancus (currently a basal Costellariidae). Perhaps the only autapomorphy of the Benthobiidae is the very unusual accessory salivary gland terminating in a large muscular bulb, described for the first time in detail by [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF], reconfirmed by [START_REF] Simone | Revision of the genus Benthobia (Caenogastropoda, Pseudolividae)[END_REF] and herein also found in Fusulculus.

Olivoidea as a clade is not significantly supported in our analyses, while the node uniting all of them except Bellolividae is highly supported (Figs 1, 2). Although Bellolividae shares a number of characters with Olividae (filament channel of the shell and correspondingly the presence of a mantle filament) and other oliviforms (anterior band of the shell, propodium morphology, parapodia), its relationships to other Olivoidea is not resolved. To check whether the ambiguous position of Bellolividae could be an artifact of the analysis induced by the inclusion of the very different Pseudolividae and Benthobiidae, we reran an analysis of the dataset after removing the two latter families. The resulting tree did not differ significantly in topology and the support of the basal node did not change much, still remaining rather low.

The trees obtained leave the deeper relationships between the major groups of Olivoidea unresolved. Therefore they do not rule out the possibility, that Benthobiidae and Pseudolividae may appear to be sister groups in future analyses with a dataset that will include more taxa. All major terminal clades are well supported and the taxonomic conclusions at the family/subfamily levels are well grounded. However attempts to reconstruct ancestral states and transformation of the characters with Mesquite software (Maddison & Maddison, 2007-2010) were unsuccessful (results not shown), as the analyses suggested multiple independent origins of obviously homologous characters. Thus, according to that analysis, the anterior band of the shell originated independently in Bellolividae, Olividae and Ancillariidae; similarly, multiple origins were suggested for the crescent-shaped propodium, parapodia, etc. We view this as an artifact of the tree due to unresolved nodes and we abstain from attempts to reconstruct the evolutionary transformations of these characters until a more resolved tree is available. The poor resolution of basal and intermediate nodes may be a result of either insufficient taxon sampling, or insufficient resolution of the genes used in the analysis.

The close relationship of Olivellinae with Olivinae and Agaroniinae is also an unexpected result of this research. [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF] pointed out characters that distinguish Olivella from Oliva, among them are a very distinctive radular morphology, the resorption of the inner shell walls in the upper whorls, the seemingly "absence" of the head, valve and gland of Leiblein, and the position of the odontophore at the base of the retracted proboscis. The present study nevertheless demonstrated that at least some of these characters are not autapomorphies of Olivellinae. Thus [START_REF] Marcus | Studies on "Olividae[END_REF] and [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF] had considered that Olivella lacks a differentiated head, since the triangular, dorso-ventrally compressed flaps are very asymmetrical and the rhynchostome opens below the right flap. However, examination of different olivoideans shows that the head is asymmetrical in nearly all of them and, even in Oliva which has a normally symmetrical head formed by vertical flaps with tentacles and eyes, the rhynchostome opens under the right flap. A similar position of the rhynchostome has been observed in Benthobiidae (but we lacked adequate material to check it in Pseudolividae). A "head" with dorso-ventrally compressed flaps is found in a number of olivoideans, including Agaronia and Ancillariidae, and thus can not be considered a unique character of Olivellinae. However, the peculiar radula really sets Olivella apart from all other members of the superfamily. The homology of the outermost plates is not clear; although the lateral teeth in Olivella look different from other olivoideans (e. g. they are curved outwards, not inwards) (Fig. 9 G-H) they are most probably homologous with the lateral teeth of other olivoideans, since they adjoin the rachidian tooth. Thus the outermost plates are either the remains of the marginal teeth, or de novo structures. Nothing similar was found in either the outgroup neogastropod taxa, or other members of Olividae. Given that the majority of basal caenogastropods have a taenioglossate radula with seven teeth per row, and that in Neogastropoda the number of teeth per row is usually three, two or even one, it is generally assumed that the evolutionary tendency in Neogastropoda is a reduction in number of teeth per row. However, there are five teeth per row in different conoidean taxa (see, e. g., [START_REF] Taylor | Foregut anatomy, feeding mechanisms and classification of the Conoidea (= Toxoglossa)(Gastropoda)[END_REF][START_REF] Bouchet | A new operational classification of the Conoidea (Gastropoda)[END_REF], and we hypothesize that the appearance of the marginal plate-like teeth is an autapomorphy of Olivellinae.

By contrast to our findings on the position of Olivellinae, confirmation that Ancillariidae represent its own well-supported clade of family rank is not unexpected. There are several conchological and morphological characters that clearly define the family. Already in the morphology-based cladistic analysis performed by [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF] Olivella and Oliva were closer to each other than to Amalda. In Ancillariidae the shell is more strongly callused than in any other Olivoidea. The suture is always overlaid with the primary callus, which can be rather narrow, but sometimes (in some Ancilla) is extremely broad and covers the entire shell surface. This is reflected in the mantle morphologythe mantle filament is absent, while the posterior mantle lobe responsible for depositing the primary callus is very large and can be hypertrophied.

The anterior foregut, on the contrary, does not differ markedly from other olivoideans. We are still missing three valid genera in our analysis -Ancillista (the species included in our analysis, A. aureocallosa, is rather distinct from the type species, A. velesiana), Eburna and Micrancillaand their inclusion may change the relationships between genera. At this moment, all recognized genera are however highly supported. The genetic heterogeneity observed in Ancilla is also paralleled by the high disparity of radula characters in that genus (Figs 13,19). The remaining Ancillariidae for which the radulae are known are characterized by a rather conservative within-genus radular morphology. The current knowledge on the radula of Ancillariidae and the range of its within-genus variability now allows clarification of the position of the deepest water olivoidean species, namely Amalda sibuetae Kantor & Bouchet, 1999, described from a depth of 1733-1885 m from off Mauritania. It possesses a radula typical of Turrancilla and should be reclassified in that genus. Kantor & Bouchet (1999) pointed out the similarities of the radula of Amalda sibuetae to the type species of Turrancilla, Ancillaria lanceolata Martens, 1901, but due to the then insufficient data on the radular morphology in Turrancilla, they did not attribute the new species to that genus.

MORPHOLOGY

One of the remarkable (although not unique) characters of oliviform shells is the great development of the different callused structures: anterior band, columellar callus (= plication plate), primary and secondary spire calluses. Correlation of the soft body anatomy with these shell characters and observations of live animals has allowed us to suggest for the first time which part of the mantle is responsible for depositing which callused feature.

As was stated above, both primary and secondary spire calluses in oliviform shells (as well as the parietal callus in pseudoliviforms) are deposited by the same mantle outgrowth, the posterior mantle lobe. Characteristic and distinctive between different groups of Olivoidea, the shell morphology is determined by the degree of extension of the posterior mantle lobe in live animals. In Oliva the lobe is extending above the aperture on the parietal part of the last whorl towards the apex, but does not extend backwards, in the direction opposite to the shell growth above the aperture. Thus, the primary callus is not overlaying the suture on the last whorl. In Ancillariidae, it is obvious that the posterior lobe is extending above but also backwards to the posterior aperture corner, and is producing both primary and secondary spire calluses. This results in the suture overlayed with a layer of primary spire callus, producing the very characteristic ancillariid appearance of the shell. In some genera (Ancilla), the primary callus covers most of the shell surface and on upper whorls it is overlaying the callus deposited during the growth of the previous whorl, producing a multi-layered callused structure. The presence of a reduced posterior mantle lobe in Pseudolividae and Benthobiidae, which deposits the callus that extends only to the parietal part of the last whorl, may represent an initial stage of development of the primary callus.

We do not have at this moment a clue to the functional significance of such prominent character as the spire callus. An obvious hypothesisthat the spire callus renders the shell more smooth to facilitate burrowing and movements in the sedimentis rejected by the fact that in Olivoidea (except Pseudolividae and Benthobiidae) the shell is nearly completely covered by the foot parapodia and is not directly in contact with the sediment.

While the callus is deposited by a subtle, but rather distinct, lobe that can be seen in preserved specimens, the anterior band is deposited by the mantle reflected at the fasciole over the shell edge. This part of the mantle (anterior mantle lobe) does not differ morphologically from the adjoining parts of the mantle. Its role can be deduced only by observations of living animals. In some preserved specimens the anterior mantle lobe forms minor folds, probably corresponding to the plicae on the anterior band, but usually it is smooth.

