Alexander E Fedosov

Nicolas Puillandre

Manfred Herrmann

Yuri I Kantor

Marco Oliverio

Polina Dgebuadze

Maria Vittoria Modica

Philippe Bouchet

Keywords: Mollusca < Taxa, new genera < Taxonomy, new species < Taxonomy, new classification < Taxonomy, species diversity < Taxonomy, molecular phylogeny

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Introduction

The neogastropod family Mitridae (miter shells, or miters) is a distinctive and familiar group of benthic gastropods, its key features being a solid, usually fusiform, colorful shell with strong columellar folds, predominant spiral sculpture and a smooth inner side of the labrum. Some iconic Mitra species like Mitra mitra (Linnaeus, 1758), Mitra papalis (Linnaeus, 1758) or Mitra cardinalis (Gmelin, 1791) were known since pre-Linnean times; the genus name established by Lamarck in 1798 and various specific epithets were named by early conchologists for their supposed resemblance with Church's ceremonial headwear.

The affinities of the miters were traditionally inferred from shell morphology. Following Reeve (1844Reeve ( -1845)), the name "mitres" was used broadly to refer to an array of marine snails that share a 'mitriform' habitus. However, as knowledge on the diversity and morphology of miters progressed, it became evident that "mitres" was in fact a heterogeneous group [START_REF] Risbec | Considérations sur l'anatomie comparée et la classification des gastéropodes prosobranches[END_REF]Cernohorsky, 1966[START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF][START_REF] Ponder | The morphology of some mitriform gastropods with special reference to their alimentary canal and productive systems (Mollusca: Neogastropoda)[END_REF] including two large, distinct families, Mitridae and Costellariidae. In our recent study [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF], we revised the composition and relationships of mitriform gastropods, demonstrating that the four mitriform families -Mitridae, Costellariidae, Ptychatractidae and Volutomitridae -do not form a single evolutionary lineage, with Mitridae appearing unrelated to the remaining three families. The distinctiveness of Mitridae is fully supported by morphological data: the organs of the digestive tract, the structure of the renal organ and the female reproductive system are drastically different in Mitridae and the other mitriform families [START_REF] Ponder | The morphology of some mitriform gastropods with special reference to their alimentary canal and productive systems (Mollusca: Neogastropoda)[END_REF]. Moreover, in the anatomy of the foregut and in the structure of the radula, the Mitridae show no apparent similarity to any other neogastropod family [START_REF] Ponder | The morphology of some mitriform gastropods with special reference to their alimentary canal and productive systems (Mollusca: Neogastropoda)[END_REF][START_REF] Taylor | Relationships of neogastropods[END_REF][START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF]. Two key features of Mitridae morphology, which set them apart from the rest of the Neogastropoda, are the epiproboscis -a complex muscular organ enclosed within the proboscis -and a radula with wide multicuspidate lateral teeth. Neither of these two features was found in species of the deep water family Pyramimitridae, a group sister to Mitridae in phylogenetic analyses [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF].

The Mitridae as currently restricted are a group flourishing in shallow waters of the Indo-Pacific, with a remarkable diversity of forms and sometimes great abundance. Although, the taxonomic diversity of ~400 Recent species worldwide (398 valid species currently indexed as valid in WoRMS) is not really impressive by neogastropod standards, it is still high, given some peculiarities of mitrid biology. Molluscs of the family Mitridae are predators with probably the most conserved dietary guilds among the Neogastropoda: all published data suggest that they feed exclusively on Sipuncula, however they do not appear to follow a strict species-specific prey specialization, and different, unrelated mitrid species appear to prey on the same sipunculan species within the same habitat [START_REF] Taylor | The diet of coral-reef Mitridae (Gastropoda) from Guam; with a review of other species of the family[END_REF][START_REF] Taylor | Dietary and anatomical specialization of mitrid gastropods (Mitridae) at Rottnest Island, Western Australia. In: The Marine Fauna and Flora of Rottnest Island, Western Australia[END_REF]. (WoRMS 2016) are classified in the genus Mitra, although the understanding that the genus, as currently circumscribed, is not consistent with a single evolutionary lineage has emerged long ago (Cernohorsky, 1996). Our previous phylogenetic analysis [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF] revealed that the species currently classified in Mitra are distributed throughout the mitrid tree, in Mitrinae, Imbricariinae and Cylindromitrinae, as well as in lineages that fell outside these three currently recognized subfamilies. Besides, obvious incongruences between relationships and current placement were uncovered in the genera Domiporta and Ziba. Thus there is no doubt that a revision of Mitridae systematics in a phylogenetic framework is needed. Several crucial mitrid lineages and type species of nominal genera were missing from the previous study, making the resulting tree insufficient for a thorough taxonomic re-evaluation. In the present study, we present a more robust tree based on an increased taxonomic coverage and revise the classification of the family based on this expanded molecular phylogenetic framework.

Material and Methods

Taxon sampling

The present study is largely based on material collected during multiple expeditions conducted by MNHN and IRD, University La Sapienza of Roma, the joint Russian-Vietnamese Tropical Center, and other ad hoc field work. The specimens studied originate from Panama, the Caribbean (Guadeloupe, French Guiana), Mediterranean (Adriatic, Aegean and Tyrrhenian Seas), the Azores, West Africa (People's Republic of Congo) and numerous Indo-Pacific localities from Madagascar to French Polynesia (see Table 1). To improve the taxon sampling of our previous study [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF], 121 additional mitrid specimens were analyzed, bringing the total number of specimens studied to 163 (Table 1). Before 2012, specimens were treated in the field with an isotonic solution of magnesium chloride until relaxed (showing no response to touch), and then a tissue clip was cut. Starting in 2012, specimens were processed using a microwave oven [START_REF] Galindo | Using microwaves to prepare gastropods for DNA Barcoding[END_REF], i.e. in most cases the entire body, or at least the last 1 -1.5 whorls, were available for study. Tissue samples were preserved in 96% ethanol.

Voucher shells are kept in MNHN, University La Sapienza and in the repository of the Laboratory of Ecology and Morphology of Marine Invertebrates of IEE.

The shells of the specimens selected for molecular analysis and morphological studies were first sorted to morphospecies and identified by the authors. Taxonomy follows WoRMS -largely based on [START_REF] Coan | Nomenclatural units in the gastropod family Mitridae[END_REF], [START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF][START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF]1991) with later updates [START_REF] Poppe | New Mitridae and Costellariidae from the Philippines and the East China Sea[END_REF][START_REF] Turner | Six new species of Mitridae from the Indian and Pacific Oceans, with remarks on Mitra abacophora Melvill, 1888 (Neogastropoda: Muricoidea)[END_REF][START_REF] Lussi | Endemic deep water Mitridae of South Africa with the description of three new species[END_REF][START_REF] Poppe | New species of Mitridae and Costellariidae from the Philippines with additional information on the Philippine species in these families[END_REF][START_REF] Huang | Calcimitra, a new genus of deep-water Mitridae (Gastropoda: Mitridae) with the description of five new species from Taiwan and the Philippines[END_REF]Huang & Salisbury 2017, etc.).

Subsequently, the barcode fragment of the COI gene was sequenced for as many as possible tissue clipped specimens. Morphospecies were thus used as preliminary hypotheses, to be verified by the analysis of the COI gene. When all specimens of a given morphospecies clustered in a single clade in the phylogenetic tree obtained with the COI gene, and this clade included only specimens of a single morphospecies, then it was considered that our species hypothesis was confirmed for subsequent analyses. When a morphospecies included only one specimen, the COI gene was used to check that it constituted a distinctive lineage of its own. Subsequently, a reduced dataset, with a selection of species best representing the currently recognized supraspecific taxa and the morphological diversity of Mitridae, was built. For these species, one only or a few specimens were sequenced for additional genes (16S rRNA, 12S rRNA and H3) to reconstruct phylogenetic relationships within the family. 12S sequence of one more species, "Mitra" carbonaria, analyzed by Colgan and coworkers (2007) was accessed from GenBank, and the photo of the voucher shell C.335424 in AMS was examined to confirm its identification and ensure that it is not conspecific with any other specimen sequenced in the present study. To complete the type series of inferred new species verified by the phylogenetic analysis, additional dry material from MNHN collections was studied to characterize their shell variation.

DNA extraction and PCR

DNA extraction and PCR were carried out as described in [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF]. All PCR fragments were pair-end sequenced by Eurofins and Macrogen (Amsterdam).

Datasets

To ensure the consistency of phylogenetic analyses with the results obtained earlier, the entire Mitridae-Costellariidae-Ptychatractidae-Volutomitridae (MCPV) datasets with multiple outgroups, as analyzed in [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF], were used and supplemented with newly obtained sequences. Six molecular datasets have been analyzed, four of them corresponding to the individual three mitochondrial (COI, 16S, 12S) and one nuclear (H3) markers. When the resulting phylogenetic reconstructions were checked for congruency, two concatenated datasets were assembled, one corresponding to the 3 mitochondrial genes (1,887 positions, 166 taxa, 7 % of missing data), referred to as "3-mit" and consistent with our earlier analysis, and the second a four genes dataset (2,215 positions, 166 taxa, 11 % of missing data), referred to as "4G" (Table 2). Finally, a reduced four genes dataset 4G-red was built to minimize the effects of missing data; only those 139 specimens that had at least three genetic markers sequenced, one of which is COI, were included in this dataset.

Phylogenetic analysis

Newly obtained COI and H3 sequences were aligned manually; 16S and 12S sequences were first aligned automatically with Muscle (Edgar, 2004) and then modified manually where necessary. In the analysis of multi-gene datasets, multiple neogastropod lineages were included as outgroups to ensure consistency with our earlier analyses [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF], Fedosov et al. 2016); in the single gene COI analysis -primarily aimed at species delimitation -, three specimens of the Pyramimitridae genera Teremitra and Vaughanites were used as outgroups based on the demonstrated sister relationship between Mitridae and Pyramimitridae [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF].

The best-fit substitution models were identified for each gene separately, using Modelgenerator V.85 [START_REF] Keane | Assessment of methods for amino acid matrix selection and their use on empirical data shows that ad hoc assumptions for choice of matrix are not justified[END_REF] (Table 2). The single gene datasets of 12S, 16S and H3 were analyzed as a single partition each; three partitions corresponding to 1st, 2nd and 3rd codon positions were analyzed independently for COI. The combined MCPV-3mit and MCPV- Best-scoring Maximum Likelihood (ML) trees were estimated using RaxML [START_REF] Stamatakis | RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models[END_REF], robustness of the nodes was assessed using the Thorough Bootstrapping algorithm [START_REF] Felsenstein | Confidence limits on phylogenies: an approach using the bootstrap[END_REF] with 1,000 iterations. Bayesian Analyses (BAs) were performed in two parallel runs in MrBayes [START_REF] Huelsenbeck | MrBayes: Bayesian inference of phylogeny[END_REF]. For 16S, 12S and H3 single gene analyses, each run consisted of six Markov chains and 10,000,000 generations with default number of chain swaps and sampling frequency of one tree in each 1,000 generations. For the COI, 3-mit, 4G, and 4G-red datasets, BA were performed with the number of chains set to 8, number of generations to 30,000,000, number of chain swaps to 5 and sampling frequency to one tree each 5,000 generations. Chain temperature of 0.02 was used in all analyses. Convergence of each analysis was checked using Tracer1.4.1 (Rambaut and Drummond, 2007); the two runs were considered converged if all ESS values exceeded 200. Consensus trees were calculated after discarding the first 25% trees as burn-in. All analyses were performed on the Cipres Science Gateway (http://www.phylo.org/portal2), using tools MrBayes 3.2.3 on XSEDE and RAXML-HPC2 on XSEDE (8.1.11). K2P genetic distances were computed using MEGA version 6.06 [START_REF] Tamura | MEGA6: Molecular Evolutionary Genetics Analysis version 6.0[END_REF].

Morphological studies

The specimens used for morphological study were, when possible, selected from among molecular vouchers to ensure an objective linkage of morphological and phylogenetic information. In other cases, conspecificity of the molecular vouchers and of specimens studied for morphology was based on examination of shell characters and preferentially those specimens that were collected from the same general localities were used for morphological studies. The snails were dissected, and the entire proboscis was extracted and transferred to distilled water.

For radula preparation, the proboscis was dissected and the buccal mass isolated from the anterior portion of proboscis; the buccal mass was suspended in diluted commercial bleach, where it was kept for 2-10 minutes until soft tissues completely dissolved; the radula was then rinsed in several shifts of distilled water and then transferred on clean 12 mm cover glass, where it was unfolded, air-dried and mounted for SEM study.

Due to the small size of studied specimens, anatomy of the proboscis was studied by serial sectioning, longitudinal in the case of Charitodoron and transversal for all other species.

The extracted proboscis was dehydrated and embedded in paraplast, subsequently 7 µm thin serial sections were prepared and stained with Masson's trichrome (Charitodoron) or standard hematoxilin-eosin (all other species). Anatomical abbreviations aoe -anterior oesophagus; bl -buccal lip; bm -buccal mass; bt -buccal tube; bw -body wall; bv -blood vessel; cm -columellar muscle; ep -epiproboscis; moe -mid-oesophagus; nnerves; nr -nerve ring; od -odontophore; odr -odontophore retractor; oe -oesophagus; poeposterior oesophagus; pt -tip of the proboscis; pw -proboscis wall; rd -radular diverticulum; sd -salivary duct; sg -salivary gland; vl -valve-like structure.

Other abbreviations SH -Shell height; w/h -ratio of shell width to shell height; a/h -ratio of aperture height to shell height; law/h -ratio of last adult whorl height to shell height; lv -live collected specimen(s); dd -dead collected specimen(s); B -Maximum likelihood bootstrap support value; PP -Bayesian Posterior Probability support value; BI -Bayesian Inference; ML -Maximum likelihood; OD -original designation of type species; SD -subsequent designation of type species; M -type species by monotypy.

Results

Single gene datasets

Transition substitution model with constant proportion of invariable sites and gamma distributed rate variation among sites (TIM+I+G) was selected as a best fit substitution model for the COI and 16S datasets, and Tamura-Nei substitution model with constant proportion of invariable sites and gamma distributed rate variation among sites (TrN+I+G) was found a best fit model for 12S and H3 (Table 2). The trees obtained after analyses of single gene datasets were largely congruent and contained no highly supported conflicting topologies. Whereas many inferred clades were well supported, in particular in the COI tree, the deeper nodes remained unstable and in most cases lacked support, being thus of little use for phylogenetic inference. Thus, we here only discuss the topology of the COI tree, as it was built upon a notably larger dataset compared to other single-gene trees and provides means for species delimitation.

The 149 mitrid specimens sequenced for COI were primarily segregated into 89 morphospecies. In all but four cases, species hypotheses based on shell morphology correspond to clades in the COI tree (or to an independent lineage when represented by a single specimen).

For the convenience of the presentation, we subdivided the COI tree in 28 clades, either moderately or well supported, or represented by single specimen; relationships among these 28 clades remained unresolved based on the analysis of COI.

Two morphospecies delineated based on the shell morphology each corresponds to two not closely related phylogenetic lineages in the COI based tree: (a) Specimens initially identified as Strigatella paupercula fell into two separate clades 7 and 11 (Figs 1, 2 A), and (b) the deep water specimens MNHN IM-2007-35618, MNHN IM-2007-35711 and IM-2013-40665, initially assigned to one morphospecies, ended up in clades 15 and 19 (Figs 1, 2 B). Conversely, morphologically diverse specimens that were initially treated as putative species complexes and identified as Cancilla rehderi / fibula (in subclade 24-7) and Mitra glaphyria (in clade 13 -Figs 1,2 C,D), in fact formed rather homogeneous molecular clades, each of them consistent with a single species: The computed K2P genetic distances varied from 0 to 0.8% among specimens of Mitra glaphyria, and from 0 to 3.9% among specimens of Cancilla rehderi / fibula. The latter, although demonstrating a rather high genetic variation, showed no bimodal distribution of pairwise distances, and we are inclined to attribute the observed genetic disparity to a consequence of its broad geographic distribution (sequenced specimens originated from localities spanning from the Philippines and Papua New Guinea to the Society Islands).