It should be noted that a significant or even hypertrophied development of the callus is not unique to Olivoidea, but is known in other, not closely related, neogastropods. In some Nassariidae [e. g. Nassarius arcularia (Linnaeus, 1758)] at the termination of growth the parietal callus produces a very thick shield that extends over all the apertural side of the shell. In the fossil Cyllenina lucinensis (Landau & Marquet, 1999) from the Pliocene of Spain, the callus completely covers the shell, including the protoconch (Lozouet & Galindo, 2015). Although nothing is known about the mantle anatomy of such species, it is logical to suggest that the callus should be deposited by some equivalent of the posterior mantle lobe of Olivoidea, sometimes only at growth termination (N. arcularia) but sometimes during the entire life (C. lucinensis). In most Marginellidae the suture is overlaid by a thin callus, producing an appearance similar to Ancillariidae. Again there are no data on the mantle morphology of marginellids. In Marginellidae the mantle edge is, as in olivoideans, reflected over the siphonal notch and a thickened shell edge is present in most species along the lip and around the fasciole. Obviously, the extra callus layer here is deposited in a similar way as in olivoideans, although it is not as intricate. [START_REF] Sterba | Bemerkungen zur Schale der Olividae[END_REF]) and, later, Tursch & Greifeneder (2001) mentioned one more unusual character of oliviform shells: they considered the morphology of the suture in Oliva to be unique in Gastropoda since the whorls of the shell do not come into direct contact with each other, but are connected by a spiral ribbon which is a part of the primary callus on the parietal whorl of the aperture. Such a peculiarity is in fact present in all oliviform shells (except Bellolividae, which do not have a spire callus and, correspondingly, no posterior mantle lobe). The mantle filament, housed in the filament channel, is indeed a unique morphological character, present in most Olividae and all Bellolividae. Several possible functions of the filament have been suggested: a sensory organ, indicating the depths of burial in the sediment (B. Tursch, pers. com. to Y. Kantor); a defensive weapon [START_REF] Zeigler | Olive shells of the world[END_REF]; a "sense organ that detects danger from behind while the animal is buried in the sand" [START_REF] Vermeij | A natural history of shells[END_REF], and others. None of these have been evaluated and remain conjectural.

Olivoidea (except Pseudolividae and Benthobiidae) have a very characteristic foot morphology. In crawling animals, the foot is large, very agile, and with a large crescent-shaped propodium, often free on external corners, and a metapodium with two large symmetrical lateral flaps, the parapodia. The propodium is subdivided by a longitudinal furrow. A similar shape of the propodium is also seen in Harpidae; however, it is not subdivided by a longitudinal cleft.

The unique propodial and metapodial morphology of olivoideans can be explained from a functional point of view. Thus the propodium is actively used for collecting food and acts like a pair of pincers, firmly seizing the prey when both halves of the propodium fold along the longitudinal furrow. The broad, thin and agile metapodium of Oliva and Agaronia serves to form the pouch in which the prey (sometimes several items according to our observations) is kept and digested. The disappearance of the operculum may be related with this ability to form a pouch by facilitating the bending of the metapodium. In Ancillariidae and Olivellidae the foot is truncated and relatively thicker, and seems to be unable to form a pouch. Correspondingly an operculum usually persists in these taxa. Some olivids can swimprobably as an escape responseusing either the propodium (as observed by [START_REF] Wilson | Use of the propodium as a swimming organ in an ancillid (Gastropoda: Olividae)[END_REF] in Ancillista cingulata (Sowerby, 1830)), or more often the parapodia (in Olivellasee [START_REF] Olsson | Studies of the genus Olivella[END_REF][START_REF] Marcus | Studies on "Olividae[END_REF]. During fieldwork at King Leopold Station in Laing Island (Papua-New Guinea), the senior author has observed swimming in many species of Oliva kept in aquaria in response to food. The propodium is also definitely used as a wedge during burrowing.

Parapodia embrace the shell in crawling snails, sometimes nearly completely. Observations were reported by [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF] on living Olivella borealis in a narrow aquarium that allowed the snail to be seen burrowed in the sediment. In the completely burrowed animal the parapodia are even more expanded than when the snail crawls on the surface of the sediment. We suggest that the lateral embrace of the shell prevents sand particles from entering the mantle cavity and thus is an adaptation to the burrowing mode of life.

Our results demonstrate that morphological data alone should be used with caution for phylogenetic reconstructions. The morphology-based phylogeny of Olivoidea [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF] was fundamentally different (and correspondingly wrong) in regards both rang of families and their relationships. For example, the radula that is otherwise considered to be of fundamental importance in the taxonomy of Neogastropoda is extremely variable within the one family Olividae, with a range of variation larger than within the entire superfamily. Similarly, the superfamily includes representatives with "normal" foot (narrow propodium, metapodium without parapodia in two families, Pseudolividae and Benthobiidae) and others with the characteristic "olivoidean" foot (crescent shaped propodium, metapodium with broad parapodia), which was hitherto considered one of the key autapomorphies for the group. with Maximal Likelihood analysis of COI, 16S rRNA, 12S rRNA and H3 genes. Only supported nodes are marked on the tree: squaresbootstrap values between 80 and 89 (supported node), circlesbootstrap values between 90 and 100 (highly supported node).

Figure 2. Phylogenetic tree of olivoidean gastropods and outgroup neogastropod taxa obtained with Bayesian analysis of COI, 16S rRNA, 12S rRNA and H3 genes. Only supported nodes are marked on the tree (squaresposterior probability values between 0.95 and 0.97, circlesposterior probability values between 0.98 and 1.00). The letters in brackets after the species names refer to photographs of vouchers in Figs 3 and4. Anterior band is strongly shagreened and its rear edge reaches the lip. D -Entomoliva mirabilis, east coast of New Caledonia, BATHUS1, st. CP713, 21°45'S, 166°37'E, 250 m. Anterior band is strongly shagreened and its rear edge reaches the lip. E-Flateral views of the tip and upper part of aperture of Oliva spp. E -Oliva miniacea, MNHN IM-2007-31973. The primary spire callus is not overlaying the filament channel. F -Oliva bulbiformis Duclos, 1835, MNHN IM-2007-31996. The primary callus overlaying the filament channel, which remains free along last whorl. The anterior border of the callus is marked by a grey line. G-Nphotographs of shells and interpretative drawings of the same specimens to show the position of the calluses. G-J -Entomoliva mirabilis, east coast of New Caledonia, BATHUS1, st. CP713, 21°45'S, 166°37'E, 250 m. K-N -Amalda fuscolingua MNHN IM-2007-43649. For collection data of registered specimens, see Table 1. Abreviations and conventions: olivoid groove. rear edge of anterior band. ABanterior band. fcfilament channel. pcprimary spire callus. scsecondary spire callus. Bouchet, 2007). Head-foot, dorsal view, mantle and visceral mass removed. G-H -Amalda aureomarginata, New Caledonia, Iles des Pins, st. CP3115, 22°48'S, 167°15'E, 440-470 m; Gmantle complex, Henlarged bipartitioned mantle lobe, dashed line shows the border of upper, flexible and reflectable partition (lobe). I -Olivella borealis Golikov, 1967, Japan Sea, Vostok Bay, 5-6 m. Mantle complex, mantle filament partly removed (after [START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF]. J -Oliva mantichora, Mozambique, Inhaca Island, INHACA 2011, st. MB5, 26°01.0'S, 32°54.1'E, 6 m. Photograph of the mantle filament and mantle lobe. Abbreviations: amlanterior mantle lobe; amtanterior mantle tentacle; ceptcephalic tentacle; cmcolumellar muscle; ctctenidium; fgfemale gonoduct; flcephalic flap; hghypobranchial gland; mfmantle filament; mlmantle lobe; mlppart of mantle lobe, producing primary callus; mlspart of mantle lobe, producing secondary callus; ososphradium; parparapodia; prproboscis seen through the body walls; prppropodium; rerectum; rsrhynchostome; ssiphon.

  the exclusively fossil Vanpalmeriidae Agedoke, 1977. [The family name Dactylidae H. & A. Adams, 1853, is invalid albeit available (ICZN, Art. 10.6) since it was based on the genus Dactylus H. & A. Adams, 1853, a junior homonym of Dactylus Schumacher, 1817 (Gastropoda: Acteonidae)].

  analyzed together with five COI sequences of other Olivoidea, used as outgroups (MNHN IM-2009-25009 Entomolliva mirabilis, MNHN IM-2007-31956 Ancillina cf. sumatrana, MNHN IM-2009-11968 Amalda hilgendorffi richeri, MNHN IM-2013-18534 Benthobia sp. and MNHN IM-2013-9727 Fusulculus crenatus). Similarly, a dataset including 56 original COI sequences of Oliva, together with two sequences from Genbank and the same outgroups, was analyzed for species delimitation. The web version (http://wwwabi.snv.jussieu.fr/public/abgd/abgdweb.html)

  Science Museum of Korea, Daejeon, Korea; BAU -Department of Animal and Human Biology, University of Rome La Sapienza, Rome, Italy; MNHN -Muséum National d'Histoire Naturelle, Paris, France; MZUSP -Museu de Zoologia, University of São Paulo, SP, Brasil; NMSA -Natal Museum, Pietermaritzburg, South Africa.

Fig. 9

 9 Fig.91F, Fig.10 2 A-B, C, H), often with additional smaller denticles or serration between the major cusps (Fig.13 D, E), to truly multicuspid (Figs 10 G, 17 J). The shape of the teeth can be very conservative within the genus; thus, in all Belloliva, there are two small additional denticles situated at the outer edge of the lateral cusps of the central tooth (Fig.9B); similarly, very distinct cusps are present in Agaronia (Fig.9 E) and Olivancillaria[START_REF] Teso | A revision of the genus Olivancillaria (Mollusca: Olividae) from the southwestern Atlantic[END_REF].

  the following way: (1) genera placed in the family on the basis of molecular and morphological data; (2) genera placed on the basis of shell and anatomical data; (3) genera tentatively attributed to the family based on shell characters alone. CLASS GASTROPODA CUVIER, 1795 SUBCLASS CAENOGASTROPODA COX, 1960 ORDER NEOGASTROPODA WENZ, 1938 SUPERFAMILY OLIVOIDEA LATREILLE, 1825

  Olivina d'Orbigny, 1841'Orbigny, (in 1834'Orbigny, -1847)). Type species: Oliva tehuelchana d'Orbigny, 1839; SD,[START_REF] Weinkauff | Catalog der Gattung Olivella Swainson[END_REF]. Orbignytesta Klappenbach, 1962. Type species: Olivella formicacorsii Klappenbach, 1962; OD. Pachyoliva Olsson, 1956. Type species: Oliva columellaris G.B. Sowerby I, 1825; OD. Parolivella Paulmier, 2007. Type species: Olivella marginelloides Paulmier, 2007; OD. Zanoetella Olsson, 1956. Type species: Oliva zanoeta Duclos, 1835; OD. SUFAMILY CALYPTOLIVINAE, subfam. nov. Type genus Calyptoliva Kantor & Bouchet, 2007. Diagnosis Plication plate either smooth or with weak spiral plicae that extend to columella. Primary callus very thin, overlaying suture. Anterior band very thin, hardly discernible, very slightly raised above the shell cloak, uniform. Filament channel absent. Anterior mantle tentacle absent. Posterior mantle lobe moderately or well developed and without anterior mantle tentacle. Head with small, broadly spaced, dorso-ventrally compressed flaps; eyes present or absent. Operculum present. Radula with 3 teeth per transverse row; central tooth medium broad, tricuspid. Lateral teeth hook-shaped. Stomach with narrow and long posterior mixing area.

  Olivancillaria d'Orbigny, 1841 (in 1834-1847) (2). Type species: Oliva auricularia Lamarck, 1811; M. Synonyms Scaphula Swainson, 1840 (preoccupied by Scaphula Benson, 1834 [Bivalvia]). Type species: Oliva patula G.B. Sowerby I, 1825; M. Olivancillaria (Lintricula) H. Adams & A. Adams, 1853 [nov. pro Scaphula Swainson, 1840, non Benson, 1834]. Type species: Oliva patula G.B. Sowerby I, 1825; by typification of replaced name. Claneophila Gray, 1858. Type species: † Oliva claneophila Duclos, 1835; by tautonymy.

FAMILY

  BELLOLIVIDAE fam. nov. Type genus: Belloliva Peile, 1922. Diagnosis Shell glossy, lacking periostracum, narrowly fusiform, with high to very high last whorl and narrow aperture tapering adapically. Siphonal canal absent, anterior end of shell distinctly notched. Anterior shell end with poorly defined anterior band, very slightly raised above the shell cloak. Plication plate limited to columella, with spiral plicae (Belloliva) or not (Olivellopsis). Primary spire callus absent. Narrow filament channel present, opened on all spire whorls. Foot with well-developed crescent-shaped propodium, subdivided by a longitudinal cleft on the dorsal side, and parapodia partially embracing the shell. Operculum present. Radula with three teeth per transverse row. Central tooth with three large, closely spaced cusps and, in Belloliva, with distinct small cusps adjoining the major lateral cusps (Fig. 9 B), that are indistinct or absent in Olivellopsis (Fig. 9 C). Mantle with mantle filament, a small anterior mantle lobe, and without anterior mantle tentacle. Head formed by dorso-ventrally compressed flaps with or without eyes, at least in Olivellopsis on very short and narrow tentacle.Genus BellolivaPeile, 1922 (1). Type species: Olivella brazieri Angas, 1877; OD Synonym: Belloliva (Gemmoliva)[START_REF] Iredale | Results from Roy Bell's molluscan collections[END_REF]. Type species: Oliva triticea Duclos, 1835; OD.Genus OlivellopsisThiele, 1929 (1). Type species: Olivella (Callianax) simplex Pease 1868; M. Synonym: Olivella (Janaoliva)Sterba & Lorenz, 2005, syn. nov. Type species: Olivella amoni[START_REF] Sterba | Olivella (Janaoliva) amoni, a new subgenus and species from the Bismark Archipelago and Indonesia (Mollusca: Gastropoda: Olivellidae)[END_REF] OD

  [START_REF] Adams | The genera of Recent Mollusca arranged according to their organisation[END_REF] Possible synonym: Vanpalmeriidae[START_REF] Adegoke | Stratigraphy and paleontology of the Ewekoro Formation (Paleocene) of southwestern Nigeria[END_REF]. Type genus: Vanpalmeria Adegoke, 1977. Monotypical family based on Vanpalmeria africana[START_REF] Adegoke | Stratigraphy and paleontology of the Ewekoro Formation (Paleocene) of southwestern Nigeria[END_REF], from the Paleocene of Nigeria.Type genus: AncillariaLamarck, 1811 [= Ancilla Lamarck, 1799] Diagnosis Shell glossy or mat, lacking periostracum, fusiform to narrowly fusiform, with high last whorl and medium broad to narrow aperture tapering adapically. Siphonal canal absent, anterior end of shell distinctly notched. Anterior shell end with well-defined anterior band, raised above the shell cloak and often strongly shagreened. Olivoid groove present (at least in some species) in all genera. Plication plate limited to columella, usually with spiral plicae. Primary spire callus well defined, covering most of, or even completely, the shell. Secondary spire callus from poorly defined to very strong. Suture always overlaid by the callus.Foot with well developed crescent-shaped propodium, subdivided by longitudinal cleft on dorsal side, and with parapodia partially or completely embracing the shell. Operculum usually present (rarely absent). Radula with three teeth per transverse row. Central tooth tricuspid or multicuspid, lateral teeth mostly simple hook-shaped, sometimes with additional serration along the inner edge (Ancillina, Fig.10 G) or bifurcated on the top (Ancilla atimovataeKantor et al., 2016 -Fig. 17 J-L). Mantle without mantle filament, with weak anterior mantle lobe, without anterior mantle tentacle, with very well developed posterior mantle lobe, sometimes subdivided in two lobes. Head formed by dorso-ventrally compressed flaps with or without eyes.

  ) Alocospira Cossmann, 1899. Type species: Ancillaria papillata Tate, 1889; OD. Austrancilla Habe, 1959. Type species: Ancilla edithae Pritchard & Gatliff, 1899; OD. Baryspira P. Fischer, 1883. Type species: Ancillaria australis G.B. Sowerby I, 1830; SD, Cossmann (1899). Exiquaspira Ninomiya, 1988. Type species: Amalda ornata Ninomiya, 1988; OD. Gracilispira Olsson, 1956. Type species: Ancillaria novaezelandiae G.B. Sowerby II, 1859; OD.Mundaspira[START_REF] Ninomiya | A new subgenus and five new species of the Ancillinae (Gastropoda: Olividae) from Australia[END_REF]. Type species: Amalda concinna[START_REF] Ninomiya | A new subgenus and five new species of the Ancillinae (Gastropoda: Olividae) from Australia[END_REF] OD. Pinguispira Finlay, 1926. Type species: Ancilla opima Marwick, 1924; OD. Genus Eburna Lamarck, 1801 (2). Type species: Eburna flavida Lamarck, 1801 [= Buccinum glabratum Linnaeus, 1758]; M.

  PseudolivaSwainson, 1840 (1). Type species: Buccinum plumbeum Dillwyn, 1817 [= Buccinum crassum Gmelin, 1791]; OD. Synonyms Mariona G.B. Sowerby III, 1890. Type species: Pseudoliva ancilla Hanley, 1859; M. Sylvanocochlis Melvill, 1903. Type species: Pseudoliva ancilla Hanley, 1859; OD; Invalid: junior objective synonym of Mariona. Fulmentum P. Fischer, 1884. Type species: Buccinum sepimentum Rang, 1832; M.

  Zemira H. Adams & A. Adams, 1853 (2). Type species: Eburna australis G.B. Sowerby I, 1833; M.

Figure 1 .

 1 Figure 1. Phylogenetic tree of olivoidean gastropods and outgroup neogastropod taxa obtainedwith Maximal Likelihood analysis of COI, 16S rRNA, 12S rRNA and H3 genes. Only supported nodes are marked on the tree: squaresbootstrap values between 80 and 89 (supported node), circlesbootstrap values between 90 and 100 (highly supported node).

Figure 3 .