Fourteen additional species with distinctive morphology were included in the molecular studies, although amplification of the COI was not successful, but they were deemed important for the study and were included in further molecular analyses. All of them, based on genetic Of the 103 species segregated, 94 are attributable to existing species names, including six that were identified with ambiguity and may (Cancilla cf. fibula, Domiporta cf. carnicolor, Subcancilla cf. leonardhilli) or may not (Mitra aff. inca, Mitra aff. labecula, Strigatella aff. paupercula) represent the taxon in question. Eight species were not identified, of which one is a subadult, and seven are adult and can confidently be recognized as undescribed.

The sequenced species represent 16 currently accepted genera; of these only two correspond to a single clade in the COI tree: Pterygia (Clade 22-4, 4 species, PP=1), and Charitodoron (Clade 28, 2 species, PP=1).

Concatenated datasets: "3-mit" based tree

The consensus trees obtained from the analyses of multi-gene datasets showed similar topologies with no supported conflicts. In maximum likelihood (ML) and Bayesian analyses (BA) of all three concatenated datasets (Figs 3 -5), the first dichotomy splits a two-species clade, corresponding to Charitodoron, from the rest of the Mitridae (from here onwards referred to as Mitridae s.s.), both clades forming very long branches, and showing maximum nodal support in all analyses. The Mitridae s.s. clade is composed of eight major groupings, marked with different colours in the molecular trees ; their compositions were entirely consistent between trees, while support values and within-clade branching pattern varied slightly depending on the dataset analyzed. At the same time, relationships among these lineages remain unresolved; weakly supported affinities identified in one analysis appear unsupported in others. Here we describe the topology of the 3-mit tree (Fig. 3) and refer to other reconstructions where necessary.

The first major lineage (orange in Fig. 3), moderately supported in BA and weakly supported in ML (PP=0.97; B=74), comprises an assortment of Mitra lineages, along with species currently assigned to Nebularia, Domiporta and Ziba. This is the largest inferred lineage of Mitridae, combining 29 of the 82 species included in our 3-mit and 4G analyses. The distinct clades included in this first major lineage are:

1.1) a well supported Mitra mitra -Mitra papalis clade (PP=1; B=100); 1.4) a clade, moderately supported in BA (PP=0.99), combining a two species Mitra avenacea-Gen. sp. MNHN IM-2013-40655 clade (PP=1; B=98) with a well-supported grouping formed by three species currently assigned to Nebularia, Nebularia connectens, N. pediculus, N. maesta and Mitra tornata (PP=1; B=99). 1.5) a clade, well supported in BA (PP=0.99), combining "Mitra" glaphyria and "Mitra" invicta from deep-water of the Indo-Pacific, and a distinct well supported clade including three New World Mitra species (PP=1; B=99).

1.6) a clade, moderately supported in BA (PP=0.97), combining the West African Ziba carinata and the Mediterranean species pair Mitra cornicula and M. zonata (PP=1; B=99).

Relationships among the six clades of the first major mitrid lineage are not resolved in 3mit or any other analysis. From a nomenclatural point of view, the following species should be noted: Mitra mitra (type species of Mitra), Domiporta filaris (type species of Domiporta), Ziba carinata (type species of Ziba), and Mitra zonata (type species of Episcomitra) are marked with asterisks on the phylogenetic tree. From here onwards we will refer to this major lineage as the Mitra-Domiporta-Ziba-Episcomitra lineage.

The second major lineage of Mitridae s.s. (yellow in is weakly supported in the 3-mit analysis (PP=0.96) and forms a sister group to the first major lineage, although their relationship is also weakly supported by BA (PP=0.96). The second major lineage comprises multiple species commonly assigned to Strigatella from both the Indo-Pacific (S. paupercula, S. litterata, S. decurtata and S. scutulata) and the Panamic (S. tristis) regions, along with some Indo-Pacific species traditionally placed in Mitra, which form three well supported clades within the second major lineage: M. aurantia plus M. vexillum (PP=1; B=99); M. ambigua plus M. fulvescens (PP=1; B=100); and M. lugubris plus M. coronata (PP=1; B=100). From a nomenclatural point of view, the following species should be noted: Strigatella paupercula (type species of Strigatella) and Mitra coronata (type species of Chrysame) are marked with asterisks on the tree. From here onwards we will refer to this major lineage as the Strigatella-Chrysame lineage.

The third major lineage of Mitridae (PP=1, B=100: green in From a nomenclatural point of view, the following species should be noted: Imbricaria conularis (type species of Imbricaria), Neocancilla papilio (type species of Neocancilla) and Scabricola variegata (type species of Scabricola). From here onwards we will refer to this major lineage as the Neocancilla-Imbricaria-Scabricola lineage.

The fourth major Mitridae lineage (grey in includes two species, Mitra tuberosa and M. bernhardina, the affinities of which are moderately supported by BA and weakly supported in ML (PP=0.98, B=81). From here onwards we will refer to this lineage as the Mitra tuberosa lineage.

The fifth and sixth major lineages show sister relationships which are well supported at least by BA (PP=0.99, B=80). The fifth major lineage (PP=1, B=98: blue in Figs 345) combines (5.1) a distinct Pterygia clade (PP=1, B=100) and (5.2) a well supported four-species clade, which includes Indo-Pacific species of Mitra (M. incompta and M. eremitarum), Mutyca (M. acuminata), and Nebularia (N. contracta) (PP=1, B=94). From a nomenclatural point of view, the following species should be noted: Pterygia dactylus (type species of Pterygia) and Nebularia contracta (type species of Nebularia), respectively included in the first and second clades. From here onwards we will refer to this lineage as the Pterygia-Nebularia lineage.

The sixth major Mitridae lineage (PP=1, B=0.98: lilac in Figs 345) combines species of Mitra and Subcancilla. The lineage includes two clades, one of which comprises three Panamic Subcancilla species (PP=1, B=0.98), and the other combines Mitra glabra from Western Australia, Mitra nigra from West Africa, Mitra cornea from European seas, and Subcancilla straminea from the western Atlantic. The latter clade has a moderate nodal support in BA, and is not supported in ML (PP=0.96; B=68). From a nomenclatural point of view, it should be noted that Mitra glabra is the type species of Isara and Mitra nigra is the type species of Fuscomitra; from here onwards we will refer to this lineage as the Isara-Fuscomitra lineage. The eighth lineage (PP=1, B=100; light brown in Figs 345) is formed by two Indo-Pacific Mitra species, Mitra peculiaris and M. typha, from here onwards referred to as the Mitra peculiaris lineage.

Other concatenated datasets

Compared to the 3-mit tree, the 4G tree (Fig. 4) shows a slightly decreased support value for the Mitra-Domiporta-Ziba lineage (PP=0.95, B=79), and an insignificant rearrangement in the internal branching order. However, the support for the Strigatella lineage is higher in the 4G analysis compared to 3-mit, (PP=1, B=71), and nine of 12 species included in this lineage form a well supported grouping (PP=1, B=81). Neither content, nor support values changed significantly for the Neocancilla-Imbricaria-Scabricola, Pterygia-Nebularia or Isara-Fuscomitra lineages, as well as for the Mitra tuberosa and Mitra peculiaris lineages. However, the support has changed in some internal nodes, in particular the Swainsonia casta-Swainsonia fusca lineage shows a higher support compared to the 3-mit analysis (PP=1, B=76). Similarly, the clade that combines Neocancilla and Scabricola gains a higher support (PP=1, B=89) and is now clearly subdivided into two subclades, one corresponding to Neocancilla (PP=1, B=99), and the other combining Scabricola variegata, S. desetangsii and S. olivaeformis (PP=0.87, B=84).

Conversely, the grouping which unified the Pterygia-Nebularia and Isara-Fuscomitra major lineages shows even a lower support compared to the 3-mit analysis (PP=0.93, B=64).

Not unexpectedly, the phylogenetic tree based on the analysis of the 4G-red dataset (Fig. 5)

shows more significant changes in contents of the inferred groupings, primarily because of the reduced number of analyzed taxa. Nevertheless, and with the exception of Mitra barbadensis which was not included in 4G-red dataset, the seven major Mitridae lineages defined in the 3-mit and 4G analyses retained their principal composition, with support values comparable to those obtained in the 3-mit and 4G analyses. Among others, the Mitra-Domiporta-Ziba-Episcomitra lineage shows the greatest decrease in BA support (PP=0.9; B=0.79), however, the six clades that compose this major Mitridae s.s. lineage remain distinctive and well supported. Similarly, the phylogenetic clades corresponding to the large Neocancilla-Imbricaria-Scabricola, Pterygia-Nebularia and Isara-Fuscomitra lineages retained acceptable support values.

The support values of phylogenetic lineages inferred in analyses of the COI, 3-mit, 4G and 4G-red datasets are summarized in Table 3; the clade names used in the table and on the trees are discussed in the section below. Only few of the currently recognized mitrid genera emerge

F o r R e v i e w O n l y
15 monophyletic in our analyses: we confirm the monophyly of Pterygia and Charitodoron, while the other genera are either found to be poly-or paraphyletic, or their identification on the molecular tree is not straightforward (see section below).

Morphological studies

Here we present data on the foregut anatomy of two diverging groups of Mitroidea that have not been addressed before and are important for understanding evolutionary pathways in Mitroidea in general (see Discussion). Although the specimen had been positioned for obtaining sagittal sections, the proboscis was diverted and sectioned nearly in frontal plane (Fig. 6 A,D). Proboscis moderately long, broad, conical, strongly contracted, so that its wall forms several telescopic folds (Figs 6 A,C,D).

Charitodoron

Mouth opening into medium long buccal tube, spanning around 1/3 of proboscis length and leading into broad buccal cavity. Buccal cavity subdivided into two chambers by the large circular fold of its wall (Fig. 6 -F). Wide anterior oesophagus leaving buccal cavity dorsally, forming rather long loop within proboscis, leaving proboscis and narrowing before passing through the nerve ring and then widening before transition to the posterior oesophagus (Fig. 6 D). Distinct valve-like structure, identifiable as a valve of Leiblein although lacking a ciliary cone, delimiting an anterior and a posterior oesophagus (Fig. 6G, vl). Anterior oesophagus empty in sectioned specimen, posterior oesophagus filled with food material (Fig. 6 D, G) in which, however, no detectable remnants were found. After the oesophageal valve, tall and lightly stained epithelium of anterior oesophagus changing to low epithelium formed by small cubic cells. Anterior oesophagus lined with thin layer of muscular fibers, which become notably stronger in posterior oesophagus (Fig. 6 G).

Thin and strongly convoluted salivary ducts adjoining anterior oesophagus in front of passage through nerve ring and passing forward being embedded into its muscular wall, leaving oesophageal wall closer to the proboscis tip and entering buccal mass. Paired odontophore cartilages well developed, not fusing with strong lateral odontophore protractor muscles attached to odontophore ventrally (Fig. 6 forming massive "lip", longer dorsally (buccal lip,E). After entering buccal cavity wall, distal salivary ducts passing inside that lip laterally on both sides of radular diverticulum opening (Fig. 6 D, E), and opening in the buccal cavity ventrally, underneath the semi-circular fold (Fig. 6 F). Epiproboscis absent.

Salivary gland large, unpaired, situated to the left of posterior oesophagus and dorsally to the nerve ring, histologically represented by a compactly packed, strongly convoluted tube with one layer of tall highgly vacuolar, obviously glandular, cells and thin outer connective tissue layer (Fig. 6 A,B).

Condylomitra tuberosa [PANGLAO 2004, Stn B5].

Proboscis moderately long, leaf-shaped, anteriorly flattened dorso-ventrally (Figs 7 A -H) and cylindrical at its base (Fig. 7 I). On transversal sections, proboscis with two lateral lobes at midlength and ventral part with rounded axial keel. Proboscis wall with thick cuticle and nearly cubic epithelium, with large basal nuclei; epithelium underlined by a layer of circular muscle fibers (Figs 7 C,E,H). Lateral lobes occupied by complex interlacement of dorso-ventral, oblique, and longitudinal muscular fibers; two blood vessels (7 F, bv) running along margins of proboscis (Figs 7 B,D,F,G). Central part of proboscis occupied with oesophagus, anterior buccal mass and epiproboscis complex forming a single stem.

Mouth opening anteriorly on the dorsal side of the proboscis in short dorsal groove; mouth surrounded by small peristome rim (Fig. 7 A Salivary ducts passing forward along the anterior oesophagus, embedded into lateral folds of oesophagus wall (Figs 7 D -I); salivary gland itself not sectioned. Salivary ducts shifting ventrally in the walls of the buccal cavity adjoining the epiproboscis sheath. More distally, salivary ducts running backwards along the epiproboscis sheath to its proximal part, where they enter the epiproboscis and run through its entire length to open at its tip (Fig. 7 E).

Discussion

Application of family-group names to the phylogenetic clades

The phylogenetic trees obtained in analyses of multiple-gene datasets were largely congruent and consistent with each other with respect to the nodal support of inferred lineages, thus providing robustness for the phylogenetic hypotheses of Mitridae. We chose the 4G based tree for further taxonomic inferences, because of (1) the better taxonomic coverage compared to the 4G-red based tree, and (2) the fewer polytomies in the 4G tree compared to the 3-mit based tree.

Our results agree in general with those obtained in earlier analyses [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF].

The assignment of names to the major lineages of Mitridae s.s. is straightforward in three cases. [START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF]1991). The Pterygia clade is unequivocally identified in our phylogenetic trees, including the type species P. dactylus. Because the Pterygia lineage shows a well supported affinity to the Nebularia contracta clade in all analyses, the name Cylindromitrinae can be applied to the resulting Pterygia-Nebularia lineage.

Beyond the names in current use above, another available name is Strigatellidae which, although little used, can be applied to the Strigatella major lineage, so here it is designated as Finally, the three major lineages "Mitra" peculiaris, "Mitra" tuberosa and "Mitra" barbadensis are here treated as separate mitrid genera, but their subfamily assignments are uncertain and we leave them as incertae sedis.

The Mitridae s.s. clade has a maximum support in all multiple-genes analyses, and is defined by a very long branch. Moreover, all studied members of the Mitridae s.s. share morphological features that are not present in Charitodoron. We thus restrict the family name Mitridae to the Mitridae s.s. clade and establish a new family for Charitodoron. The obvious morphological distinctiveness of Charitodoron, and its position on the molecular tree as a sister group to all other Mitridae justify its inclusion in a separate family. The lineage comprising Mitridae, Charitodoronidae fam. nov. and Pyramimitridae is well supported in all our phylogenetic analyses, and is consistent with earlier results [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF]; therefore we rank it as superfamily Mitroidea stat. nov.

Application of genus-group names to the phylogenetic clades

Our 4G molecular dataset includes the type species of 13 nominal genera: nine (Domiporta, Imbricaria, Mitra, Nebularia, Neocancilla, Pterygia, Scabricola, Strigatella and Ziba) currently treated as valid and four (Chrysame, Episcomitra, Fuscomitra and Isara) treated as synonyms [of Mitra]. Here we elaborate a new classification of the Mitridae based on genetically defined monophyletic groups. The inferred lineages including one (or more) of the type species mentioned above are named with certainty, but for the other clades we have to apply names based on their common usage and in agreement with shell and radular characters. The clades for which there is no available name are treated as new genera.

We unequivocally ascribe the genera Mitra, Domiporta and Ziba to the subfamily Mitrinae, based on their inclusion in the Mitrinae lineage of the phylogenetic trees. Based on the analysis of the COI gene, we transfer Neocancilla circula to the genus Domiporta. As here circumscribed the genus Ziba corresponds to a West African clade, and numerous Indo-Pacific imbricariine species assigned to Ziba after [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF] are here transferred to Imbricaria. Two other clades within the Mitrinae lineage have available names: Mitra zonata is the type species of Episcomitra, subsequently synonymized [START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF] with Mitra. Based on the sequenced type species, we reinstate Episcomitra at the rank of genus with two Mediterranean species assigned to it, "Mitra" zonata and "Mitra" cornicula. Based on shell and radula characters we tentatively attribute the Indo-Pacific deep water clade "Mitra" glaphyria -"Mitra" invicta to the genus Calcimitra. Although the Mitrinae lineage comprises the six distinct clades described above, some of them show morphological, geographical, bathymetric, and molecular disparities that lead us to further subdivide them. Thus we delineate on the molecular tree twelve well supported clades (some being single-species) that we rank as genera (marked light grey on the trees). Five of them have applicable names, and the remaining seven are here treated as new genera:

Quasimitra n. gen. for the Indo-Pacific clade of "Mitra" sophiae, "Mitra" puncticulata, "Mitra" cardinalis, and "Mitra" sanguinolenta; Roseomitra n. gen. for the Indo-Pacific clade of "Mitra" rosacea and "Mitra" Pseudonebularia n. gen. for the Indo-Pacific "lesser" Nebularia clade of "Nebularia" pediculus, "N." connectens, "N." maesta, and "Mitra" tornata, which show no affinity to either Mitra or Nebularia as circumscribed here (the latter genus as typified by N. contracta is found to be closely related to Pterygia -see below); Gemmulimitra n. gen. for the Indo-Pacific clade of "Mitra" avenacea and MNHN IM-

2013-40655;

Neotiara n. gen. for the clade of New World species "Mitra" lens, "Mitra" nodulosa and "Mitra" cf. inca.

Cancillopsis n. gen. for the Indo-Pacific clade 1 of our COI based tree, which shows no affinity to any other mitrid lineage in the analysis.

Five species of Strigatella, including the type species, S. paupercula, are included in the Strigatellinae lineage of our trees, along with numerous "Mitra" species. As Strigatella and "Mitra" species do not form clades but are well intermixed, we recognize within this subfamily only one genus Strigatella and assign to it all the species in the Strigatellinae lineage. However, the genus name Chrysame (the type species Mitra coronata is included in our dataset) would apply to the Mitra coronata -Mitra lugubris subclade, which forms a very long branch and has a peculiar monoserial radula (see below) which differs remarkably from the radula of the rest of the Strigatellinae. From a cladistic point of view, recognition of Chrysame would render Strigatella paraphyletic, or many new genera would need to be established. We think the latter approach would be premature and, despite the unique position of Chrysame in the molecular trees, we treat it as a synonym of Strigatella. The first clade of the Imbricariinae includes Imbricaria conularis (type species of the genus) and an assortment of "Ziba" and "Subcancilla" species from the Indo-Pacific, which we thus transfer to Imbricaria. The second inferred imbricariine clade combines Cancilla rehderi / fibula and "Mitra" schepmani. Based on the similarity in shell and radula morphology between the Cancilla species included in this clade and the type species C. isabella (as examined recently by [START_REF] Salisbury | Notes on Cancilla isabella (Swainson, 1831) (Neogastropoda: Mitridae) with emphasis on the radula and generic assignment within Mitridae[END_REF], we tentatively attribute the name Cancilla to this clade. The third imbricariine molecular clade combines Neocancilla with Scabricola, Swainsonia and two former "Imbricaria" species. As Neocancilla species form a well supported clade with N. papilio (the type species) included in it, we confidently allocate this clade to Neocancilla. In sister position to the Neocancilla, the well supported grouping of Scabricola variegata (type species of Scabricola), S. desetangsii and S. olivaeformis is recovered; here we allocate it to the genus Scabricola. The Swainsonia fusca -Swainsonia casta clade is recovered as a sister group to the Neocancilla-Scabricola. As the type species of Swainsonia, Mitra fissurata, closely resembles Scabricola fusca in characters of the shell and radula, we consider that the name Swainsonia is available for it. Neocancilla, Scabricola and Swainsonia (as circumscribed here) form a well supported clade that is sister to the single species clade formed by "Imbricaria" punctata; for the latter, we establish Imbricariopsis n. gen.

Apart from Pterygia, the newly circumscribed Cylindromitrinae includes a closely related clade which combines "larger" Mitra species, "M." eremitarum and "M." incompta, with Mutyca acuminata and Nebularia contracta; the latter is the type species of Nebularia, and hence we attribute the name Nebularia to this four-species clade.

The names Isara and Fuscomitra, hitherto considered synonyms of Mitra, are available for the well supported "Mitra" glabra -"Mitra" nigra -"Mitra" cornea -Subcancilla straminea clade, as the type species of these nominal genera are included in it. Here we allocate the name Isara to this clade, and treat Fuscomitra as a synonym. The Isara clade forms a sister group to another well supported clade formed by three Panamic species of Subcancilla. As circumscribed by [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF], the genus Subancilla combined species from the Panamic and Caribbean regions, as well as from the Indo-Pacific, and the monophyly of the genus remained unconfirmed and doubtful [START_REF] Thorsson | Panamic and Caribbean Subcancilla[END_REF]. The type species Subcancilla sulcata is Panamic and, according to [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF], it differs significantly from the Indo-Pacific species in the morphology and dimensions of radula. Our analysis confirms the non-monophyly of the Subcancilla species from the Panamic region with those from the Indo-Pacific. Based on biogeographical grounds and radular morphology, we here attribute the name Subcancilla to the clade of Panamic species, which is recovered as the sister group to the newly circumscribed Isara. No genus group names are available for the inferred Mitra tuberosa clade and Mitra peculiaris clade. Thus we establish two new genera, Condylomitra n. gen. and Carinomitra n. gen., respectively for these distinctive lineages.

Three further lineages showed no supported affinity to any of the inferred major lineages that we rank as subfamilies; at present we abstain from giving them any subfamilial assignment, and treat them as incertae sedis. Two genera that are not represented in our dataset, Atrimitra Dall, 1918 andDibaphimitra Cernohorsky, 1970, also cannot be attributed to any of the newly identified subfamilies based on the published morphological data [START_REF] Bayer | Observations on Mitra florida Gould[END_REF][START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF], thus at present we also treat them as incertae sedis, pending further molecular studies to clarify their affinities.

In conclusion, the combined molecular and morphological data lead to the revised classification below.

Superfamily Mitroidea* Swainson, 1831

Family Charitodoronidae new family Genus Charitodoron Tomlin, 1932.

Family Mitridae Swainson, 1831

Subfamily Mitrinae Swainson, 1831

Genera Mitra Lamarck 1798, Ziba H. Adams & A. Adams, 1853, †Clifdenia Laws, 1932, Eumitra Tate, 1889, Episcomitra Monterosato, 1917, Domiporta Cernohorsky, 1970, Calcimitra Huang, 2011, Fusidomiporta n. gen., Gemmulimitra n. gen

Incertae sedis

Genera Atrimitra Dall, 1918, Vicimitra Iredale, 1929, Dibaphimitra Cernohorsky, 1970, Probata Sarasúa, 1989, Magnamitra Huang & Salisbury, 2017, Carinomitra n. gen Diagnosis: Shell medium-sized to large (50-170 mm), thick-walled, fusiform or elongatefusiform, with truncated base and colour pattern of multiple rounded or squarish red spots or bands on white background. Protoconch multispiral, of about 3 smooth slightly convex whorls. Suture deeply impressed. Teleoconch whorls gently convex or angulated adapically, bearing a row of short spiny bumps at shoulder. Rest of whorl surface sculptured with weak, evenly spaced, spiral grooves, well pronounced on early spire whorls and often vanishing later. Siphonal canal very short with robust siphonal fasciole delineated from shell base by deep depression.

Aperture rather wide, with deep siphonal notch; outer apertural lip nearly straight adapically and strongly convex in its lower portion, bearing numerous fine denticles throughout its length or limited to its abapical margin. Inner lip reflected, with 3-4 subequal oblique columellar folds.

Radula with rather narrow rachidian bearing 6 subequal cusps, and very wide laterals bearing cusps throughout their width (Fig 10 A). Cusps rather strong near rachidian and becoming progressively weaker laterally.

Distribution: Indo-Pacific, shallow intertidal and subtidal to upper bathyal depths, sand.

Species included: Mitra mitra (Linnaeus, 1758) 1 , Mitra deprofundis Turner, 2001 3 , Mitra papalis (Linnaeus, 1758) 1 , Mitra stictica (Link, 1807) 3 .

Remarks.

A critical reassessment of the contents of the genus Mitra is one of the main outcomes of the present study, and our phylogenetic analysis demonstrates that the name Mitra may be confidently applied to a handful of species only. The redefined genus Mitra is morphologically Quasimitra cardinalis and Q. puncticulata resemble species of Mitra. However, Q. cardinalis has a notably wider shell with lower spire, and Q. puncticulata lacks the characteristic colour pattern of Mitra and bears no denticle on the outer aperture lip. Nebularia incompta also resembles Mitra in shell proportions and the presence of denticles on the margin of the outer lip, but has a stronger sculpture and its siphonal canal is not clearly delineated from the shell base; besides, it has a typical Nebularia-type colour pattern instead of a Mitra-type. Some Strigatella species, especially S. imperialis, also closely resemble Mitra in the morphology of the abapical part of the shell (aperture and siphonal canal), but they have a notably lower spire and lack the Mitra-type colour pattern as well. Diagnosis: Shell medium-sized to large (35-125 mm), elongate-fusiform, with high orthoconoid or slightly acuminate spire and slender, tapering siphonal canal. Protoconch narrowly conical, multispiral, of about 3 smooth convex whorls. Suture deeply impressed or canaliculate. Spire whorls flattened to evenly convex, smooth, or sculptured with weak and irregular, or distinct, evenly set cords. Interspaces between spiral cords sometimes ornamented with fine collabral growth lines or dense riblets sometimes forming fine cancellate sculpture pattern. Siphonal canal long, tapering, with shallow or indistinct siphonal notch. Aperture elongate, its outer lip smooth, evenly convex. Inner aperture lip often calloused, sometimes reflected, bearing 3-4 fine columellar folds. Shell white, pale to orange, sometimes bearing pattern of reddish or brownish axial bands; fresh specimens often covered with brown periostracum. Radula with rather narrow rachidian, bearing 6-8 subequal cusps; laterals wide, bearing 12-20 pointed cusps (Figs 10 B,C), two proximal cusps rather short, weaker than the next five ones that are strongest; marginal 5-8 cusps gradually diminishing.

Genus

Distribution: Indo-Pacific, bathyal depths, mud, sometimes on sunken wood.

Species included: Calcimitra arnoldeyasi (Poppe, Tagaro & Salisbury, 2010) 3 comb. nov., C. brinkae (Salisbury & Kilburn, 1996) 3 comb. nov., C. brunetta (Chino & Herrmann, 2015) (Turner & Salisbury, 2007) 3 comb. nov., C. marrowi (Turner, 2001) 3 comb. nov., C. meyeriana (Salisbury, 1992) Remarks. The five species included in clade 13 of the COI based tree show considerable variation in shell morphology, both intra-and interspecifically. Six sequenced specimens of Calcimitra glaphyria display a range of sculpture types, from smooth shells resembling Calcimitra kingtsio to distinct, cancellated ones, and differently sculptured specimens can be found in the same general locality. While typical Calcimitra has a medium-sized, thin-walled shells, our phylogenetic analysis placed 'Mitra' invicta in the same phylogenetic clade. The latter is a recently described species representative of a group of large deep-water mitrids (here referred to as the C. morchii complex) that possess heavy, strongly sculptured shells [START_REF] Huang | Magnamitra n. gen. and nomenclatural remarks on large Cancilla and Mitra from Taiwan and the Philippines (Gastropoda: Mitridae)[END_REF]). The close affinity of these conchologically diverse species is consistent with a generally conservative radular morphology in Calcimitra kingtsio [START_REF] Huang | Calcimitra, a new genus of deep-water Mitridae (Gastropoda: Mitridae) with the description of five new species from Taiwan and the Philippines[END_REF], C. morchii, C. chiangfucius, C. philosopha (Huang & Salisbury 2017), C. glaphyria and C. invicta (herein). The intraspecific variation in sculpture pattern demonstrated by C. glaphyria challenges the status of the species in the Calcimitra morchii complex accepted by [START_REF] Huang | Magnamitra n. gen. and nomenclatural remarks on large Cancilla and Mitra from Taiwan and the Philippines (Gastropoda: Mitridae)[END_REF]. The distinction of C. chiangfucius, C. taiwanbale and C. invicta is based on subtle differences in shell morphology; they may prove to represent variants of a single species, and more comprehensive sampling is necessary to clarify species boundaries.

Magnamitra sandrogorii strongly resembles some of the larger species of Calcimitra, and the distinction of these two genera based on shell characters is problematic. The spiral sculpture in M. sandrogorii is even stronger than in species of the Calcimitra morchii complex, and the main distinction between the two genera is in the radula. In Magnamitra, the rachidian has a very strong unpaired central cusp that notably exceeds lateral cusps, whereas all studied species of Calcimitra possess a rachidian with equal or subequal cusps. Furthermore, rachidian and laterals have comparable width in Magnamitra sandrogorii, whereas in species of Calcimitra the width of the rachidian does not exceed half of the laterals width. Species of Profundimitra n. gen. superficially resemble Calcimitra, but the latter genus can be recognized by the generally larger shells with proportionally longer siphonal canal and often calloused inner aperture lip. The radulae of species of Calcimitra and Profundimitra n. gen. differ greatly, the latter having rachidian and marginals of about equal width. Species of Eumitra are also similar to Calcimitra in shell proportions and sculpture, however Eumitra can be distinguished by its columellar folds, which are reduced to not more than 2 very weak folds, while in Calcimitra there are four (very rarely 3) distinct ones. Some species of Calcimitra, in particular C. triplicata, resemblance the Miocene fossil genus Clifdenia, both in whorl outline with evenly rounded inflated shoulders, and in shape and orientation of columellar plaits (A. Beu, personal communication). Based on this resemblance, we tentatively allocate Clifdenia to the subfamily Mitrinae.

Genus Eumitra Tate, 1889 (Fig. 12) Type species: †Mitra alokiza Tenison-Woods 1879; SD (Ludbrook, 1958: 71).

Diagnosis: Shell small to medium-sized (15-58 mm), fusiform, thin-walled, with high orthoconoid spire and slender siphonal canal. Protoconch very low, cyrtoconoid, paucispiral, with glossy whorls and rather small nucleus in all known species. Suture distinct, impressed.

Early teleoconch whorls subcylindrical, later evenly convex, sometimes resulting in a slightly acuminate spire profile. Shell smooth, sometimes with very weak axial folds on first teleoconch whorl or fine grooves on shell base and siphonal canal. Siphonal canal, slender, tapering.

Siphonal notch absent. Aperture elongate, outer lip evenly convex, smooth. Inner aperture lip either smooth, bearing one very weak or sometimes two distinct, but weak, columellar folds.

Shell uniformly pale to light brown. Radula with very small, narrow rachidian, bearing 5 short robust cusps, only slightly extending beyond preceding tooth margin, but forming ridges, notably elevated above surface of rachidian. Laterals wide with 8-10 strong, pointed subequal cusps.

Distribution: New Caledonia, Coral Sea and the Philippines at bathyal depths, and a further undescribed species referable to Eumitra from Indonesia. Fossil species widely distributed in the Miocene of Europe and in the Australia -New Zealand region.

Recent species included: Eumitra apheles Lozouet, 1991 2 , E. caledonica Lozouet, 1991 3 , E. imbricata Lozouet, 1991 3 , E. richeri Lozouet, 1991 3 [START_REF] Lozouet | Eumitra récentes de la région néo-calédonienne et Charitodoron fossiles de l'Oligocène supérieur d'Aquitaine (Mitridae)[END_REF], although additional material from the West Pacific has been examined. The placement of Eumitra in the newly circumscribed Mitrinae is thus tentative and based on the plesiomorphic radula of Eumitra and the presence of a well developed epiproboscis (YK, unpublished). Based on shell characters, Eumitra resembles Calcimitra and Profundimitra n. gen., however in the latter two genera species commonly have 3+ strong columellar folds. Besides, all studied species of Profundimitra n. gen. are characterized by a radula with a rachidian as broad as the laterals. Species of Eumitra lacking shell sculpture somewhat resemble the costellariid genera Latiromitra and Costapex [START_REF] Fedosov | Systematics, diversity and evolutionary history of the family Costellariidae (Gastropoda: Neogastropoda)[END_REF], which however can be distinguished by the presence of an operculum, although reduced, and a longer siphonal canal (Latiromitra) or 3+ distinct columellar folds (Costapex).

Moreover, in both Latiromitra and Costapex, the early teleoconch whorls are sculptured with strong axial ribs that are absent in Eumitra.

'Eumitra' suduirauti Bozzetti, 1997 (Fig. 12 E), shows some features (such as a wide aperture, especially wide in its anterior portion, a distinct subsutural ramp, pronounced on late teleoconch whorls, as well, as on early spire whorls) that are uncommon not only in the genus Eumitra, but in the family Mitridae in general. However, as no data on radula morphology or DNA of this odd species is available, we maintain it in the genus Eumitra pending for better placement to be determined in future.