 3 Figure 3. Shells of sequenced species. Conspecific specimens of sequenced vouchers (in cases of poor preservation of the voucher) are marked with asterisk. Shells not to scale. For collection data of registered specimens, see Table 1. A -Belloliva exquisita, MNHN IM-2007-34786, SL 7.2 mm. B* -Olivellopsis cf. amoni, Papua New Guinea, Kavieng Lagoon, S coast of Baudison I., KAVIENG 2014, st. KB20 02°45,2'S, 150°41,7'E, 8 m, SL 3.4 mm. C* -Olivellopsis amoni, Papua New Guinea, Kavieng Lagoon, NE corner of Nusa I., KAVIENG 2014, st. KS19, 02°34'S, 150°47,1'E, 10 m SL 3.9 mm. D* -Fusulculus crenatus, MNHN IM-2007-34041, Solomon Islands, SALOMON 2, st. CP2213, 7°38'42''S; 157°42'53.9856''E, 495-650 m, SL 20.6 mm. E -Benthobia sp., MNHN IM-2009-31002, SL 10.3 mm. F -Calyptoliva bbugeae, MNHN IM-2007-39266, SL 8.2 mm. G -Agaronia acuminata, MNHN IM-2013-52002, SL 25.3 mm; H -Agaronia annotata, MNHN IM-2013-52004, SL 18.0 mm. I -Olivella minuta, MNHN IM-2009-24350, SL 9.1 mm. J -Olivella olssoni, MNHN IM-2009-24370, SL 5.4 mm. K -Olivella exilis, MNHN IM-2009-24367, SL 6.6 mm. L -Oliva lacanientai, MNHN IM-2007-33275, SL 16.4 mm. M -Pseudoliva sepimenta, MNHN IM-2013-52005, SL 8.5 mm.

Figure 5 .

 5 Figure 5. Interpretative drawings and photos of the anterior end of the shell of actual specimens in different Olivoidea, showing terminology and proposed homologies.Homologous areas are similarly tiled. Olivadrawing after Tursch and Greifeneder (2001), shell of Oliva caroliniana, MNHN IM-2009-15462. Olivelladrawing of O. (Niteoliva) nitidula (after Olsson, 1956), shell of Olivella minuta, MNHN IM-2009-24350. Agaronia -A. acuminata, MNHN IM-2013-52002. Bellolivadrawing after Kantor and Bouchet (2007), shell -B. exquisita, MNHN IM-2007-34786. Ancilladrawing redrawn after Kilburn (1981), shell -Ancilla thomassini, MNHN IM-2000-15460. Turrancilla -Turrancilla sp. 2, MNHN IM-2007-31954, Philippines, the olivoid groove (depicted by dashed line) is present in this species, but absent in other studied ones. Amaldadrawing modified from Kilburn (1977), shell -Amalda aureomarginata, MNHN IM-2007-43655. Eburna -E. glabrata (Linnaeus, 1758), Venezuela, YK collection. Abbreviations and conventions: olivoid groove. rear edge of anterior band. fasciolar ridge. rear edge of plication plate. ABanterior band. CLlast whorl cloak. fufalse umbilicus. lablower anterior band. ldlabral denticle. obolivoid band. PPplication plate. uabupper anterior band.

Figure 6 .

 6 Figure 6. Details of the shell morphology of Olivoidea. A-Ddorso-anterior views of the anterior band. A -Oliva miniacea, MNHN IM-2007-31973.An arrow indicated the place, where the anterior band is merging with the siphonal notch callus. B -Amalda fuscolingua, MNHN IM-2007-43649. Rear edge of the anterior band nearly reaches the lip. C -Turrancilla sp. 1, Solomon Islands, SW Choiseul Island, SALOMON 2, st. CP2212, 7°38'S, 157°42'E, 400-475 m. Anterior band is strongly shagreened and its rear edge reaches the lip. D -Entomoliva mirabilis, east coast of New Caledonia, BATHUS1, st. CP713, 21°45'S, 166°37'E, 250 m. Anterior band is strongly shagreened and its rear edge reaches the lip. E-Flateral views of the tip and upper part of aperture of Oliva spp. E -Oliva miniacea, MNHN IM-2007-31973. The primary spire callus is not overlaying the filament channel. F -Oliva bulbiformis Duclos, 1835, MNHN IM-2007-31996. The primary callus overlaying the filament channel, which remains free along last whorl. The anterior border of the callus is marked by a grey line. G-Nphotographs of shells and interpretative drawings of the same specimens to show the position of the calluses. G-J -Entomoliva mirabilis, east coast of New Caledonia, BATHUS1, st. CP713, 21°45'S, 166°37'E, 250 m. K-N -Amalda fuscolingua MNHN IM-2007-43649. For collection data of registered specimens, see Table 1. Abreviations and conventions: olivoid groove. rear edge of anterior band. ABanterior band. fcfilament channel. pcprimary spire callus. scsecondary spire callus.

Figure 7 .

 7 Figure 7. Live animals of different Olivoidea. A. Oliva tesselata Lamarck, 1811; Papua-New Guinea, Kavieng Lagoon, Nago Island, KAVIENG 2014, sta. KR06, 02°36,3'S, 150°46,2'E, 3-12 m, MNHN IM-2013-46528 (Photo L. Charles). B-C. Oliva miniacea, Papua-New Guinea, Kavieng Lagoon, Nago Island, KAVIENG 2014, st. KD01, 02°37,5'S, 150°46,5'E, MNHN IM-2013-47178 (Photo L. Charles); Benlarged anterior end of crawling snail to show anterior mantle tentacle and anterior mantle lobe, forming the anterior band. D -Oliva sericea, Philippines, Panglao Island (photo S. Tagaro), crawling animal, ventral view of the foot to show the foot pouch. E -Olivella biplicata (Sowerby, 1825), Bodega Bay, California, USA (after Troost et al., 2012, photocourtesy Winfried S. Peter. F -Olivella semistriata (Gray, 1839), Playa Grande, Costa Rica (after Troost et al., 2012, photo W.S. Peters). The species possess very long free propodial tips (marked by an arrow), that form the mucus net during feeding. G -Agaronia acuminata, Senegal, Dakar, Dakar'09, st. 18, 7-12 m (photo P. Maestrati). H -Amalda contusa, Mozambique, Inhaka Island, MNHN IM-2009-22263. I -Ancillina sp., Philippines, AURORA 2007, st. CP2730, 358-378 m, 15°21'N, 121°34'E (photo B. Buge). J -Olivellopsis amoni, Papua-New Guinea, Kavieng, MNHN IM-2013-51055 (Photo L. Charles). K -Ancilla ventricosa, MNHN IM-2009-15537 (photo P. Maestrati). Abbreviations: amlanterior mantle lobe; amtanterior mantle tentacle; ceptcephalic tentacle; fpfoot pouch; mfmantle filament; parparapodia; prppropodium; ssiphon.

Figure 8 .

 8 Figure8. Mophology of head-foot and mantle of different Olivoidea. A -Oliva bulbosa(Röding, 1798). anterior view of the head and propodium, mantle removed (after[START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF]. B -Entomoliva mirabilis, New Caledonia, EXBODI, st. DW3787, 22°13'S, 167°06'E, 223-249 m. Anterior view of the head-foot, mantle removed. C-D -Pseudoliva sepimenta, Congo, MNHN IM-2013-52003. C -Anterior view of the head-foot, mantle removed; Dmantle, internal view. E -Agaronia acuminata, Senegal, Casamance, MNHN. Anterior view of the head-foot, mantle removed. F -Belloliva alaos Kantor & Bouchet, 2007, North of New Caledonia. MUSORSTOM 4, st. DW160, 18°42'S, 163°13'E, 668 m (afterKantor & Bouchet, 2007). Head-foot, dorsal view, mantle and visceral mass removed. G-H -Amalda aureomarginata, New Caledonia, Iles des Pins, st. CP3115, 22°48'S, 167°15'E, 440-470 m; Gmantle complex, Henlarged bipartitioned mantle lobe, dashed line shows the border of upper, flexible and reflectable partition (lobe). I -Olivella borealisGolikov, 1967, Japan Sea, Vostok Bay, 5-6 m. Mantle complex, mantle filament partly removed (after[START_REF] Kantor | On the morphology and relationships of some oliviform gastropods[END_REF]. J -Oliva mantichora, Mozambique, Inhaca

Figure 9 .

 9 Figure 9. Radulae of Bellolividae (A-C) and Olividae (E-H).A, B -Belloliva exquisita, Northern New Caledonia, MUSORSTOM 4, st. DW 184, 19°04'S, 163°27'E, 260 m (after Kantor & Bouchet, 2007). C -Olivellopsis amoni, Papua New Guinea, Kavieng Lagoon NE corner of Nusa I., KAVIENG 2014, sta. KS19, 02°34'S, 150°47,1'E, 10 m. D -Calyptoliva bbugea, holotype, MNHN IM-2007-38919. E -Agaronia acuminata, MNHN IM-2013-52002, central tooth. F -Oliva rufofulgurata Schepman, 1904, MNHN IM-2007-36132. G -Olivella minuta, MNHN IM-2009-24350. H -Olivella exilis, MNHN IM-2009-24367.

Figure 10 .

 10 Figure 10. Radulae of Pseudolividae (A-B) and Ancillariidae (C-I). A -Pseudoliva sepimenta, MNHN IM-2013-52005. B -Pseudoliva crassa (Gmelin, 1791), Baia do Baba, Moçâmedes, Angola, 6-20 m, coll. S. Gofas, SL 39.5 mm, AL 31.5 mm. C -Entomoliva mirabilis New Caledonia: Canal de la Havannah, EXBODI, st. DW3787, 22°13'S, 167°06'E, 223-249 m, SL 11.2 mm. D -Amalda contusa, MNHN IM-2009-22264. E, F -Amalda cf hilgendorfi richeri, MNHN IM-2007-43661, Fenlarged lateral tooth to show bifurcation on the tip. G -Ancillina cf sumatrana, Madagascar, between Majunga and Cap Saint-André, MIRIKY, st. CP3278, 15°24'S, 45°56'E, 750-780 m. H -Turrancilla sp. 2, MNHN IM-2007-31947. I -Eburna lienardi (Bernardi, 1859), MZSP 40140, Proc. Areia Branca, RN, Brazil, 04°45'S, 36°58'W, 9 m.