All known species of Eumitra share a paucispiral protoconch that distinguishes them from most other mitrid genera. Nevertheless, given the wide distribution of Eumitra in the fossil record, we cannot rule out the possibility of multiple transitions to non-planktotrophic development in the evolutionary history of the genus, and thus some members of Eumitra might have retained planktotrophic development, and thus a multispiral protoconch. Diagnosis: Shell small to large (20-75 mm), solid, coarsely sculptured, broadly fusiform to about turriform. Protoconch multispiral, rather wide, cyrtoconoid, of about 3.5 smooth, convex whorls.

Genus

Spire high; suture deeply impressed. Teleoconch whorls flattened to strongly convex or distinctly shouldered. Sculpture of slightly prosocline, wide and broadly spaced axial folds, or rather dense ribs; axials overridden by spiral cords which are typically wide and flattened, but may be narrow and elevated. Rounded or spirally elongated beads sometimes present at the intersection of spiral and axial sculptural elements. Siphonal canal rather short, stout, with weak or no fasciole. Remarks. According to Cernohorsky (1976: 454), the type of Mitra lens, which used to be in Mawe's collection, can no longer be traced. In order to fix the identity of the species and of the genus Neotiara n. gen., we here designate our sequenced specimen BAU00800 from Panama City, Panama, measuring 40.0 x 15.5 mm (Fig. 13 A), as neotype of Mitra lens, MNHN IM-2000-33145. As circumscribed here, the genus Neotiara can be recognized among New World Mitridae primarily by the well developed, typically coarse sculpture with distinct axial elements. Other Caribbean and Panamic Mitridae of the genera Isara, Subcancilla, Probata and Atrimitra lack axial sculpture, and the spiral elements are either lacking as well, or represented by fine cords.

As indicated by our phylogenetic analysis, the Panamic Neotiara lens and N. aff. inca, and the Caribbean N. nodulosa are closely related and Notiara is at present the only mitrid genus with both Pacific and Atlantic species. Although only three species are included in our phylogenetic analysis, we classify all heavily sculptured American mitrids in Neotiara n. gen.

Genus Ziba H. Adams & A. Adams, 1853 (Figs 10 F, 14 A -D)

Type species: Mitra carinata Swainson, 1824;SD, Wenz (1943: 1292). Early teleoconch whorls evenly convex, with weak cancellate sculpture, later whorls typically distinctly carinated. Sculpture of weak grooves on late spire whorls, sometimes missing on last adult whorl. Last adult whorl high, with shell base gradually extended into tapering siphonal canal, typically sculptured with wide and low rounded cords. Siphonal notch weakly or moderately shallow. Aperture elongate, narrow, its outer lip smooth, gently convex or flattened.

Inner lip with three fine, oblique columellar folds. Rachidian with five, equally developed, pointed, median cusps, flanked by two notably smaller marginal cusps (Fig. 10 F). Lateral teeth multicuspidate, third proximal cusp strongest, and subsequent cusps gradually diminishing; distal third of tooth smooth, with no cusps.

Distribution: West Africa, continental shelf, on mud.

Species included: Ziba carinata (Swainson, 1824) 1 , Z. gambiana (Dohrn, 1861) 3 .

Remarks. The present study has lead to a drastic reconsideration of the position and contents of the genus Ziba. Based on the results of our phylogenetic analysis, Ziba is now classified in the subfamily Mitrinae and its contents exclude all the Indo-Pacific and West American species hitherto referred to Ziba by [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF]. An array of Indo-Pacific Ziba species are here transferred to Imbricaria, whereas the Panamic Ziba attenuata is reclassified in Subcancilla, neither of these genera showing an affinity to the true Ziba, which ends up being confined to the tropical eastern Atlantic; to our knowledge, there is no Indo-Pacific species referable to this genus. The radula of Ziba carinata shows a plesiomorphic morphology which is commonly found in Mitridae and, although supporting placement of Ziba in Mitrinae, adds little to understanding its relationships within the subfamily. The moderately supported lineage Ziba carinata -Episcomitra inferred in the 3-mit tree (though absent in other analyses) may reflect an affinity that makes sense from the biogeographical point of view; however, further studies are needed to determine the relationships of Ziba. Species included: Episcomitra cornicula (Linnaeus, 1758) 1 comb. nov., E. zonata (Marryat, 1818) 1 .

Genus

Remarks. The genus Episcomitra includes two of the three mitrid species inhabiting the Mediterranean, which share a smooth shell with well-developed periostracum. However, the same uninformative shell morphology characterizes Isara cornea, which occurs sympatrically with Episcomitra in the Mediterranean. Episcomitra and Isara are equally indistinguishable in radular characters: both share the same general morphology, and the intrageneric variation in Isara by far exceeds the subtle differences between typical Episcomitra and Isara radula morphologies. Thus, there are no morphological characters that substantially tell apart the two genera, as represented by Episcomitra zonata and E. cornicula and Isara cornea. This remarkable similarity in shell morphology demonstrates the limited use of shell characters for supraspecific assignments. -M). Laterals rather wide, with 10-14 widely set and triangular, or dense and rather long, pointed cusps distributed throughout tooth width, proximal 1-2 cusps notably smaller than succeeding ones.

Genus

Distribution: Indo-Pacific, upper subtidal depths, mostly on sand.

Species included: Quasimitra albocarnea (Bozzetti, 2016) 3 comb. nov., Q. barbieri (Poppe & Tagaro, 2006) 3 comb. nov., Q. bovei (Keiner, 1838) 3 comb. nov., Q. brettinghami (E.A. Smith, 1906) 3 comb. nov., Q. cardinalis (Gmelin, 1791) 1 comb. nov., Q. floccata (Reeve, 1844) 3 comb. nov., Q. fulgurita (Reeve, 1844) 3 comb. nov., Q. lacunosa (Reeve, 1844) 3 comb. nov., Q. lamarcki (Deshayes, 1832) 3 comb. nov., Q. latruncularia (Reeve, 1844) 3 comb. nov., Q. leforti (Turner, 2007) 3 comb. nov., Q. manuellae (Cossignani & Cossignani, 2006) 3 comb. nov., Q. marmorea [START_REF] Turner | Six new species of Mitridae from the Indian and Pacific Oceans, with remarks on Mitra abacophora Melvill, 1888 (Neogastropoda: Muricoidea)[END_REF] (Turner, 1993) (Poppe, 2008) 3 comb. nov., Q. solida (Reeve, 1844) 3 comb. nov., Q. sophiae (Crosse, 1862) 1 comb. nov., Q. stossieri (Herrmann, 2016) 3 comb. nov., Q. structilis (Herrmann & Salisbury, 2016) 3 comb. nov., Q. variabilis (Reeve, 1844) 3 comb. nov.

3 comb. nov., Q. nubila (Gmelin, 1791) 3 comb. nov., Q. nympha (Reeve, 1845) 3 comb. nov., Q. propinqua (A. Adams, 1853) 3 comb. nov., Q. pseudobovei (T. Cossignani & V. Cossignani, 2005) 3 comb. nov., Q. puncticulata (Lamarck, 1811) 1 comb. nov., Q. punctostriata (A. Adams, 1855) 3 comb. nov., Q. rinaldii
3 comb. nov., Q. roselinae (Martin & Salisbury, 1987) 3 comb. nov., Q. rossiae (Reeve, 1844) 3 comb. nov., Q. sanguinolenta (Lamarck, 1811) 1 comb. nov., Q. sarmientoi
Etymology: Latin quasi, meaning appearing like, which reference to the overall shell resemblance with species of the genus Mitra. Gender feminine.

Remarks. The genus Quasimitra n. gen. is defined primarily based on molecular characters, and there are no apomorphies or clear diagnostic morphological features that support its recognition.

Shell proportions and sculpture vary extensively, with two extremes represented by Q. cardinalis and Q. sophiae, the close affinity of which is evidenced by the molecular data. The radulae of studied species generally retain a plesiomorphic morphology and also do not present any notable feature.

We tentatively assign Mitra solida Reeve, 1844 to Quasimitra, because of its characteristic 33 sequenced species here assigned to Quasimitra. At present, we treat Vicimitra prosphora as a separate species in a monotypic genus Vicimitra, but the name Vicimitra may prove to be a senior synonym of Quasimitra. Some species of Strigatella, like S. luctuosa or S. aurantia, bear a remarkable resemblance to Quasimitra in shell outline, and can only be distinguished by the generally more flattened spire whorls; at the same time there is a consistent difference in radular morphology between the two groups: the laterals bear cusps throughout their width in Quasimitra, whereas the distal third of laterals is devoid of cusps in the species of Strigatella mentioned.

Genus Domiporta Cernohorsky, 1970 (Figs 16 A -E, 17 A -D)

Type species: Voluta filaris Linnaeus, 1771; OD.

Diagnosis: Shell small to medium-sized (12-65 mm), fusiform to elongate-fusiform, with high spire and rather short aperture. Protoconch pointed, narrowly-conical, of 2.5 -3.5 smooth, slightly convex whorls to large, cyrtoconoid, of nearly 5 smooth whorls. Suture distinct, impressed. Teleoconch whorls evenly convex in outline, with dominating spiral sculpture represented by regular, elevated, widely interspaced cords that are fine or (usually) strong, sometimes slightly gemmate. Interspaces between cords further sculptured by fine spiral grooves intersected by axial grooves of equal strength, resulting in fine, cancellated background pattern. (Herrmann & Salisbury, 2012) 3 comb. nov., C. manoui (Huang, 2011) 3 comb. nov., D. praestantissima (Röding, 1798) 1 , D. shikamai Habe, 1980 3 , D. sigillata (Azuma, 1965) 3 Remarks. The genus Domiporta is a rather compact and conchologically moderately variable group, however the colour and sculpture pattern in some species are close to those found in imbricariine genera. For example, Domiporta circula and D. praestantissima are superficially very much like Imbricaria as circumscribed here, while D. granatina and D. carnicolor resemble species of Neocancilla. It was probably this general similarity that led Cernohorsky to assign Domiporta to the subfamily Imbricariinae. Nevertheless, the species of Domiporta mentioned can be recognized by the higher spire, more convex whorls, and the presence of a concavity which delimits the shell base from the siphonal canal. Among Mitrinae, Roseomitra n. gen. is closest to Domiporta. The two closely related genera differ, however, in shell proportions: Domiporta is characterized by a notably higher spire and lower aperture compared to Roseomitra Species included: R. citharoidea (Dohrn, 1862) 3 comb. nov., R. earlei (Cernohorsky, 1977) 3 comb. nov., R. fluctuosa (Herrmann & Salisbury, 2013) 3 comb. nov., R. honkeri [START_REF] Poppe | New species of Mitridae and Costellariidae from the Philippines with additional information on the Philippine species in these families[END_REF] 3 comb. nov., R. incarnata (Reeve, 1845) 3 comb. nov., R. millepunctata (G. B. Sowerby III, 1889) 1 comb. nov., R. reticulata (A. Adams, 1853) 3 comb. nov., R. rosacea (Reeve, 1845) 1 comb. nov., R. roseovitta (Huang, 2011) 3 comb. nov., R. strangei (Angas, 1867) 3 comb. nov., R. tagaroae (Poppe, 2008) 3 comb. nov.

Etymology:

The genus name refers to the pink(ish) colour that characterizes some species, such as R. incarnata or R. rosacea. Gender feminine.

Remarks. As the type specimen of Mitra millepunctata is lost [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF] noted that the type specimen cannot be traced, and the first author failed to find it in NHMUK as well), we here designate our sequenced specimen IM-2013-40686, measuring 30.0 x 11.4 mm (Fig. 16 F) as neotype of M. millepunctata. This specimen was collected off Nocnocan Is., Central Philippines, at the type locality of Mitra terryni, a junior synonym [START_REF] Herrmann | A New Neocancilla Species (Gastropoda: Mitridae) from the Indian Ocean and remarks on Neocancilla clathrus (Gmelin, 1791) and Domiporta carnicolor (Reeve, 1844)[END_REF] of M. millepunctata. Siphonal canal rather long, straight, tapering, not notched at its tip. Aperture elongate. Outer lip very thin, convex, smooth or slightly undulating. Inner lip with three columellar folds, usually fine but well pronounced, sometimes very weak. Shell white, pale to yellowish, sometimes covered by light brown periostracum. Radula with laterals and rachidian of equal width, with evenly convex anterior margin bearing 12-14 equal pointed cusps (Figs 17 G,H).

Species of the genus

Distribution: Indo-Pacific, bathyal depths, mostly on mud.

Species included: Profundimitra abyssicola [START_REF] Schepman | The Prosobranchia of the Siboga expedition[END_REF] 3 comb. nov., P. planofilum [START_REF] Huang | Calcimitra, a new genus of deep-water Mitridae (Gastropoda: Mitridae) with the description of five new species from Taiwan and the Philippines[END_REF] 1 comb. nov., P. taylori n. sp. 1 .

Etymology:

The name Profundimitra refers to the fact that all known species of this new genus inhabit deep water. Gender feminine.

Remarks. Whereas all the sequenced species of Calcimitra were identifiable to known species, the new deep-water genus Profundimitra (as well as the related Fusidomiporta) appears to represent a kind of Mitridae terra incognita. Of four species recognized in our phylogenetic analysis, only Profundimitra planofilum has a name. The latter was initially described in Cancilla, and indeed the sculpture pattern and light brown marks on spiral cords of Profundimitra planofilum strikingly resemble the pattern seen in species of Cancilla. P. planofilum, however, differs from Cancilla by the shell base outline, which is distinctly concave, and by a slightly higher spire. The immature lectotype of C. abyssicola figured by [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF] is very close to species of the Profundimitra clade in shell characteristics, and we tentatively assign this species to Profundimitra. It should be noted, though, that C. abyssicola as circumscribed by Cernohorsky combined a variety of forms, some of them (like the specimen figured by Cernohorsky 1970: pl. 6 fig. 7, which is undoubtedly conspecific with our specimens IM-2007-35711 andIM-2007-35618) referable to Profundimitra n. gen., but most likely not conspecific with the lectotype, while others obviously represent true Cancilla or Imbricaria as circumscribed here.

The genus Profundimitra n. gen. can be recognized by the elongate, typically weakly sculptured shell, either white or pale, sometimes covered with light brown periostacum. The genus Calcimitra is closest to Profundimitra n. gen. in shell morphology, however species in the latter genus have proportionally higher spire and, correspondingly, lower last adult whorl and siphonal canal. Another genus which seems superficially close to Profundimitra n. gen. is

F o r R e v i e w O n l y 37
Eumitra, which however is characterized by a smooth columella, typically with no folds or with only one or two distinct folds, while species of Profundimitra n. gen. have 3+ columellar folds, albeit sometimes slightly sunken in the aperture. Another notable difference between the two genera is the morphology of protoconch: it is narrow and narrowly-conical in Profundimitra, very distinctly demarcated from the much wider first teleoconch whorl, whereas the protoconch of Eumitra is very low, cyrtoconoid, and not clearly delimited from teleoconch. Moreover, Profundimitra n. gen. has a characteristic radula morphology with rachidian and laterals of equal width and similar morphology, as opposed to the very small, narrow rachidian and wide laterals of Eumitra. Species included: P. atjehensis (Oostingh, 1939) 1 comb. nov., P. chrysalis (Reeve, 1844) Etymology: Pseudonebularia means "false Nebularia" and refers to the long accepted, and erroneous, placement of the included species in Nebularia, which it resembles superficially.

Genus

Gender feminine.

Remarks. Pseudonebularia n. gen. comprises a group of popular, colourful shallow water or even intertidal miters that are traditionally referred to Nebularia but, as demonstrated by our analysis, are not related to it, as Nebularia is recovered in a sister position to Pterygia. Species of Pseudonebularia n. gen. typically have a small, but broad and robust, brightly coloured shell.

While red and variegated biconical shells in species of the P. fraga group are quite distinctive, more elongated species, for example P. atjehensis, resemble species of Domiporta or Imbricaria as newly circumscribed here. The elongated Pseudonebularia species, however, can be distinguished by generally broader, rounded spiral cords that bear no darker lines on their crests.

The three species of Pseudonebularia studied anatomically show a significant disparity in radular morphology, with rachidian varying considerably from bow-shaped with multiple densely set cusps to very narrow, but robust, bearing a single short, pointed cusps on a bulbous shaft. Species included: Strigatella amaura (Hervier, 1897) 3 , S. assimilis (Pease, 1868) 3 , S. aurantia (Gmelin, 1791) 1 comb. nov., S. auriculoides (Reeve, 1845) Remarks. The sculpture of strong rounded cords of some species of the Strigatella clade may lead to a wrong generic allocation; for instance, [START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF] placed S. vexillum, S. aurantia, S. lugubris and even S. fulvescens in Nebularia, whereas our phylogenetic analysis unequivocally places them in Strigatella.

Genus

There are three conchologically more or less distinct groups of Strigatella, (1) a S. paupercula group with typical broadly fusiform Strigatella shell, (2) a S. fulvescens group with more elongate shell, distinctly recurved siphonal canal and lack of inner callous varix on the outer aperture lip, and (3) a S. coronata group with a fusiform shell also lacking a callous varix on the outer aperture lip, and also distinguished by a uniserial radula. S. coronata and S. lugubris forms a very distinct clade in our phylogenetic tree, and the name Chrysame is available for it.

Given its distinctive morphology and its long branch on the molecular tree, the Strigatella coronata group could be treated as a subgenus of Strigatella. However we prefer to abstain from recognizing Chrysame, as it would necessitate to also recognize as subgenera the other two Strigatella groups -a level of taxonomic splitting that we are not currently prepared to defend. As circumscribed here, Imbricaria appears rather heterogeneous morphologically, with shell varying from conical to narrowly-fusiform, most species having a notably higher spire than I. conularis. Shell sculpture may be faint, like in the type species, or represented by strong elevated cords, and depressions between cords are often sculptured with dense riblets. Moreover, species of Imbricaria display a remarkable disparity in radular morphology, as seen in the degree of cusp differentiation in both the rachidian and lateral teeth. In this respect, the eight studied species form a continuous grade from radulae with subequal cusps (presumably a plesiomorphic state, found in I. conularis and I. pugnaxa -see Discussion) to more derived ones with one or two cusps on each tooth greatly exceeding the others. Remarks. The typification of Cancilla renders the identity of the genus a little weak: the specimen of Mitra isabella originally illustrated by Swainson (1831) was selected as lectotype by [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF], but, like many types of Swainson relevant to this study, this specimen is lost [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF]. Taking in consideration frequent parallelisms in shell morphology demonstrated in Mitridae, the utility of Swainson's original illustration for taxonomic decisions is limited and it will be desirable in the future to designate a neotype with associated molecular data.

Genus

The radula of C. isabella was studied by [START_REF] Salisbury | Notes on Cancilla isabella (Swainson, 1831) (Neogastropoda: Mitridae) with emphasis on the radula and generic assignment within Mitridae[END_REF]. Although the two specimens dissected by these authors differ notably in shell proportions, their radulae are morphologically close to each other, and rather peculiar. The rachidian bears two pairs of long pointed cusps (the lateral cusps being notably weaker than the central ones), flanked by two pairs of very weak cusps; the laterals bear long pointed cusps that progressively increase in size towards tooth mid-width, whereas the distal halves bear no cusps and show only weak serration Taking into account this observed radular disparity among sequenced (and thus confirmed related) Cancilla species, we believe that the divergent radular morphology of Cancilla isabella does not contradict its affinity to the other species studied herein. Species of Domiporta, can be distinguished from the 'robust' Cancilla species by the sculpture pattern with narrow, elevated, widely interspaced spiral cords, whereas a distinct concavity at the transition of shell base to siphonal canal distinguishes species of Domiporta from 'slender' Cancilla species. Moreover, the two genera differ greatly in radular morphology.

Besides, there is a clear resemblance between some species of Imbricaria and 'slender' Cancilla: I. fulgetrum and C. rehderi / fibula (especially our immature specimen IM-2013-19073) or the I. yagurai group and C. cf. fibula (represented by IM-2013-40646). In these pairs, the species of Cancilla can be recognized by the more convex outline of the shell base, which gives the shell a rather fusiform shape compared to the rather biconical shell of Imbricaria. The new genus Cancillopsis can be differentiated from the 'robust' Cancilla species by the higher spire with rather subcylindrical whorls, and from the 'slender' Cancilla species by the presence of a concavity at the base of the siphonal canal. Finally, the radula with rachidian and lateral teeth of similar morphology differentiates Profundimitra n. gen. and Fusidomiporta n. gen. from Cancilla. Remarks. Few mitrids are characterized by an almost involute conical shell, and these were formerly allocated to either Imbricaria or Pterygia. However, we found Imbricaria as traditionally construed to be paraphyletic, with Imbricaria punctata forming a lineage of its own, not closely related to I. conularis. Although a single species has been sequenced, we hypothesize a close relationship between Imbricaria punctata, I. conovula and I. vanikorensis, largely based on shell characters. Imbricariopsis n. gen., as we circumscribe it, constitutes a rather distinctive group of small shallow water mitrids: species of Imbricariopsis can be recognized by conical, weakly sculptured, or not sculptured at all, pale shells with a minute, pointed apex (that is much lower than in Imbricaria conularis). The radulae of the three species that we assign to Imbricariopsis n. gen. were illustrated by [START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF] and are again figured here; all three share the same general morphology of lateral radular teeth, with one robust major and several minor cusps situated on both sides of the major one.

Genus

The enigmatic West African Mitra carbonacea Hinds, 1844 resembles species of Imbricariopsis n. gen. in overall shell morphology, and may prove to be related to this genus.

However, given the disparity in geographical distribution, we place this species in Imbricariopsis n. gen. with a question mark, pending future studies. Minor differences in the position of the columellar folds (situated in the abapical portion of the columella in Imbricariopsis and at midheight in M. carbonacea) may hint at a not-so-close relationship. conularis; based on this resemblance, he placed conoidea in Imbricaria, although in a separate subgenus Sohlia. According to [START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF], "I." conoidea is the earliest known fossil Imbricaria (as circumscribed by him), although it shows some differences to the Recent members of that genus, in particular the presence of only few columellar folds that are "placed rather low on columellar pillar" (Cernohorsky 1970: 44). Noteworthy, the columellar folds, despite numbering five in Imbricariopsis n. gen., are also positioned anteriorly, whereas the species of Imbricaria (as circumscribed herein), are characterized by columelar folds set at midheight of columella. This may indicate that species of Imbricariopsis n. gen. are in fact the closest living relatives of the ancient "Imbricaria" conoidea, which agrees with the rather underived position of Imbricariopsis n. gen. in the imbricariine segment of the mitrid tree. The range of sculpture patterns is equally considerable: whereas most species of Scabricola are characterized by wide and low cords or punctate grooves, some (e.g., Scabricola padangensis) have a "Subcancilla-type" sculpture pattern [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF] of narrow, high cords, and still others, like S. coriacea, show a distinctly crenulated sculpture. In the treatment of Cernohorsky, the more slender species of Scabricola, with weaker sculpture, were separated in the subgenus Scabricola (Swainsonia). It should be noted, though, that the boundaries of the two subgenera appear blurred, primarily because S. variegata (the type species of Scabricola) is conchologically closer to Swainsonia than to many species of Scabricola (Scabricola). One of the important diagnostic characters of Scabricola is the morphology of the radula lateral teeth, unique for the Mitridae, "with 3-5 strong cusps on the inward facing angular plate" (Cernohorsky, 1991: 115) (Figs 28 D -S. variegata). This radular morphology is shared by seven species of Scabricola (Scabricola) and Scabricola (Swainsonia) studied by [START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF], whereas radulae with divergent morphologies are found in S. desetangsii (Fig. 28 E) and S. coriacea (Cernohorsky, 1970, fig. 120): the former has unicuspidate laterals, the latter lacks the major cusp on laterals which, therefore, have a somewhat intermediate morphology between typical Scabricola and Neocancilla.

Genus

Because the boundaries of the genus remain poorly understood, we are only confident about the placement of those species for which molecular and radular morphology data are available, and the remaining species are placed in Scabricola with a question mark. Given this major uncertainty on its contents, we refrain from giving a formal diagnosis of the genus. Remarks. The genus Neocancilla is one of the few lineages the composition of which is left rather unchanged after our phylogenetic analysis. The group is well delineated thanks to the characteristic shell and conservative radular morphology. The radula of the West African "Neocancilla" hebes depicted by [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF] rather resembles the radula of Strigatellinae, and led Cernohorsky to question the relation of that species to the Indo-Pacific Neocancilla. Here we remove this species from Neocancilla and provisionally place it in

Genus

Domiporta.

Species of Domiporta are closest to Neocancilla primarily because of similar shell proportions and sculpture pattern. Species of Neocancilla, however, can be distinguished by the shell base and siphonal canal, which form a continuous, usually slightly convex, outline with no concave waist. Besides, there are consistent differences between the two genera in radular morphology: in Neocancilla the radula has few short, blunt and robust cusps on the rachidian and laterals, while in Domiporta the rachidian and laterals bear multiple pointed cusps that are not differentiated in size or shape. The Scabricola -Swainsonia grade is closely related to Neocancilla, and the species of these three genera are conchologically similar. However, shells of Scabricola and Swainsonia can be differentiated by a notably weaker sculpture, with no or very weak axial elements; besides, most species of Scabricola are wider, with a higher last adult whorl compared to Neocancilla. Distribution: Indo-Pacific, subtidal, mostly on coral rubble, one species occurs on mud.

Species included: Pterygia arctata (G. B. Sowerby II, 1874) 3 , P. conus (Gmelin, 1791) 3 , P. crenulata (Gmelin, 1791) 2 , P. dactylus (Linnaeus, 1767) 1 , P. fenestrata (Lamarck, 1811) 1 , P. gorii [START_REF] Turner | Six new species of Mitridae from the Indian and Pacific Oceans, with remarks on Mitra abacophora Melvill, 1888 (Neogastropoda: Muricoidea)[END_REF] 3 , P. japonica Okutani & Matsukuma, 1982 3 , P. jeanjacquesi Bozzetti, 2010 3 , P. nucea (Gmelin, 1791) 2 , P. purtymuni Salisbury, 1998 3 , P. scabricula (Linnaeus, 1767) 3 , P. sinensis (Reeve, 1844) 1 , P. undulosa (Reeve, 1844) 3 .

Remarks. The genus Pterygia is morphologically distinctive thanks to its ovate shell with multiple columellar folds, and monoserial radula. In the present study we leave the contents of the genus unchanged; however, some species currently classified in other genera (for example, Scabricola potensis) may actually belong to Pterygia. Conical shells resembling those of Pterygia are found in Imbricaria and Imbricariopsis n. gen., which are however phylogenetically very distant from Pterygia, and are characterized by a triserial radulae. 3 , N. semperi [START_REF] Poppe | New species of Mitridae and Costellariidae from the Philippines with additional information on the Philippine species in these families[END_REF] 3 comb. nov., N. thachi [START_REF] Turner | Six new species of Mitridae from the Indian and Pacific Oceans, with remarks on Mitra abacophora Melvill, 1888 (Neogastropoda: Muricoidea)[END_REF] 3 , N. ustulata (Reeve, 1844) 3 comb. nov.

Remarks.

Our phylogenetic analysis has led to a considerable rearrangement of the genus Nebularia. The "lesser" Nebularia species (N. fraga and N. tornata groups) appeared unrelated to the true Nebularia typified by N. contracta. On the other hand, some large Mitra species (M. incompta, M. eremitarum) showed a close affinity to N. contracta, and so did 'Strigatella' acuminata. The genus Nebularia, as circumscribed herein, is supported by several distinctive morphological shell (colour pattern, a subsuturally thickened outer aperture lip, and an often recurved tip of the siphonal canal) and radula characters (a very strong unpaired central cusp on the rachidian). The same features characterize a number of species classically included in Mitra (M. ancillides, M. coarctata, M. edentula, M. ferruginea, M. nebulosa, M. nivea, M. ustulata), which we thus classify in Nebularia. As a consequence, we also consider the nominal genera Dibaphus, Mitroidea and Mutyca to be synonyms of Nebularia. Both shell proportions and development of sculpture vary considerably among the species of Nebularia, the two extremes being N. incompta (Fig. 33 D) and N. ancillides (Fig. 33 G). An even more cylindrical shell characterizes Mauritia barclayi, which we tentatively place in Nebularia as well.

Subfamily Isarinae Fedosov, Herrmann, Kantor & Bouchet, new subfamily (Figs 34 -36)

Diagnosis: Shell small to large (15-125 mm) mm, biconical, fusiform to turriform, pale to dark brown due to well developed periostracum. smooth, or sculptured with fine to very strong spiral cords,. Suture distinct, impressed. Spire moderately to very high; teleoconch whorls gently convex, distinctly shouldered due to pronounced spiral sculpture, or flattened. Sculpture absent, or represented by regular spiral cords, ranging from wide and flattened to rounded, or narrow, elevated, separated by wide, concave interspaces. Microsculpture of fine collabral growth lines.

Siphonal canal short and stout with well developed fasciole, to moderately long, strait and tapering. Siphonal notch rarely deep, commonly shallow or absent. Aperture elongate, varying from narrow to moderately wide; outer aperture lip evenly convex or straight in its adapical portion and strongly convex anteriorly. Inner lip with four columellar folds, the anteriormost notably weaker than succeeding. Rachidian narrow, less than half as wide as the laterals, bearing 5 or 6 cusps of varying shape (Figs 33 A -C). Laterals wide, with 8-40 cusps, of varying strength. , Cox (1936: 66). Pallary referred to the type species as "Mitra (Fuscomitra) fusca (Swainson) Reeve", i.e. the species identified by Reeve (1844) as Mitra fusca Swainson, 1824. Reeve did not establish a new species, and there is no such thing as "Mitra fusca Reeve, 1844". Instead, this is a case of a deliberate misidentification of a nominal species used for the fixation of the type species and, under Art. 67.13, Pallary is deemed to have established a new nominal species, Mitra fusca Pallary, 1900.

Diagnosis: Shell medium-sized to large (30-100 mm), fusiform to turriform, lightly sculptured, light or dark brown due to well developed periostracum. Suture distinct, impressed. Spire high to very high; teleoconch whorls gently convex to flattened, sculptured by regular or occasional spiral grooves, or smooth, often with a microsculpture of fine collabral growth lines. Siphonal canal short with well developed fasciole to moderately long and tapering. Siphonal notch shallow or absent. Aperture elongate, rather wide; outer aperture lip evenly convex or straight in its adapical portion and strongly convex anteriorly. Inner lip with four columellar folds, the anteriormost notably weaker. Rachidian narrow, less than half as wide as the laterals, bearing 5 or 6 cusps of varying shape (Figs 33 A -C). Laterals wide, with 8-40 cusps, subequal and densely set throughout, or proximal strongest and subsequent ones weakening towards tooth edge.

Species included: Isara aerumnosa (Melvill, 1888) 3 comb. nov., I. aikeni [START_REF] Lussi | Endemic deep water Mitridae of South Africa with the description of three new species[END_REF] Remarks. The type species of Isara, Mitra bulimoides (Fig. 34 B), was considered by [START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF] a synonym of M. glabra, and indeed shows conchological resemblance to it. [START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF] provided no whereabouts of the type specimen of Mitra glabra Swainson, 1821, and designated two unnumbered figures on plate 2 of the "Exotic Conchology" as the lectotype of M. glabra. Although this designation is nomenclaturally valid, ICZN article 74.4 makes it clear that this designation is to be treated as designation of the specimen illustrated. As this specimen cannot currently be traced, we herein designate the holotype of Mitra bulimoides NHMUK 1966656 as neotype of M. glabra. Thus Mitra bulimoides Reeve, 1845 becomes an objective synonym of the Mitra glabra Swainson 1821.

The genus Isara, as circumscribed herein, is a group largely defined based on molecular characters. Although the included species share a similarity in shell morphology (weak sculpture and, in many cases, well developed periostracum), the shell proportions are drastically different in I. straminea and I. glabra, and their relatedness would hardly be guessed based on overall shell appearance. The sequenced Isara species altogether show an unusually broad distribution: Azores and Mediterranean (I. cornea), West Africa (I. nigra), Caribbean (I. straminea) and Western Australia (I. glabra and I. carbonaria). Based on the overall shell morphology we tentatively assign Mitra chinensis to Isara, as well as an array of Australian species (M. badia, M. chalybeia, M. peterclarcksoni and M. slacksmithae).