Figure 11 .

 11 Figure 11. Phylogenetic tree of Oliva and outgroup olivoidean taxa obtained with Bayesian analysis of COI gene sequences. Only supported nodes with posterior probability values between 0.98 and 1.00 are marked on the tree with circles. The names of the corresponding subgenera (after Petuch & Sargent, 1986 and Hunon et al., 2009) are given after the species names. Abbreviations of the subgenera: Ac -Acutoliva; An -Annulatoilva; C -Carmione; G -Galeola; Mi -Miniaceoliva; Mu -Musteoliva; O -Omogymna; Ol -Oliva; P-Parvoliva; R -Rufoliva.

Figure 12 .

 12 Figure 12. Phylogenetic tree of Ancilla and outgroup olivoidean taxa obtained with Bayesian analysis of COI gene sequences. Only supported nodes with posterior probability values between 0.98 and 1.00 are marked on the tree with circles. (A) -Ancilla aureocallosa, MNHN IM-2007-39045, SL 21.7 mm. (B) -Ancilla sp., MNHN IM-2007-36606, SL 5.6 mm. (C) -A. thomassini, MNHN IM-2009-15460, SL 6.3 mm. (D) -A. morrisoni, holotype, MNHN IM-2013-55401, SL 10.0 mm. (E) -A. ventricosa, MNHN IM-2009-15544, SL 10.7 mm. (F) -A. cf. giaquintoi, MNHN IM-2009-15526. (G) -A. adelphe, MNHN IM-2009-15519, SL 11.1 mm. (H) -A. sp., MNHN IM-2009-15527, SL 12.7 mm. (I) -A. lhaumeti, holotype, MNHN IM-2009-15513, SL 15.3 mm.

Figure 13 .

 13 Figure 13. Radulae of species of Ancilla. The shells of the specimens are depicted on Fig. 12 (except Ancilla cinamomea). (A) -Ancilla aureocallosa, MNHN IM-2007-39045, SL 21.7 mm. (B) -Ancilla sp., MNHN IM-2007-36606, SL 5.6 mm. (C) -A. thomassini, MNHN IM-2009-15460, SL 6.3 mm. (D) -A. morrisoni, holotype, MNHN IM-2013-55401, SL 10.0 mm. (E) -A. ventricosa, MNHN IM-2009-15544, SL 10.7 mm. (G) -A. adelphe, MNHN IM-2009-15519, SL 11.1 mm. (I) -A. lhaumeti, holotype, MNHN IM-2009-15513, SL 15.3 mm. (J) -A. cinamomea Lamarck, 1801, Southern India, Rameshwaran (after Kantor & Bouchet, 2007: fig. 30 B).

Figure 1 .

 1 Figure 1. Phylogenetic tree of Ancilla and outgroup olivoidean taxa obtained with Bayesian analysis of COI gene sequences. Only supported nodes with posterior probability values between 0.98 and 1.00 are marked on the tree with circles. (A) -Ancilla aureocallosa, MNHN IM-2007-39045, SL 21.7 mm. (B) -Ancilla sp., MNHN IM-2007-36606, SL 5.6 mm. (C) -A. thomassini, MNHN IM-2009-15460, SL 6.3 mm. (D) -A. morrisoni, holotype, MNHN IM-2013-55401, SL 10.0 mm. (E) -A. ventricosa, MNHN IM-2009-15544, SL 10.7 mm. (F) -A. cf. giaquintoi, MNHN IM-2009-15526. (G) -A. adelphe, MNHN IM-2009-15519, SL 11.1 mm. (H) -A. sp., MNHN IM-2009-15527, SL 12.7 mm. (I) -A. lhaumeti, holotype, MNHN IM-2009-15513, SL 15.3 mm. 167x139mm (300 x 300 DPI)

Figure 2 .

 2 Figure 2. Phylogenetic tree of Oliva and outgroup olivoidean taxa obtained with Bayesian analysis of COI gene sequences. Only supported nodes with posterior probability values between 0.98 and 1.00 are marked on the tree with circles. The names of the corresponding subgenera (after Petuch & Sargent, 1986 and Hunon et al., 2009) are given after the species names. Abbreviations of the subgenera: Ac -Acutoliva; An -Annulatoilva; C -Carmione; G -Galeola; Mi -Miniaceoliva; Mu -Musteoliva; O -Omogymna; Ol -Oliva; P-Parvoliva; R -Rufoliva. 167x199mm (300 x 300 DPI)

Figure 3 .

 3 Figure 3. Phylogenetic tree of olivoidean gastropods and outgroup neogastropod taxa obtained with Maximal Likelihood analysis of COI, 16S rRNA, 12S rRNA and H3 genes. Only supported nodes are marked on the tree: squares -bootstrap values between 80 and 89 (supported node), circles -bootstrap values between 90 and 100 (highly supported node).199x237mm (300 x 300 DPI)

Figure 4 .

 4 Figure 4. Phylogenetic tree of olivoidean gastropods and outgroup neogastropod taxa obtained with Bayesian analysis of COI, 16S rRNA, 12S rRNA and H3 genes. Only supported nodes are marked on the tree (squares -posterior probability values between 0.95 and 0.97, circles -posterior probability values between 0.98 and 1.00). The letters in brackets after the species names refer to photographs of vouchers in Figs 5 and 6. 199x238mm (300 x 300 DPI)

Figure 5 .

 5 Figure 5. Shells of sequenced species. Conspecific specimens of sequenced vouchers (in cases of poor preservation of the voucher) are marked with asterisk. Shells not to scale. For collection data of registered specimens, see Table 1. A -Belloliva exquisita, MNHN IM-2007-34786, SL 7.2 mm. B* -Olivellopsis cf. amoni, Papua New Guinea, Kavieng Lagoon, S coast of Baudison I., KAVIENG 2014, st. KB20 02°45,2'S, 150°41,7'E, 8 m, SL 3.4 mm. C* -Olivellopsis amoni, Papua New Guinea, Kavieng Lagoon, NE corner of Nusa I., KAVIENG 2014, st. KS19, 02°34'S, 150°47,1'E, 10 m SL 3.9 mm. D* -Fusulculus crenatus, MNHN IM-2007-34041, Solomon Islands, SALOMON 2, st. CP2213, 7°38'42''S; 157°42'53.9856''E, 495-650 m, SL 20.6 mm. E -Benthobia sp., MNHN IM-2009-31002, SL 10.3 mm. F -Calyptoliva bbugeae, MNHN IM-2007-39266, SL 8.2 mm. G -Agaronia acuminata, MNHN IM-2013-52002, SL 25.3 mm; H -Agaronia annotata, MNHN IM-2013-52004, SL 18.0 mm. I -Olivella minuta, MNHN IM-2009-24350, SL 9.1 mm. J -Olivella olssoni, MNHN IM-2009-24370, SL 5.4 mm. K -Olivella exilis, MNHN IM-2009-24367, SL 6.6 mm. L -Oliva lacanientai, MNHN IM-2007-33275, SL 16.4 mm. M -Pseudoliva sepimenta, MNHN IM-2013-52005, SL 8.5 mm. 167x184mm (300 x 300 DPI)

Figure 6 .

 6 Figure 6. Shell of sequenced species. Conspecific specimens of sequenced vouchers (in cases of poor preservation of the voucher) are marked with asterisk. Shells not to scale. For collection data of registered specimens, see Table 1. N -Amalda bellonarum, MNHN IM-2007-33273, SL 16.6 mm. O -Amalda contusa, MNHN IM-2009-22263, SL 24.9 mm. P -Amalda aureomarginata, MNHN IM-2007-43655, SL 31.5 mm. Q -Amalda cf hilgendorfi richeri, MNHN IM-2009-11963, SL 14.9mm. R -Amalda hilgendorfi richeri, MNHN IM-2009-11968, SL 32.5 mm. S* -Ancillina cf sumatrana, BIOPAPUA, st. CP3731, 07°50'S, 148°04'E, 895-1150 m, SL 7.3 mm. T* -Turrancilla sp. 2., MNHN IM-2007-31954, PANGLAO 2005, st. CP2360, 8°48'54''N; 123°37'36.0156''E, 357-372 m, SL 12.3 mm. U -Turrancilla sp., MNHN IM-2013-19425, SL 20.5 mm. V -Turrancilla glans, MNHN IM-2007-31949, SL 36.4 mm. W -Turrancilla sp. 1, MNHN IM-2007-33282, SL 34.1 mm. 167x157mm (300 x 300 DPI)

Figure 7 .