The species of Isara are commonly large, and their protoconchs are often corroded in (sub)adults; we managed to find an intact protoconch only in juveniles of I. chalybeia from southwestern Australia. It is characterized by a shape unusual for Mitridae -paucispiral, cyrtoconoid, broad, with a very small nucleus, and strongly convex whorls. I. chalybeia lives in subtropical / warm temperate waters and we doubt that this protoconch morphology also occurs in the tropical species of Isara, which are expected to have the plesiomorphic multispiral protoconch present throughout the family.

The pantropical distribution of Isara does not include the East Pacific. Whereas no "smooth" Atrimitra species from the Panamic province was included in our phylogenetic analysis, the obvious resemblance between the Panamic "Mitra" orientalis, and "M." caliginosa, and the Atlantic Isara nigra and I. cornea, should be noted. Another species which also remains unstudied, and may prove to be related to Isara, is Atrimitra idae. At present we treat these New World species as Atrimitra, pending availability of molecular data. Remarks. [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF] remarked that the placement of the New World Subcancilla sulcata and of Indo-Pacific species in the same genus Subcancilla was inconsistent with their disparate radular morphology, however no alternative was suggested. Thorsson and Salisbury (2008: 3) stressed that the classification of the Panamic and Caribbean species of Subcancilla, including the type species S. sulcata, has not been addressed sufficiently, and that the "exact placement of Subcancilla into the family Mitridae remain[ed] to be absolutely determined". Based on our phylogenetic analysis we demonstrate a close affinity of three East Pacific and one Caribbean species of Subcancilla, and reconsider the contents of the genus, so that only New World species are here assigned to it. The species of Subcancilla can be readily distinguished from other American mitrids by their characteristic sculpture of narrow, elevated, usually widely interspaced, spiral cords, often bearing a brown line on their crests; the same sculptural pattern characterizes Indo-Pacific species in the genera Domiporta and Imbricaria.

Members of these three genera are also characterized by similar shell proportions, so that their shells share the same characteristic habitus that has long confused conchologists. Our results demonstrate that the three genera represent three major phylogenetic lineages of Mitridae in the subfamilies Isarinae, Mitrinae and Imbricariinae, and their resemblance is obviously a result of convergence.

The radula differs remarkably in the three studied species, with those of S. attenuata and S.

cf. leonardhilli being overall unremarkable underived mitrid radulae, whereas that of S. erythrogramma demonstrates signs of cusp differentiation, resembling imbricariine radulae The small and unconspicious radula of S. sulcata pictured by Cernohorsky (1991: fig. 79) undoubtedly belongs to the first type.

Subfamily Pleioptygmatinae Quinn, 1989

Genus Pleioptygma Conrad, 1863

Type species: †Voluta carolinensis Conrad 1840: 387; M.

Diagnosis: As no material on Pleioptygma was available for the present study, we do not provide our own diagnosis to this enigmatic and poorly studied genus, and refer to [START_REF] Quinn | Pleioptygmatidae, a new family of mitriform gastropods (Prosobranchia: Neogastropoda)[END_REF].

Species included: Pleioptygma helenae (Radwin & Bibbey, 1972) 2 .

Distribution: Caribbean, subtidal to upper bathyal depths.

Remarks. For remarks on the placement of Pleioptygma, see section 'Implications of the morphological studies for the taxonomy of the Mitridae' below. Etymology: The name Condylomitra combines the Latin word condylus, meaning knob, with the suffix Mitra, in reference to the characteristic shell sculpture of tubercles that is shared by the species in this newly recognized lineage. Gender feminine.

Incertae sedis

Genus

Remarks. Although each on a quite long branch, Condylomitra bernhardina and C. tuberosa form a clade in our molecular trees and share features of the shell and radula, and the characteristic sculpture pattern allows a rather easy recognition of this group. Because of their superficial similarity, we initially considered Strigatella lugubris and "Mitra" pudica to be possibly related to Condylomitra; however the former is closely related to Strigatella coronata in the Strigatella clade, and the latter, although it was not sequenced, is excluded from Condylomitra by its triserial radula.

Magnamitra Huang & Salisbury, 2017

Type species: Magnamitra sandrogorii Huang & Salisbury, 2017; OD.

Diagnosis: As no material on Magnamitra sandrogorii was available for the present study, we do not provide our own diagnosis to this newly described genus, and refer to [START_REF] Huang | Magnamitra n. gen. and nomenclatural remarks on large Cancilla and Mitra from Taiwan and the Philippines (Gastropoda: Mitridae)[END_REF]. Species included: Probata barbadensis (Gmelin, 1791) 1 comb. nov., P. espinosai (Sarasúa, 1978) 3 .

Remarks. Mitra espinosai, the type species of Probata, has been considered a junior synonym of Mitra barbadensis by [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF]. Indeed, the immature holotype of the former very much resembles Mitra barbadensis and, even if not conspecific, they are definitely closely related. Thus we tentatively allocate the name Probata to the Mitra barbadensis clade. The affinities of Probata remain unclear and obviously need further studies. In terms of sculpture pattern, typical Atrimitra, as typified by Mitra idae, resembles Probata; however the latter Diagnosis: As no material on Dibaphimitra florida was available for the present study, we do not provide our own diagnosis to this poorly studied genus, and refer to [START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF].

Description

Shell medium sized (holotype 24.15 mm, paratype 35.5 mm), fusiform, with high, cyrtoconoid, slightly tilted spire and rather short siphonal canal. Protoconch pointed, subcylindrical, of 2+ whorls (corroded in all specimens studied). Teleoconch of 7.5 gently convex whorls, suture linear, distinctly impressed. First teleoconch whorls with reticulate sculpture formed by four low, very weak, rounded spiral cords, intersected by even weaker rounded spiral folds. Interstices rather deep, forming characteristic pattern of four spirally aligned punctures, well pronounced on two succeeding whorls and rather gentle, widely spaced on later whorls. Penultimate whorl with three distinct spiral rows of punctures on adapical portion, followed by three very weak rows below. Last adult whorl evenly convex, gradually extending to rather short, straight tapering siphonal canal. Adapical portion and periphery of last adult whorl sculptured with ten spiral rows of punctures. On shell base spiral rows of punctures becoming denser, as well as punctures themselves merging to form narrow grooves on abapical part of shell base and strong, oblique grooves on siphonal canal.

Siphonal canal slender, moderately long, tapering, not notched at its tip. Aperture rather low, lanceolate, its outer lip thin, evenly convex. Inner apertural lip evenly concave in its adapical half and straight below, with 3 oblique columellar folds at mid-height, slightly deepened in aperture and weak in appearance. Remarks: The identification of Fusidomiporta ponderi n. sp. is uneasy, mainly because of its striking resemblance to (still undescribed) species of Profundimitra n. gen., represented in our analysis by specimens IM-2007-35618 andIM-2007-35711 [both mislaid]. The single live taken specimen of F. ponderi n. sp. seems to have a proportionally slightly lower spire and a somewhat stepped appearance of spire whorls, with more elevated spiral cords. At the same time, the distinctive and heavy sculpture pattern makes it quite recognizable from among most Mitridae. Some deep water species of Gemmulimitra n. gen. have a somewhat similar, beaded sculpture, which however is nowhere as strong as in Fusidomiporta ponderi n. sp. Besides, Gemmulimitra boucheti has a proportionally broader shell with more convex whorls compared to F. ponderi n.

sp. On the contrary, G. neocaledonica n. sp. has a rather biconical shell with subcylindrical whorls and a rather reticulate sculpture pattern. However, at this moment, many of the deep water mitrids are still represented in our dataset by too few specimens to assess their intraspecific variation and elaborate robust shell delimitation criteria.

Gemmulimitra neocaledonica

Type data

Holotype. MNHN IM-2013-40655, New Caledonia, Kouakoué Canyon, EXBODI Stn 3821, 21°53'S, 166°50'E, 211-440 m, lv, 24.25 mm (Figs 21 A, B).

Paratypes: New Caledonia, EXBODI, Stn 3800, East coast off Thio, 21°32'S, 166°22 'E,[279][280][281][282]1 lv (Paratype 3,21.5 mm); Stn 3807, East coast off Toupeti, 21°43'S, 166°36 'E, 352-372

Other material examined

New Caledonia, EXBODI Stn 3795, 21°32'S, 166°21 'E, 240-245m, 1 lv (16.4 mm); Stn 3800, 21°32'S, 166°22 'E, 279-282m, 12 lv (16.7 mm, 17.1 mm, 17.3 mm, 15.0 mm, 16.2 mm, 16.2 mm, 16.55 mm, 16.8 mm, 15.05 mm, 23.1 mm, 17.65 mm, 17.8 mm); Stn 3805, 21°42'S, 166°34 'E, 302 m, 1 lv (20.9 mm); Stn 3806, 21°42'S, 166°34 'E, 307-309 m, 7 lv (22.0 mm, 18.4 mm, 15.9 mm, 16.3 mm, 14.0 mm, 9.15 mm, 9.8 mm) (22.6 mm, 20.1 mm, 21.05 mm, 20.7 mm, 20.0 mm, 19.2 mm, 17.4 mm, 20.9 mm, 18.1 mm, 18.2 mm, 19.65 mm); Stn 3810, 21°44'S, 166°38 'E, 384-385 m, 8 lv (25.5 mm, 25.85 mm, 29.3 mm, 25.5 mm, 27.4 mm, 21.65 mm, 25.5 mm, 23.3 mm); Stn 3814, 21°49'S, 166°44 'E, 331-344 m, 6 lv (22.8 mm, 21.5 mm, 20.6 mm, 21.6 mm, 18.9 mm, 14.1 mm); Stn 3821, 21°53'S, 166°50 'E, 211-440 m, 2 lv (25.85 mm, 25.15 mm); Stn 3823, 21°55'S, 166°55 'E, 246-255m, 2 lv (15.5 mm, 14.5 mm); Stn 3825, 21°58'S, 166°59 'E, 349-405m, 1 lv (16.2 mm); Stn 3828, 22°00'S, 167°01 'E, 300-302m, 1 lv (16.4 mm); Stn 3829, 22°02'S, 167°05 'E, 350-360m, 1 lv (23.0 mm); TERRASSES Stn 3095, 22°02'S, 167°06 'E, 320-380 m, 1 lv (21.5 mm).

Papua New Guinea, BIOPAPUA Stn CP3634, 07°29'S, 147°31 'E, 279-290 m, 1 dd (21.75 mm).

Description

Shell small to medium-sized (holotype 24.25 mm), elongate-fusiform to almost biconical, heavily sculptured, with narrow aperture and stepped appearance of late teleoconch whorls.

Protoconch pointed, cyrtoconoid, translucent, of 3+ slightly convex, glossy whorls. Protoconch / teleoconch transition distinct. Teleoconch of about 9 whorls; suture canaliculated. Subsutural ramp forming distinct shelf, flat or even slightly inclined inside, giving teleoconch whorls a subcylindrical profile. Whorl periphery slightly convex, early spire whorls sculptured with three strong, closely set, spiral cords overriding low indistinct axial ribs to form series of prominent, axially aligned, beads. On third teleoconch whorl, adapical spiral cord splits into two, and succeeding spire whorls sculptured with four gemmate cords, adapical cord bordering subsutural ramp wavy, slightly narrower than others, delineated from second cord by narrower interspace.

On later teleoconch whorls, interspaces between gemmate spiral cords gradually broadening, axial ribs becoming more distinct and sculpture rather reticulate. Last adult and penultimate whorls with 26 axial ribs each. Interspaces between axial ribs and spiral cords forming deep quadrangular depressions with microsculpture of fine co-axial growth lines, sometimes retaining fragments of periostracum forming squamiform projections. Shell base extended to moderately long, tapering, slightly notched siphonal canal. Four gemmate spiral cords on adapical portion of last adult whorl, and 12 cords on shell base and siphonal canal. According to our earlier results [START_REF] Fedosov | Phylogeny and systematics of mitriform gastropods (Mollusca: Gastropoda: Neogastropoda)[END_REF], the families Mitridae, Charitodoronidae and Pyramimitridae form a well supported phylogenetic group, which we designate as a superfamily Mitroidea, although they are rather heterogeneous morphologically, as summarized in Table 4.

The morphological distinctiveness of the Mitridae has been appreciated by many authors and, when anatomical descriptions scattered in many papers are brought together, the family ranks among the better studied among neogastropods. Beside multiple illustrations of radulae [START_REF] Risbec | Contribution à l'étude anatomique de quelques espèces de mitres de la presqu'ile de Nouméa[END_REF][START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF][START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF][START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF], data on the general body morphology and/or foregut anatomy are available, among Mitrinae, for Mitra mitra [START_REF] Ponder | The morphology of some mitriform gastropods with special reference to their alimentary canal and productive systems (Mollusca: Neogastropoda)[END_REF], Ziba carinata [START_REF] Simone | Anatomical description of Ziba carinata from Ghana (Caenogastropoda, Mitridae)[END_REF], Pseudonebularia cucumerina [START_REF] Risbec | Contribution à l'étude anatomique de quelques espèces de mitres de la presqu'ile de Nouméa[END_REF] and Episcomitra zonata (Vayssière 1901); among Strigatellinae, for Strigatella paupercula [START_REF] Ponder | The morphology of some mitriform gastropods with special reference to their alimentary canal and productive systems (Mollusca: Neogastropoda)[END_REF], S. retusa and S. scutulata [START_REF] Risbec | Contribution à l'étude anatomique de quelques espèces de mitres de la presqu'ile de Nouméa[END_REF]; among Imbricariinae, for Imbricariopsis conovula [START_REF] Ponder | The morphology of some mitriform gastropods with special reference to their alimentary canal and productive systems (Mollusca: Neogastropoda)[END_REF]; among Cylindromitrinae, for Pterygia crenulata [START_REF] Risbec | Contribution à l'étude anatomique de quelques espèces de mitres de la presqu'ile de Nouméa[END_REF]; among Isarinae, for Isara cornea [START_REF] Harasewych | Anatomy and biology of Mitra cornea Lamarck 1811 (Mollusca, Caenogastropoda, Mitridae) from the Azores[END_REF]; and for Atrimitra idae (West 1990), A. catalinae (West 1991), and Condylomitra tuberosa (herein), that we treat as incertae sedis. All the species that were studied with sufficient attention revealed an epiproboscis (referred to as "tube à venin" by [START_REF] Risbec | Contribution à l'étude anatomique de quelques espèces de mitres de la presqu'ile de Nouméa[END_REF], either in combination with a radula of underived morphology (in most of the species mentioned) or with a uniserial radula (Pterygia crenulata, Condylomitra tuberosa). Meanwhile, an epiproboscis is not present in the Charitodoronidae, the Neogastropoda clade closest to the Mitridae. There is no doubt that the epiproboscis represents an autapomorphy of the Mitridae and can be used for the anatomical circumscription of the family.

In this connection, the morphology of the Caribbean Pleioptygma helenae, as addressed by [START_REF] Quinn | Pleioptygmatidae, a new family of mitriform gastropods (Prosobranchia: Neogastropoda)[END_REF], deserves special attention. Quinn's description of the foregut anatomy raises many questions; in particular the position of the buccal mass is not described explicitly, as well as the nature of the "proboscis bulb", and we believe that the homologies of these organs may have been misinterpreted by him. This assumption is further reinforced by the fact that the two specimens dissected by Quinn displayed some variation in foregut morphology, which may be an artifact of poor fixation (with the use of rhum suggested by the author!). Some parts of the description are of special interest. In particular, the "proboscis bulb attached to the cephalic cavity floor by a broad, rather thick band of muscles originating in the foot" (Quinn 1989: 14) more likely refers to the large buccal mass and a radula / odontophore retractor muscle. Then the peculiar introvert, which "invaginates and runs back through the outer tube / sheath and enters the proboscis bulb [?=buccal mass]" may not be anything but an epiproboscis. Undoubtedly, the anatomy of Pleioptygma helenae is peculiar and its homologies need to be clarified based on investigations of additional material. However, given its rarity and the small chance of obtaining live-taken specimens in the near future, we dare reassess the systematics of Pleioptygma based on currently available data. Several lines of evidence support a placement of Pleioptygma as a separate subfamily within the Mitridae, including 1) the mitriform shell; 2) the radula with rachidian and laterals of about equal morphology -similar to the one in Domiporta and 3) the presence of a structure that, based on topology and morphology, is closely comparable to the mitrid epiproboscis, and most likely is an epiproboscis.

Evolution of radular morphology in the Mitridae

Our studies revealed only two major radula types in the family Costellariidae, with a very simple scenario of two independent transitions from plesiomorphic to derived [START_REF] Fedosov | Systematics, diversity and evolutionary history of the family Costellariidae (Gastropoda: Neogastropoda)[END_REF]). The situation is incomparably more complex in the Mitridae. Here we recognize six major types of radula; each is refered to a genus that typically represents it (Fig. 41).

1 -Mitra-type: radula with laterals notably wider than rachidian tooth and bearing multiple equal or subequal cusps. This underived radula morphology is widespread across the Mitridae tree. It characterizes the family Charitodoronidae, most Mitrinae (except the Domiporta group and some Pseudonebularia), most Isarinae, as well as the incertae sedis genera Probata, Carinomitra, Atrimitra [START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF]West 1990;1991) and Dibaphimitra [START_REF] Bayer | Observations on Mitra florida Gould[END_REF]. 1a -Strigatella-type: radula with slightly curved laterals bearing short robust cusps on their medial convex portion and lacking cusps laterally; rachidian always with central unpaired cusp. This type of radula is found in all Strigatellinae (except in the S. lugubris -S. coronata clade). The

Strigatella-type of radula represents a modified Mitra-type, and intermediate morphologies with

Strigatella-like laterals but different rachidian are found outside Strigatellinae in Neotiara nodulosa (Mitrinae) and Cancilla schepmani (Imbricariinae). 1b -Nebularia-type: is also a slightly modified version of the Mitra-type, with a characteristic rachidian bearing only five cusps, the central unpaired cusp being notably enlarged. This type of morphology characterizes the genus Nebularia as circumscribed herein.

2 -Profundimitra-type: very wide rachidian, roughly attaining the width of laterals and bearing equal number of cusps. This type of radula characterizes the mitrine genera Profundimitra and Fusidomiporta, and is also found in some Domiporta (but not in its type species, D. filaris), Pseudonebularia maesta and, outside Mitrinae, in the genus Pleioptygma [START_REF] Quinn | Pleioptygmatidae, a new family of mitriform gastropods (Prosobranchia: Neogastropoda)[END_REF].

3 -Imbricaria-type: radula characterized by moderately wide multicuspidate laterals, with one of the medial cusps notably enlarged compared to the others. Cusps of laterals are also differentiated in size, with a central unpaired cusp retained, although often reduced in size. This type of radula is found in all Imbricaria species except I. fulgetrum. 72 3a -Cancilla-type: resembles the Imbricaria-type, but the laterals are more than twice as wide as the rachidian, and the central unpaired cusp of the rachidian is absent. This type of radula is found in Cancilla isabella [START_REF] Salisbury | Notes on Cancilla isabella (Swainson, 1831) (Neogastropoda: Mitridae) with emphasis on the radula and generic assignment within Mitridae[END_REF] and other species of this genus studied herein, as well as in Imbricariopsis, Subcancilla erythrogramma and Imbricaria fulgetrum.

4 -Scabricola-type: radula characterized by strongly modified laterals in which the lateralmost cusp is greatly enlarged to form a robust spine. Typically, the rachidian is comb-like (Scabricola variegata, Swainsonia spp.) but it may also bear a single strong cusp. This type of radula is found in species of Scabricola and Swainsonia. Although the diversity of radular morphologies in the family Mitridae is impressive, the understanding of its evolution is greatly hampered by homoplasies. Radula morphology does not clearly align with the inferred phylogenetic groupings of the Mitridae (Fig. 41) because of an ubiquitous retention, in advanced lineages, of the plesiomorphic state and because of multiple convergences. In fifteen genera of Mitridae, all studied species presented an underived Mitratype radula. Seven genera appeared heterogeneous in radula morphology, combining several general types. Thus specific apomorphies can be identified in only few cases, and in even fewer cases the apomorphic state is shared by all members of the lineage. Only eight genera (Profundimitra, Fusidomiporta, Neocancilla, Pterygia, Nebularia, Condylomitra and, with some reservations, Imbricariopsis and Swainsonia) are supported by distinct apomorphies in radular morphology. Among them, only Nebularia and Neocancilla show autapomorphic radula types.

Undoubtedly, the subfamily Imbricariinae shows the greatest diversity of radular morphologies, and the pattern of radular evolution is most obvious and consistent in that subfamily. The progressive differentiation of cusps on both the laterals and the rachidian is noteworthy, along with a general tendency to the reduction of the number of cusps. The radulae of Cancilla spp. are closest to the underived Mitra-type, and alternate courses of radula transformation are observed in the Imbricaria and Scabricola-Neocancilla clades.

In addition to a blurred phylogenetic signal of radula characters, our understanding of radula evolution in the Mitridae is impeded by a lack of evidence on the functionality of different

Taxonomy of Mitridae: historical considerations and new arrangement

The current Mitridae species list (as indexed in WORMS consulted on May 30 2017) comprises 398 accepted Recent species, plus the three new species described in the present work. The baseline of Mitridae systematics is the fundamental revisions by Walter Cernohorsky, who first dealt with the subfamily Mitrinae [START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF]), and later with the Imbricariinae and Cylindromitrinae [START_REF] Cernohorsky | The Mitridae of the World. Part 2. The Subfamily Mitrinae Concluded and Subfamilies Imbricariinae and Cylindromitrinae[END_REF]). Cernohorsky's monographs had a profound impact on mitrid taxonomy and systematics, despite the general lumping attitude prevailing in his time, which resulted in an extensive taxonomic graveyard for many of the species level taxa he accepted as valid. Several cases of abusive synonymization are demonstrated in the present study. For instance, Mitra morchii A. Adams, 1855, considered by Cernohorsky a synonym of Cancilla isabella, is, based on [START_REF] Huang | Magnamitra n. gen. and nomenclatural remarks on large Cancilla and Mitra from Taiwan and the Philippines (Gastropoda: Mitridae)[END_REF] and on our data, not related to Cancilla or even to Imbricariinae. Another example is Mitra millepunctata G.B. Sowerby III, 1889, synonymyzed [START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF] with Domiporta carnicolor despite notable disparity in shell proportions, and later described as Mitra terryni Poppe, 2008. Altogether the number of valid species of Mitridae was greatly diminished by Cernohorsky, but the rate of species description increased notably in the following decades and over 100 species (i.e. almost 30% of the currently accepted number of species) were described since 1991. This burst of activity in mitrid taxonomy is also notable for being almost entirely accounted for by the amateur community: only ten out of 112 species described in the last 25 years were described by academics. Amateurs perhaps more reluctantly establish new supraspecific taxa, and only two genera Calcimitra [START_REF] Huang | Calcimitra, a new genus of deep-water Mitridae (Gastropoda: Mitridae) with the description of five new species from Taiwan and the Philippines[END_REF]Magnamitra Huang &[START_REF] Huang | Magnamitra n. gen. and nomenclatural remarks on large Cancilla and Mitra from Taiwan and the Philippines (Gastropoda: Mitridae)[END_REF] were established in the last 25 years, whereas 65 species described over this period were originally placed in Mitra.

A concise review of changes in mitrid genus-level taxonomy was presented by [START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF], in a study that itself contained the description of four new genera and subgenera did not cluster with any other lineage and were not successfully sequenced for genetic markers other than COI. Thus, they at present remain "hanging" in the list of unallocated Mitridae and may potentially represent two more new genera.

Phylogeographic patterns in the Mitridae

It has been widely known that the Indo-West Pacific harbours the greatest diversity of mitrid species and lineages, and therefore a good representation of IP localities at various depths was a primary requirement to our work. 88 of the 103 inferred species of Mitridae and Charitodoronidae in our dataset originate from Indo-Pacific localities. Besides, three species were sampled from the Mediterranean and NE Atlantic, two from West Africa, four from the Caribbean, and six from the Panamic province (Fig. 42). 23 mitrid genera were sampled in the Of all studied genera, Charitodoron probably demonstrates the narrowest range, being confined to deep waters in the Mozambique Chanel and off South Africa. According to some authors (e.g., [START_REF] Obura | Evolutionary mechanisms and diversity in a western Indian Ocean center of diversity[END_REF], starting in the Eocene, this area served as a main refuge for relict lineages of Tethyan origin which, for some reason, did not give rise to new radiations. The long branch that separates Charitodoron on the molecular tree, characters such as the underived morphology (with only the radula indicating its relatedness to Mitridae), and its low diversity in the Recent fauna, all point to the relict nature of Charitodoron. The paucispiral, bulbous protoconch of Charitodoron indicates non-planktotrophic development, which correlates well with its restricted distribution.

Conversely, the distribution of Isara is the widest among mitrid genera: this is the only genus recorded in more than two major zoogeographical regions of the shelf (as defined by Briggs and Bowen 2012) -Indo-West Pacific, East Atlantic and West Atlantic (Fig. 42) -, and it may be also present in the East Pacific (see Remarks under Isara). Isara species contribute significantly to mitrid diversity in the peripheral Indo-Pacific (South Australia, presumably South Africa) (see Fig. 8), but they are rare and not really diverse in the Central Indo-West Pacific. In our understanding, Isara is undoubtedly an old and underived lineage of Mitridae, and this pattern may be interpreted in two different ways. According to one possible scenario, it was once widely distributed in tropical seas and subsequently was replaced by younger and derived lineages in the Central Indo-West Pacific while maintaining its diversity in peripheral areas. The other possible scenario suggests that the primarily Indo-Pacific Isara, once "forced" into subtropical waters by growing competition with younger evolutionary lineages of Mitridae, has adapted to new temperatures and, through this, was capable of spreading beyond the biogeographical limits of the Indo-Pacific. Although most known species of Strigatella occur in the Indo-Pacific, an amphi-Pacific distribution characterizes this genus, with at least one species inhabiting shallow water in the Panamic province ending up close to the Indo-Pacific species in both molecular and morphological characters. Finally, if 'Mitra' hebes is related to the species of Domiporta, as suggested by shell and radular morphology, the range of that genus would also include West Africa.

The generally low overlap in regional lists of mitrid genera may be interpreted as a consequence of the relatively late major diversification of mitrid lineages, dating back to the time when modern biogeographical barriers were already established, and largely separate We have attempted to circumscribe the bathymetric distribution of the newly delineated genera, based primarily on the locality data of sequenced specimens. Given the patchiness of our data, we arbitrarily divided the sampled depth range from 0 to about 1800 meters into four depth intervals: 1) from 0 to 40 meters, 2) from 41 to 80 meters, 3) from 81 to 300 m and 4) >300 m.

Of the 103 species of Mitridae studied herein based on molecular characters, 72 were sampled from the 0-40 meters interval, seven from the 41-80 meters interval, nine from the 81-300 meters interval, and 20 from depths in excess of 300 meters. This accounting does not, however, strictly reflects changes in species richness with depths, as it is strongly biased by the distribution of collecting efforts (Fig. 43). Intertidal and upper subtidal zones (at diveable depths) as well as outer slopes from about 200 down to about 1800 meters were sampled incomparably better than the mesophotic and abyssal zones. Therefore our data do not necessarily demontrate a drop in species richness in the 41-300 m interval. Nevertheless, despite the sampling bias, there is an obvious pattern of greater mitrid diversity in shallow water, decreasing with depth. This pattern can be explained by a greater diversity of habitats, often fragmented and intermixed at a small scale, in shallow water ecosystems. Another related factor is the multitude of biological interactions in these shallow habitats, driven by the complex ecological structure of reefs and associated habitats, providing a wealth of niches to be explored by mitrids and their preys.

The bathymetric distribution of the newly delineated mitrid genera, is shown on Figure 43 based on our results (black vertical bars) and literature (grey vertical bars). Because of the very fragmentary data, some of the displayed results are inconsistent, like the disjunct bathymetric range of Gemmulimitra and Subcancilla. We in fact fully expect that many genera, which in our results are restricted to the 0-40 meters depth interval, in fact reach deeper, probably down to 80-120 meters. Nevertheless, some general conclusions can be drawn from the observed bathymetric distribution of the genera. The representation of mitrid genera in shallow water is notably higher than at depths in excess of 100 -150 meters, which mirrors the distribution of species diversity discussed above. Whereas Gemmulimitra, Imbricaria, Isara, Roseomitra, and Subcancilla have extensive bathymetric ranges, with congeneric species sampled from the intertidal down to 300-400 meters, the mitrid fauna from the greatest depths is essentially represented by specialized lineages (treated here as genera), that do not occur in shallower water.

The genera Calcimitra, Cancilla, Cancillopsis, Fusidomiporta and Profundimitra represented in our dataset by 15 species (plus 2 species of Charitodoron) and the genus Eumitra (not included in our phylogenetic studies) constitute several separate radiations that thrive in the deep sea.

World diversity of Mitridae

Of 103 species included in our molecular dataset, 89 were identified with confidence -these constitute 22% of the described diversity of the family Mitridae. Of the remaining 15 species, four were either not identified ("sp.") or were attributed to a described species with an indication of some disparity in morphological or molecular characters (referred to as "cf."); ten proved to be unnamed -although three of them showed some similarity to described species and were allocated to tentative species complexes (referred to as "aff."). Remarkably, of the 60 mitrid species from the Indo-Pacific sampled in the 0-40 meters depth interval only one (Strigatella aff. paupercula), represented a new species -the notable divergence of our material from the otherwise similar Strigatella paupercula was only revealed in the phylogenetic analysis. The proportion of unidentified / undescribed species grows with increasing depth and, of 20 species from depths greater than 300 meters in tha analysis, only 11 were identified with confidence, and seven (i.e. 35%) were undoubtedly new, all representing previously undescribed lineages of Mitrinae. Three of these species were here described as Profundimitra taylori n. gen. and sp., Fusidomiporta ponderi n. gen. and sp., and Gemmulimitra neocaledonica n. gen. and sp. Thus, an estimate of 30 to 100 species of Mitridae remaining to be described from the deep waters of the Indo-Pacific seems reasonable to us. Recognition of new deep water mitrid species may be hampered by the high intraspecific variability of shell sculpture and, simultaneously, rampant convergences leading to hardly distinguishable shells in not closely related species. Likewise, the radula appears generally useless for species delimitation, as in most cases it retains plesiomorphic morphology, and the significance of slight variations in the number of cusps is unknown. Thus, molecular data will remain essential for a reliable recognition of new deep water taxa of Mitridae.

Whereas a significant proportion of the deep water mitrids remains undescribed, the overall diversity of the family in deep water pales in comparison with the well documented diversity of the family in shallow water. These contrasting proportions of described Mitridae in shallow versus deep water can be viewed as a result of recent efforts by collectors and amateur taxonomists. Since the first half of the 19 th century and the iconic monograph by Reeve (1844Reeve ( -1845)), the comparatively easily accessible species from shallow water are collected steadily throughout many Indo-Pacific locales and then carefully scrutinized by a dedicated community of amateurs, leading to the present day state of mitrid taxonomy, with a pretty well inventoried shallow water fauna. Undoubtedly, there are areas with high local endemism that still hold undescribed species, especially in peripheral locales (e.g., southern Australia, South and East Africa and the Arabian Sea), but this is not the general situation. The diversity of mitrids in the Mediterranean is low, and all three species known from that sea were included in the present study. Conversely, we have included only a limited number of species from West Africa and the New World, which makes any judgment on diversity in these regions untimely. Two Panamic species in our dataset were not confidently identified but were represented by subadults, the identification of which is always troublesome, and the uncertainty of our identifications cannot be viewed as an indication of undescribed species. A more thorough sampling in both the Panamic and Caribbean region is definitely required to assess the proportion of undescribed species in these areas and uncover the relationships of their endemic phylogenetic lineages. [START_REF] Lozouet | Eumitra récentes de la région néo-calédonienne et Charitodoron fossiles de l'Oligocène supérieur d'Aquitaine (Mitridae)[END_REF][START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF]1991;[START_REF] Huang | Magnamitra n. gen. and nomenclatural remarks on large Cancilla and Mitra from Taiwan and the Philippines (Gastropoda: Mitridae)[END_REF] NMW.1955.158.00066, Point Loma, Lower California, Mexico, 60.0 mm;B. Atrimitra effusa, lectotype, NHMUK 1966415, Guacomayo, Central America, 31.6 mm;C. Atrimitra caliginosa, holotype, NHMUK 1966718, locality unknown, 34. [START_REF] Lozouet | Eumitra récentes de la région néo-calédonienne et Charitodoron fossiles de l'Oligocène supérieur d'Aquitaine (Mitridae)[END_REF][START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF]1991;[START_REF] Huang | Magnamitra n. gen. and nomenclatural remarks on large Cancilla and Mitra from Taiwan and the Philippines (Gastropoda: Mitridae)[END_REF]. Numbers next to each bar indicate number of species involved. Blue horizontal bars show the proportion of stations in each depth interval.