 7 Figure 7. Interpretative drawings and photos of the anterior end of the shell of actual specimens in different Olivoidea, showing terminology and proposed homologies. Homologous areas are similarly tiled. Olivadrawing after Tursch and Greifeneder (2001), shell of Oliva caroliniana, MNHN IM-2009-15462. Olivelladrawing of O. (Niteoliva) nitidula (after Olsson, 1956), shell of Olivella minuta, MNHN IM-2009-24350. Agaronia -A. acuminata, MNHN IM-2013-52002. Belloliva -drawing after Kantor and Bouchet (2007), shell -B. exquisita, MNHN IM-2007-34786. Ancilla -drawing redrawn after Kilburn (1981), shell -Ancilla thomassini, MNHN IM-2000-15460. Turrancilla -Turrancilla sp. 2, MNHN IM-2007-31954, Philippines, the olivoid groove (depicted by dashed line) is present in this species, but absent in other studied ones. Amalda -drawing modified from Kilburn (1977), shell -Amalda aureomarginata, MNHN IM-2007-43655. Eburna -E. glabrata (Linnaeus, 1758), Venezuela, YK collection. Abbreviations and conventions: -olivoid groove. -rear edge of anterior band. -fasciolar ridge.rear edge of plication plate. AB -anterior band. CL -last whorl cloak. fu -false umbilicus. lab -lower anterior band. ld -labral denticle. ob -olivoid band. PP -plication plate. uab -upper anterior band.

Figure 8 .

 8 Figure 8. Details of the shell morphology of Olivoidea. A-D -dorso-anterior views of the anterior band. A -Oliva miniacea, MNHN IM-2007-31973. An arrow indicated the place, where the anterior band is merging with the siphonal notch callus. B -Amalda fuscolingua, MNHN IM-2007-43649. Rear edge of the anterior band nearly reaches the lip. C -Turrancilla sp. 1, Solomon Islands, SW Choiseul Island, SALOMON 2, st. CP2212, 7°38'S, 157°42'E, 400-475 m. Anterior band is strongly shagreened and its rear edge reaches the lip. D -Entomoliva mirabilis, east coast of New Caledonia, BATHUS1, st. CP713, 21°45'S, 166°37'E, 250 m. Anterior band is strongly shagreened and its rear edge reaches the lip. E-F -lateral views of the tip and upper part of aperture of Oliva spp. E -Oliva miniacea, MNHN IM-2007-31973. The primary spire callus is not overlaying the filament channel. F -Oliva bulbiformis Duclos, 1835, MNHN IM-2007-31996. The primary callus overlaying the filament channel, which remains free along last whorl. The anterior border of the callus is marked by a grey line. G-N -photographs of shells and interpretative drawings of the same specimens to show the position of the calluses. G-J -Entomoliva mirabilis, east coast of New Caledonia, BATHUS1, st. CP713, 21°45'S, 166°37'E, 250 m. K-N -Amalda fuscolingua MNHN IM-2007-43649. For collection data of registered specimens, seeTable 1.

Figure 9 .

 9 Figure 9. Live animals of different Olivoidea. A. Oliva tesselata Lamarck, 1811; Papua-New Guinea, Kavieng Lagoon, Nago Island, KAVIENG 2014, sta. KR06, 02°36,3'S, 150°46,2'E, 3-12 m, MNHN IM-2013-46528 (Photo L. Charles). B-C. Oliva miniacea, Papua-New Guinea, Kavieng Lagoon, Nago Island, KAVIENG 2014, st. KD01, 02°37,5'S, 150°46,5'E, MNHN IM-2013-47178 (Photo L. Charles); B -enlarged anterior end of crawling snail to show anterior mantle tentacle and anterior mantle lobe, forming the anterior band. D -Oliva sericea, Philippines, Panglao Island (photo S. Tagaro), crawling animal, ventral view of the foot to show the foot pouch. E -Olivella biplicata (Sowerby, 1825), Bodega Bay, California, USA (after Troost et al., 2012, photo -courtesy Winfried S. Peter. F -Olivella semistriata (Gray, 1839), Playa Grande, Costa Rica (after Troost et al., 2012, photo W.S. Peters). The species possess very long free propodial tips (marked by an arrow), that form the mucus net during feeding. G -Agaronia acuminata, Senegal, Dakar, Dakar'09, st. 18, 7-12 m (photo P. Maestrati). H -Amalda contusa, Mozambique, Inhaka Island, MNHN IM-2009-22263. I -Ancillina sp., Philippines, AURORA 2007, st. CP2730, 358-378 m, 15°21'N, 121°34'E (photo B. Buge). J -Olivellopsis amoni, Papua-New Guinea, Kavieng, MNHN IM-2013-51055 (Photo L. Charles). K -Ancilla

Figure 10 .

 10 Figure 10. Mophology of head-foot and mantle of different Olivoidea. A -Oliva bulbosa (Röding, 1798). anterior view of the head and propodium, mantle removed (after Kantor, 1991). B -Entomoliva mirabilis, New Caledonia, EXBODI, st. DW3787, 22°13'S, 167°06'E, 223-249 m. Anterior view of the head-foot, mantle removed. C-D -Pseudoliva sepimenta, Congo, MNHN IM-2013-52003. C -Anterior view of the headfoot, mantle removed; D -mantle, internal view. E -Agaronia acuminata, Senegal, Casamance, MNHN. Anterior view of the head-foot, mantle removed. F -Belloliva alaos Kantor & Bouchet, 2007, North of New Caledonia. MUSORSTOM 4, st. DW160, 18°42'S, 163°13'E, 668 m (after Kantor & Bouchet, 2007). Headfoot, dorsal view, mantle and visceral mass removed. G-H -Amalda aureomarginata, New Caledonia, Iles des Pins, st. CP3115, 22°48'S, 167°15'E, 440-470 m; G -mantle complex, H -enlarged bipartitioned mantle lobe, dashed line shows the border of upper, flexible and reflectable partition (lobe). I -Olivella borealis Golikov, 1967, Japan Sea, Vostok Bay, 5-6 m. Mantle complex, mantle filament partly removed (after Kantor, 1991). J -Oliva mantichora, Mozambique, Inhaca Island, INHACA 2011, st. MB5, 26°01.0'S, 32°54.1'E, 6 m. Photograph of the mantle filament and mantle lobe. Abbreviations: aml -anterior mantle lobe; amt -anterior mantle tentacle; cept -cephalic tentacle; cmcolumellar muscle; ct -ctenidium; fg -female gonoduct; fl -cephalic flap; hg -hypobranchial gland; mfmantle filament; ml -mantle lobe; mlp -part of mantle lobe, producing primary callus; mls -part of mantle lobe, producing secondary callus; os -osphradium; par -parapodia; pr -proboscis seen through the body walls; prp -propodium; re -rectum; rs -rhynchostome; s -siphon. 170x172mm (300 x 300 DPI)

Figure 11 .

 11 Figure 11. Radulae of Bellolividae (A-C) and Olividae (E-H). A, B -Belloliva exquisita, Northern New Caledonia, MUSORSTOM 4, st. DW 184, 19°04'S, 163°27'E, 260 m (after Kantor & Bouchet, 2007). C -Olivellopsis amoni, Papua New Guinea, Kavieng Lagoon NE corner of Nusa I., KAVIENG 2014, sta. KS19, 02°34'S, 150°47,1'E, 10 m. D -Calyptoliva bbugea, holotype, MNHN IM-2007-38919. E -Agaronia acuminata, MNHN IM-2013-52002, central tooth. F -Oliva rufofulgurata Schepman, 1904, MNHN IM-2007-36132. G -Olivella minuta, MNHN IM-2009-24350. H -Olivella exilis, MNHN IM-2009-24367. 167x199mm (300 x 300 DPI)

Figure 12 .

 12 Figure 12. Radulae of Pseudolividae (A-B) and Ancillariidae (C-I). A -Pseudoliva sepimenta, MNHN IM-2013-52005. B -Pseudoliva crassa (Gmelin, 1791), Baia do Baba, Moçâmedes, Angola, 6-20 m, coll. S. Gofas, SL 39.5 mm, AL 31.5 mm. C -Entomoliva mirabilis New Caledonia: Canal de la Havannah, EXBODI, st. DW3787, 22°13'S, 167°06'E, 223-249 m, SL 11.2 mm. D -Amalda contusa, MNHN IM-2009-22264. E, F -Amalda cf hilgendorfi richeri, MNHN IM-2007-43661, F -enlarged lateral tooth to show bifurcation on the tip. G -Ancillina cf sumatrana, Madagascar, between Majunga and Cap Saint-André, MIRIKY, st. CP3278, 15°24'S, 45°56'E, 750-780 m. H -Turrancilla sp. 2, MNHN IM-2007-31947. I -Eburna lienardi (Bernardi, 1859), MZSP 40140, Proc. Areia Branca, RN, Brazil, 04°45'S, 36°58'W, 9 m. 167x199mm (300 x 300 DPI)

Figure 13 .

 13 Figure 13. Radulae of species of Ancilla. The shells of the specimens are depicted on Fig. 1 (except Ancilla cinamomea). (A) -Ancilla aureocallosa, MNHN IM-2007-39045, SL 21.7 mm. (B) -Ancilla sp., MNHN IM-2007-36606, SL 5.6 mm. (C) -A. thomassini, MNHN IM-2009-15460, SL 6.3 mm. (D) -A. morrisoni, holotype, MNHN IM-2013-55401, SL 10.0 mm. (E) -A. ventricosa, MNHN IM-2009-15544, SL 10.7 mm. (G) -A. adelphe, MNHN IM-2009-15519, SL 11.1 mm. (I) -A. lhaumeti, holotype, MNHN IM-2009-15513, SL 15.3 mm. (J) -A. cinamomea Lamarck, 1801, Southern India, Rameshwaran (after Kantor & Bouchet, 2007: fig. 30 B).167x199mm (300 x 300 DPI)

  No family group name being available for the clade uniting Belloliva and Olivellopsis, we propose for it the new family name Bellolividae fam. nov. [see Taxonomy part]. Similarly, no name being available for the clade including representatives of Benthobia and Fusulculus,previously (e. g.,[START_REF] Vermeij | Generic revision of the neogastropod family Pseudolividae[END_REF] included in Pseudolividae, we propose for it the new family Benthobiidae fam. nov. [see Taxonomy part].

  . Type species: Oliva panniculata Duclos, 1835; OD. Americoliva Petuch, 2013. Type species: Oliva sayana Ravenel, 1834; OD. Annulatoliva Petuch & Sargent, 1986. Type species: Oliva annulata Gmelin, 1791; OD. Arctoliva Petuch & Sargent, 1986. Type species: Oliva arctata Marrat, 1871; OD. Cariboliva Petuch & Sargent, 1986. Type species: Oliva scripta Lamarck, 1811; OD. Carmione Gray, 1858. Type species: Voluta ventricosa Dillwyn, 1817 [= O. bulbosa Galeola Gray, 1858. Type species: Oliva galeola Duclos, 1835; by tautonymy. [The identity of the type species is unclear. Gray (1858) placed it in the synonymy of Galeola avellana (Lamarck, 1811), a nomen dubium according to Tursch & Greifeneder (2001), who treated O. galeola as a sujective synonym of O. sericea (Röding, 1798), a species attributed to Miniaceoliva]. Miniaceoliva Petuch & Sargent, 1986. Type species: Oliva miniacea Röding, 1798; OD. Musteloliva Petuch & Sargent, 1986. Type species: Oliva mustelina Lamarck, 1811; OD. Neocylindrus P. Fischer, 1883. Type species: Oliva tessellata Lamarck, 1811; M.

	Lamarck, 1811]; M.
	Multiplicoliva Petuch & Sargent, 1986. Type species: Oliva multiplicata Reeve, 1850;
	OD.

Omogymna Martens, 1897. Type species: Oliva paxillus Reeve, 1850 [= O. nitidula Duclos, 1835]; M. Parvoliva Thiele, 1929. Type species: Oliva dubia Schepman, 1904; M. Porphyria Röding, 1798. Type species: Voluta porphyria Linnaeus, 1758; by tautonymy. Note. The name Strephona requires special mention, as it is broadly used in the literature (e.g. by

  , selecting Voluta porphyria Linnaeus, 1758 as type species.Rufoliva[START_REF] Petuch | Atlas of the living olive shells of the world[END_REF]. Type species: Oliva rufula Duclos, 1835; OD.Strephonella[START_REF] Dall | Contributions to the Tertiary paleontology of the Pacific coast. I. The Miocene of Astoria and Coos Bay, Oregon[END_REF]. Type species: Oliva undatella Lamarck, 1810; OD.DiagnosisPlication plate not distinctly subdivided, with distinct spiral plicae. Olivoid groove present, shallow. Olivoid band differing or not in colour from cloak of last whorl. Filament channel well defined, free on most spire whorls. Operculum absent. Anterior mantle tentacle well defined.Head with dorso-ventrally compressed flaps, eyes absent in all studied species. Radula with three teeth per transverse row, central tooth with very long lateral flaps and three large major cusps. Short but very distinct small denticles situated externally from lateral cusps, separated from them by deep narrow grooves.Anazola Gray, 1858. Type species: Oliva acuminata Lamarck; SD, Cossmann (1899).

	Genus Agaronia Gray, 1839 (1). Type species: Voluta hiatula Gmelin, 1791; M.
	Synonyms
	Hiatula Swainson, 1831 (Invalid: junior homonym of Hiatula Modeer, 1793 [Bivalvia]).
	Utriculina Gray, 1847. Type species: Voluta utriculus Gmelin, 1791 [= Voluta gibbosa
	Born,1778]; OD.
	Strephona Gray, 1847, is thus a junior objective synonym of Porphyria.
	Proxoliva Petuch & Sargent, 1986. Type species: Oliva caldania Duclos, 1835; OD [a
	nomen dubium formerly applied to O. bretingbami Bridgman, 1909, according to
	Tursch & Greifeneder, 2001].
	Viduoliva Petuch & Sargent, 1986. Type species: Oliva vidua Röding, 1798; OD.
	SUBFAMILY AGARONIINAE OLSSON, 1956
	Type genus: Agaronia Gray, 1839.

  had previously illustrated the radula of the type species, which proved its placement in Olivellidae]. Dactylidia H. Adams & A. Adams, 1853. Type species: Oliva nana Lamarck, 1811; SD, Weinkauff (1878: 124, as Dactyliola, an incorrect subsequent spelling). This type designation was overlooked and later Cossmann (1899: 54) designated Olivella mutica (Say, 1822) [an originally included species] as the type species. Olsson followed Cossmann, but did not include Oliva nana in his monograph. Therefore, the subgeneric position of O. nana as well as the composition of the subgenus remain unclear. Micana Gray, 1858. Type species: Oliva nana Lamarck, 1811; M. A junior objective synonym of Dactylidia. Dactylidella Woodring, 1928. Type species: Oliva anazora Duclos, 1835; OD. Lamprodoma Swainson, 1840. Type species: Oliva volutella Lamarck, 1811; M [=Ramola Gray, 1858, Type species: Oliva volutella Lamarck, 1811; M]. Macgintiella Olsson, 1956. Type species: Olivella watermani McGinty, 1940; OD.Minioliva[START_REF] Olsson | Studies of the genus Olivella[END_REF].Type species: Olivella perplexa[START_REF] Olsson | Studies of the genus Olivella[END_REF] OD.

Niteoliva

[START_REF] Olsson | Studies of the genus Olivella[END_REF]

. Type species: Porphyria minuta Link, 1807; OD.

  . Type species: Ancilla cinnamomea Lamarck, 1801; by subsequent monotypy. Synonyms Anaulax Roissy, 1805. Unnecessary substitute name for Ancilla Lamarck, 1799, by Roissy believed to be preoccupied by the genus "ancille" used by Geoffroy for freshwater limpets [= Ancylus]. Ancillaria Lamarck, 1811. Substitute name for Ancilla Lamarck, 1799, treated by Lamarck as a junior homonym of "Ancylus Geoffroy" [Ancylus O.F. Müller, 1773]. Ancillus Montfort, 1810. Unjustified emendation of Ancilla Lamarck, 1799. Under ICZN Art. 33.2, Ancillus is an objective synonym of Ancilla, and Montfort's designation of Ancilla buccinoides Lamarck, 1803, as type species of Ancillus is invalid. Sparellina P. Fischer, 1883. Type species: Ancilla candida Lamarck, 1811 [= Voluta ampla Ancillista Iredale, 1936. Type species: Ancillista velesiana Iredale, 1936; OD.

	Sparella Gray, 1857. Type species: Ancillaria ventricosa Lamarck, 1811; SD, Cossmann
	(1899).
	Gmelin, 1791]; M.
	Possibly valid (sub)genera
	Chilotygma H. Adams & A. Adams, 1853. Type species: Ancilla exigua G.B. Sowerby I,
	1830; M.
	Hesperancilla Kilburn, 1981. Type species: Ancilla matthewsi J.Q. Burch & R.L. Burch,
	1967; OD.

?

Table 1

 1 Calyptoliva bbugeae, MNHN IM-2007-39266, SL 8.2 mm. G -Agaronia acuminata, MNHN IM-2013-52002, SL 25.3 mm; H -Agaronia annotata, MNHN IM-2013-52004, SL 18.0 mm. I -Olivella minuta, MNHN IM-2009-24350, SL 9.1 mm. J -Olivella olssoni, MNHN IM-2009-24370, SL 5.4 mm. K -Olivella exilis, MNHN IM-2009-24367, SL 6.6 mm. L -Oliva lacanientai, MNHN IM-2007-33275, SL 16.4 mm. M -Pseudoliva sepimenta, MNHN IM-2013-52005, SL 8.5 mm.

	. A -Belloliva exquisita, MNHN IM-

Figure 4. Shell of sequenced species.

Conspecific specimens of sequenced vouchers (in cases of poor preservation of the voucher) are marked with asterisk. Shells not to scale. For collection data of registered specimens, see Table 1. N -Amalda bellonarum, MNHN IM-2007-33273, SL 16.6 mm. O -Amalda contusa, MNHN IM-2009-22263, SL 24.9 mm. P -

Table 1 .

 1 List of specimens analyzed.

	Family (Subfamily)	Genus Species	Inventory number	Expedition	Station	Locality	BOLD	COI	16S	12S	28S	Source
	Ancillariidae	Amalda aureomarginata Kilburn & Bouchet, 1988	IM-2007-43655	TERRASSES	DW3123	New Caledonia, Ille des Pins, 22°53'S, 167°13'E, 420-450 m						original sequence
	Ancillariidae	Amalda bellonarum Kilburn & Bouchet, 1988	IM-2007-33273	EBISCO	CP2572	New Caledonia, North Bellona, 20°23'S, 158°45'E, 324-330 m
	Ancillariidae	Amalda cf hilgendorfi richeri Kilburn & Bouchet, 1988	IM-2009-11963	NORFOLK 2	DW2147	New Caledonia, Ille des Pins, 22°50'S,167°16'N, 496 m						original sequence
	Ancillariidae	Amalda contusa (Reeve, 1864)	IM-2009-22263	INHACA 2011	MR4	Mozambique, 26°6'18''S; 32°58''E, 17-19 m						original sequence
	Ancillariidae	Amalda fuscolingua Kilburn & Bouchet, 1988	IM-2007-43649	TERRASSES	DW3122	New Caledonia, Ille des Pins, 22°47'S, 167°12'E, 390-410 m						original sequence
	Ancillariidae	Amalda hilgendorffi richeri Kilburn & Bouchet, 1988	IM-2009-11968	CONCALIS	DW2979	Northern New Caledonia, 18°16'S, 162°54'E, 350 m						original sequence
	Ancillariidae	Ancilla adelphe Kilburn, 1981	IM-2009-15475	ATIMO VATAE	BM06	South Madagascar, 25°27'55''S, 44°57'38"E, 0-1 m
	Ancillariidae	Ancilla adelphe Kilburn, 1981	IM-2009-15491	ATIMO VATAE	BM07	South Madagascar, 25°27'55''S, 44°57'38"E, 0-1 m
	Ancillariidae	Ancilla adelphe Kilburn, 1981	IM-2009-15510	ATIMO VATAE	BV16	South Madagascar, 25°25'33''S, 44°55'50.9953"E
	Ancillariidae	Ancilla adelphe Kilburn, 1981	IM-2009-15519	ATIMO VATAE	BP42	South Madagascar, 25°22.8'S, 44°51.1'E, 18-20 m						original sequence
	Ancillariidae	Ancilla adelphe Kilburn, 1981	IM-2009-15524	ATIMO VATAE	BV16	South Madagascar, 25°25'33''S, 44°55'50"E
	Ancillariidae	Ancilla adelphe Kilburn, 1981	IM-2009-15535	ATIMO VATAE	BP41	South Madagascar, 25°22.9'S, 44°51.0-51.6'E, 19-21 m						original sequence
	Ancillariidae	Ancilla adelphe Kilburn, 1981	IM-2009-15538	ATIMO VATAE	BP36	South Madagascar, 25°21'52''S, 44°50'13''E, 10-17 m
	Ancillariidae	Ancilla atimovatae Kantor et al., 2016	IM-2009-15511	ATIMO VATAE	BS01	South Madagascar, 25°28.2'S, 44°56.4'E, 12-14 m						original sequence
	Ancillariidae	Ancilla aureocallosa (Kilburn & Jenner, 1977)	IM-2007-39045	MAINBAZA	CP3143	Mozambique Channel, 23°32'S, 35°46'E, 264-277 m						original sequence
	Ancillariidae	Ancilla cf giaquintoi Bozzetti, 2006	IM-2009-15526	ATIMO VATAE	BP22	South Madagascar, 25°23'26.5164''S, 44°51'39.42''E, 20-22 m
	Ancillariidae	Ancilla kaviangiensis Kantor et al., 2016	IM-2013-51964	KAVIENG 2014	KR82	Papua-New Guinea, 02°39,6'S, 150°39,9'E, 2-12 m						original sequence
	Ancillariidae	Ancilla kaviangiensis Kantor et al., 2016	IM-2013-55102	KAVIENG 2014	KS57	Papua-New Guinea, 02°38,9'S, 150°40,1'E, 7-9 m						original sequence
	Ancillariidae	Ancilla kaviangiensis Kantor et al., 2016	IM-2013-55130	KAVIENG 2014	KB62	Papua-New Guinea, 02°36,4'S, 150°42,4'E, 19 m						original sequence
	Ancillariidae	Ancilla kaviangiensis Kantor et al., 2016	IM-2013-55138	KAVIENG 2014	KD70	Papua-New Guinea, 02°38,3'S, 150°45,6'E, 10-15 m						original sequence
	Ancillariidae	Ancilla lhaumeti Kantor et al., 2016	IM-2009-15481	ATIMO VATAE	BP21	South Madagascar, 25°23.1'S , 44°51.4'E, 20-23 m

Table 2 .

 2 Alignments and substitution models data

		Primers	Reference	Annealing temperature	Amplification length bp	Alignment length bp	N of sequences	substitution (hLRTs) model	I	gamma
	Fw: HCOI-1490 GGTCAACAAATCATAAAGAYATGYG	Folmer et al. (1994)	48-50	658	658	53	GTR+I+G	0,48	0,39
	Rv: LCOI-2198 TAAACTTCAGGGTGACCAAARAAYCA	Folmer et al. (1994)
	Fw: 16SH	CCGGTCTGAACTCAGATCACG	Palumbi (1996)	55	~550	577	52	TVM+I+G	0,16	0,26
	Rv: 16LC	GTTTACCAAAAACATGGCTTC	Palumbi (1996)
	Fw: 12SA	AAACTGGGATTAGATACCCACTAT	Palumbi (1996)	57	~380	426	52	GTR+I+G	0,22	0,52
	Rv: 12SB	GAGGGTGACGGGCGGTGTGT	Palumbi (1996)
	Fw: H3F	ATGGCTCGTACCAAGCAGACVGC	Colgan et al. (2000)	55	328	328	37	TrN+I+G	0,47	0,37
	Rv: H3R	ATATCCTTRGGCATRATRGTGAC	Colgan et al. (2000)

Table 3 .

 3 Homology of different structures of the anterior oliviform shell in Olivoidea and correlation of terminology used for its description in different genera. Abbreviations and conventions correspond to Figure5.

	Major and secondary areas	Grooves and	Oliva	Olivella	Amalda	Ancilla	Eburna
	(accepted here)	borders	(Tursch,	(Ollsson, 1956)	(Kilburn, 1977)	(Kilburn, 1981)	(Voskuil, 1991)
			(accepted	Greifeneder, 2001)
			here)
	Body whorl cloak (CL)	Olivoid band		Body whorl cloak absent	General surface absent	Ancillid band	(when present) Ancillid band	Ancillid band
		Upper			Upper			Posterior			Posterior
		anterior band			anterior			fasciolar			fasciolar
	Anterior band (AB)	(uab) Lower		Anterior band	band Lower	Fasciolar band	Fasciolar band	band Anterior	Fasciolar band	Fasciolar band	band Anterior
		anterior band			anterior			fasciolar			fasciolar
		(lab)			band			band			band
					Parietal
	Plication plate (PP)			Plication plate	plate belt	Columella pilar + parietal callus	Columella pilar	Columella pilar	Columella pilar
					shoe
			Olivoid	absent	absent	Ancillid groove	Ancillid groove	Ancillid groove
			groove 						(when present)
			Rear edge of	Rear edge of anterior	Not specified	Not specified	Posterior	Not specified
			anterior band	band				fasciolar groove
			
			Fasciolar	absent	Submedian line Posterior fasciolar	absent	Posterior fasciolar
			ridge 				groove		groove
			Rear edge of	Rear edge of belt	Not specified	Anterior fasciolar	Anterior	Anterior fasciolar
			plication plate				groove	fasciolar groove	groove
			

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Table 1 .

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Acknowledgements

The MNHN Indo-Pacific material originates from a series of expeditions and workshops, conducted in the context of the "Our Planet Reviewed" programme with Pro-Natura International (SANTO 2006 to Vanuatu; ATIMO VATAE to South Madagascar in partnership with Université de Tuléar, and its spin-off INHACA 2011 at Estação de Biologia Maritima, Universidade Eduardo Mondlane; MAINBAZA to Mozambique in partnership with Instituto Español de Oceanografia; PAPUA NIUGINI 2012, BIOPAPUA and KAVIENG 2014 to Papua New Guinea, in partnership with University of Papua New Guinea and the National Fisheries College), of the "Tropical Deep-Sea Benthos" programme with Institut de Recherche pour le Développement (CONCALIS, TERRASSES, NORFOLK 2, EXBODI, EBISCO, TARASOC, SALOMON 2 and SALOMONBOA3 cruises in the South Pacific; PANGLAO 2005 and AURORA 2007 cruises to the Philippines, in partnership with University San Carlos and the Philippines Bureau of Fisheries and Aquatic Resources), or stand-alone projects (KARUBENTHOS with Parc National de la Guadeloupe). The organizers thank the Total Foundation, Prince Albert II of Monaco Foundation, Stavros Niarchos Foundation, Fondation

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