F o r R e v i e w O n l y

99x55mm (300 x 300 DPI)

  2017, 180 of the 398 accepted Recent species of Mitridae indexed in WoRMS

  separated into five and six unlinked partitions respectively: 16S, 12S, three codon positions of the COI gene (both datasets) plus H3 gene (4G and 4G-red datasets).

8

 8 Abbreviations of repositories and facilities used in text

9

  Decisions on the generic assignment of individual species are based on: 1 -molecular evidence and radula morphology, or molecular data only; 2 -shell characters and radula morphology;

  COI, appeared sufficiently divergent from other inferred lineages to exclude possible conspecificity. Therefore, our dataset includes 103 species as defined by both morphological and molecular characters.

  1.2) a rather supported Mitra sanguinolenta -M. sophiae -M. cardinalis -M. puncticulata clade (PP=1; B=59); 1.3) a Domiporta grouping, moderately supported in BA (PP=0.98), combining (1.3a) a distinctive four species Domiporta s.s. clade (PP=1; B=100); (1.3b) a two species Mitra . rosacea clade (PP=100; B=1); and two clades formed by undescribed deepwater mitrids: (1.3c) IM-2013-40651 and IM-2013-40654 (PP=1; B=100), and (1.3d) the singlespecies clade of IM-2013-40665.

  Figs 3 -5) consists of three distinct, well supported clades that include 20 Indo-Pacific species commonly classified in the Imbricariinae: ) a first clade (PP=1, B=99) combining Imbicaria conularis, two species of Subcancilla, and five species of shallow-water Ziba from the Indo-West Pacific, Z. bantamensis, Z. flammigera, Z. fulgetrum, Z. insculpta and Z. verrucosa; 3.2) a second clade (PP=1, B=100) including the morphologically variable deep-water species Cancilla fibula, represented in our analysis by three specimens, and Swainsonia schepmani; 3.3) a third, distinctive and well supported, grouping including: (3.3a) a well supported Neocancilla clade, with (3.3b) weakly supported Scabricola clade showing supported affinity to Neocancilla, (3.3c) the two-species Swainsonia clade and (3.3d) Imbricaria punctata, which is recovered as a sister group to the rest of 3.3.

  Mitridae lineage comprises a single species, the shallow water Caribbean Mitra barbadensis.

  veneris. Foregut anatomy based on NMSA D4172 [off Durban, 29°50.4'S, 31°14.0'E, 150 m, sandstone gravel, 14.1 mm], radula based on NMSA E9237 [northern Natal, off Shellfield Beach, 29°39.8'S, 31°30.16'E, 150 m].

  D, arrow), both chambers lined with tall, densely ciliated epithelium, epithelial cells with large basal or central nuclei. Wall of the posterior chamber bearing very tall longitudinal folds (Figs 6 D

  D). Strong and massive odontophore retractor adjoining the columellar muscle shortly behind the nerve ring. Radular diverticulum opening into buccal cavity ventrally in posterior position, behind thick circular fold of the buccal cavity wall,

  ). Compact buccal mass situated shortly after mouth, furnished with thick muscular walls and triangular lumen; radular diverticulum opening ventrally into buccal cavity (Figs7 B, C). Anterior oesophagus quadrangular on transversal sections, lying dorsally in the anterior part of the proboscis, attached to proboscis wall by thin muscular fibers (Figs7 D -H). Dorsal and ventral walls of oesophagus very thin, lateral walls with two large longitudinal folds jutting out into the lumen of oesophagus. Lateral folds of oesophagus disappearing at mid proboscis height and thick layer of muscular fibers evenly covering it (Fig.7I). Radular diverticulum rather short; odontophore formed by two small cartilages (Figs7 D, E), not fused anteriorly. Odontophore retractor very strong, lying in a muscular sheath in its anterior portion (Figs7 F -H), and dividing into two symmetrical branches at mid proboscis length (Fig.7 I).Epiproboscis short, formed by densely packed longitudinal muscle fibers, with no epithelium. Epiproboscis lying ventrally from radular diverticulum, enclosed inside its own muscular sheath, which is attached to the radular diverticulum along most of its length (Figs7 D -H). Both epiproboscis and its sheath originating from the ventral part of the radular retractor 7 G, H). Epiproboscis sheath opening ventrally on the anterior border of the buccal mass.

  (A) Based on the inclusion of Mitra mitra (type species of Mitra, itself the type genus of Mitrinae), we designate the Mitra-Domiporta-Ziba-Episcomitra lineage as subfamily Mitrinae. Within it, the Mitra mitra-Mitra papalis clade is distinct and well supported, and shows no supported affinity to any other clade within the Mitrinae; thus we restrict the genus Mitra to this Mitra mitra-Mitra papalis clade (with a few other, presumed closely related, species). The genus Tiarella thus becomes a junior synonym of Mitra. The numerous species (even including Mitra cardinalis !) traditionally classified in Mitra that fall outside this clade are assigned below to other genera. (B) The Neocancilla-Imbricaria-Scabricola lineage, well supported in all phylogenetic analyses, includes the type species of Imbricaria, I. conularis, thus we designate it as subfamily Imbricariinae. (C) Pterygia (with its subjective synonym Cylindromitra) is the only genus classified in the subfamily Cylindromitrinae

  Conversely, there is no name available for the Isara-Subcancilla major lineage, for which we establish below the new subfamily Isarinae.

  n. gen. and sp. for the single-species clade of MNHN IM-2013-40665 (Indo-Pacific, deep water); Profundimitra n. gen. for the Indo-Pacific deep water clade of MNHN IM-2013-40651 and MNHN IM-2013-40654;

  the name Probata to the single species clade of "Mitra" barbadensis, following[START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF], although we question the synonymy of Mitra espinosai (type species of Probata) with M. barbadensis, as suggested by Cernohorsky.

  the conservative shell shape with rather high spire and low aperture, presence of denticles on the outer aperture lip and a recognizable colour pattern. At the same time, its radular morphology is overall the same as in many other mitrid clades and adds little to the diagnosis of the genus.

  Calcimitra Huang, 2011 (Figs 10 B-C, 11) Type species: Calcimitra kingtsio Huang, 2011; OD.

  Neotiara Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Figs 10 D, E, 13) Type species: Mitra lens W. Wood, 1828; OD, herein.

Diagnosis:

  , elongate to fusiform, usually light brown to olive.Protoconch narrowly conical of about 2.5-3 smooth convex whorls. Suture distinct, impressed.

  Episcomitra Monterosato, 1917 (Figs 10 G, H, 14 E -H) Type species: Mitra zonata Marryat, 1818; M.Diagnosis: Shell small to large (15-110 mm), fusiform or elongate fusiform, smooth, with welldeveloped periostracum. Protoconch large, paucispiral, cyrtoconoid, of about 2 smooth glossy convex whorls. Spire tall or rather low; suture impressed. Spire whorls evenly convex in outline, sculpture represented by fine, dense, collabral lines. Last adult whorl typically evenly convex with rather short siphonal canal bearing a distinct fasciole, separated from shell base by deep notch deep or rather shallow. Outer aperture lip evenly convex, or convex in its adapical portion and straight throughout most of its length. Inner aperture lip with four, fine, subequal columellar folds. Shell covered with periostracum, light brown to dark brown or olivegreen, uniform or with multiple dark freakles, or (typically) adapical 2/3 of spire whorls slightly lighter than their abapical portions. Inside of aperture white or orange. Rachidian with 6-8 pointed cusps, increasing in length towards center; laterals wide, bearing rather long and pointed cusps throughout their length (Figs10 G, H).

  Quasimitra Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Figs 10 I -M, 15) Type species: Mitra sanguinolenta Lamarck, 1811; OD, herein. Diagnosis: Shell small to large (20-90 mm), solid, broadly fusiform, vividly coloured. Protoconch pointed, narrowly-conical, of about 3.5 slightly convex, smooth whorls. Suture deeply impressed. Shell proportions variable, spire moderately high to rather low, with ortho-or cyrtoconoid profile. Spire whorls evenly convex to subcylindrical, sometimes distinctly shouldered. Sculpture of very weak and low, flat or rounded, cords separated by shallow depressions or fine grooves. Subsutural area smooth or with one row of short spiny warts or tubercles, giving whorl a somewhat shouldered appearance. Siphonal canal short, with well developed fasciole usually separated from shell base by narrow concave waist. Siphonal notch deep. Aperture elongate, wide or moderately wide. Outer lip smooth, inner lip with 4 distinct columellar folds. Rachidian narrow (1/3 of laterals width) to rather wide (exceeding half of

  sculpture and colour pattern, although varying notably in shell proportions. Cernohorsky treated Vicimitra prosphora Iredale, 1929 as a junior synonym of Mitra solida, despite V. prosphora has a smooth, uniformly pale shell, and only superficially resembles Mitra solida and the other

  Siphonal canal moderately long, tapering or robust, with well developed fasciole, delimited from shell base by distinct narrow waist, or extended concave stretch of shell base. Aperture rather short, elongate, with distinct siphonal notch. Outer lip gently convex, inner lip with 3-4 fine columellar folds, subequal or adapical strongest. Shell usually white or light with contrasting spiral cords, either uniformly dark brown, or dashed-brown. Rachidian from narrow (about 40% of laterals width) with 11 short, blunt cusps to almost as wide as laterals, arched, with 20+ pointed cusps. Laterals broad, bearing 20-35 equal, pointed cusps set on narrow, straight or arched, plate (Figs 17 A -D). Distribution: Indo-Pacific, perhaps West Africa (D. hebes), shallow subtidal to bathyal depths, on sand. Species included: Domiporta aglais 3 (Li & Zhang, 2005), D. carnicolor (Reeve, 1844) 1 , D. circula (Kiener, 1838) 1 comb. nov., D. daidaleosa (Li & Li, 2005) 3 comb. nov., D. filaris (Linnaeus, 1771) 1 , D. gloriola (Cernohorsky, 1970) 3 , D. granatina (Lamarck, 1811) 1 , D.hebes(Reeve, 1845) 2 comb. nov., D. latistriata(Herrmann & Salisbury, 2012) 3 comb. nov., D.

  n. gen. Four species of Domiporta depart notably in radular morphology, showing a transition from rather typical mitrine radula in D. filiaris to the remarkable radulae of D. granatina and D. praestantissima with arched laterals and rachidian of equal width, bearing 25+ cusps; an intermediate morphology is observed in D. cf. carnicolor. Genus Roseomitra Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Figs 16 F -I, 17 E, F) Type species: Mitra millepunctata G. B. Sowerby III, 1889; OD, herein. Diagnosis: Shell small to medium-sized (10-45 mm) thin-walled, fusiform with rather low spire and high aperture. Protoconch pointed, narrowly conical, of 3+ smooth, glossy, slightly convex whorls. Suture distinct, slightly impressed. Spire rather low, orthoconoid; spire whorls with convex outline, sculptured with narrow, rounded, widely set cords. Interspaces between spiral cords bearing finer and lower secondary cords, or fine, widely interspaced, axial grooves. Last adult whorl very high, evenly convex, with shell base extended without clear distinction into straight, stout siphonal canal. Siphonal fasciole not pronounced, siphonal notch shallow or indistinct. Aperture high and moderately wide, its outer lip evenly convex or strongly convex adapically and nearly straight in its mid-width. Inner lip with five fine folds, adapicalmost strongest and succeeding ones gradually diminishing. Shell white, cream or pink, with brown blotches or with regular brown dots or dashes on spiral cords. Radula with narrow rachidian bearing 5-8 subequal cusps and wide laterals with narrow base and 15-21 moderately long pointed cusps (Figs 17 E, F).

  Indo-Pacific, shallow subtidal to bathyal depths, on sand or mud.

  Roseomitra n. gen. can be recognized by the combination of rather thin walled shell with proportionally low spire and high aperture, distinctly rounded whorls and Domiporta-like sculpture. The shell proportions with low spire differentiate Roseomitra n. gen. from closely related Domiporta and most other mitrid genera. Species of Pterygia and Imbricariopsis have a proportionally even higher aperture, but in these genera the shell is conical or ovate, whereas it is fusiform in Roseomitra n. gen. In turn, Condylomitra tuberosa, C. bernhardina and Quasimitra sophia have a notably stronger sculpture compared to Roseomitra spp. Genus Profundimitra Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Figs 17 G, H, 18 A -H) Type species: Profundimitra taylori n. sp.; OD, herein. Diagnosis. Shell small (20-35 mm), thin-walled, fusiform or elongate-fusiform, with high orthoconoid or slightly acuminate spire. Protoconch pale, semi-translucent, narrowly conical with 3+ smooth whorls [usually strongly eroded]. Suture adpressed, sometimes shallowly canaliculated. Teleoconch whorls gently convex, early teleoconch whorls sometimes subcylindrical. Sculpture variable, from very weak, represented by indistinct spirally aligned deep spiral grooves separating flat, rounded or distinctly gemmate spiral cords.

  Fusidomiporta Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Figs 17 I, 18 I -L) Type species: Fusidomiporta ponderi n. sp.; OD, herein. Diagnosis: As the genus is established primarily based on the molecular data, and its recognition based on the morphology is problematic, we do not provide a formal diagnosis. Description (based on type species): Shell small (about 25 mm), fusiform, heavily sculptured, with rather high spire and stout siphonal canal. Protoconch multispiral, of 3+ whorls usually strongly corroded or missing. Suture canaliculated; subsutural shelf well developed, rather narrow, bordered by projecting margin of adapical spiral cord. Sculpture of strong, elevated, gemmate spiral cords separated by deep interspaces; microsculpture of fine growth lines in interspaces between spiral cords. Spiral cords become distinctly beaded on penultimate and last adult whorls, forming rows of closely set, axially expanded, glossy granules. Siphonal rather long and stout, slightly recurved. Aperture elongate, its outer lip gently convex throughout its height, wavy, reflecting arrangement of spiral cords. Inner lip strongly calloused, reflected, with three strong oblique columellar folds at its mid-height. Rachidian and laterals multicuspidate, of about same width. Distribution: Indo-Pacific, bathyal depths in about 400-500 m. Species included: F. ponderi n. sp. 1Etymology: The name Fusidomiporta reflects the close affinity of the new genus to Domiporta, and refers to the fusiform shell of the type species. Gender feminine.

  Morphologically, Fusidomiporta ponderi n. sp. falls within the range of variation seen in Profundimitra n. gen., both in the shell and radula, and we even initially thought that specimen IM-2013-40665 (now the holotype of F. ponderi) was conspecific with IM-2007-35711 (an unknown species of Profundimitra). The molecular distinctiveness of Fusidomiporta ponderi n. sp. was thus totally unexpected. However, none of the single-gene analyses placed Fusidomiporta ponderi n. sp. in the same molecular clade as Profundimitra, and a Fusidomiporta-Profundimitra grouping is only present in the 3 genes based tree. Thus the genus is largely defined by molecular characters; the morphological basis for the separation between Fusidomiporta and Profundimitra needs to be more carefully elaborated, for which more specimens need to be examined. A sculpture pattern close to that present in Fusidomiporta n. gen. is also observed in species of Gemmulimitra n. gen. (G. neocaledonica n. sp. and G. boucheti); however, Gemmulimitra differs notably from Fusidomiporta in radula morphology, showing a classical mitrid radula with a narrow rachidian (Figs 20 F, G). Genus Cancillopsis Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Fig. 18 M) Type species: Cancilla meimiaoae Huang & Salisbury 2017; OD, herein. Diagnosis: As the genus is established primarily based on the molecular data, and its recognition based on the morphology is problematic, we do not provide a formal diagnosis. Description (based on the type species, adapted from Huang & Salisbury (2017): Shell mediumsized (50-82 mm), fusiform, with high spire and long, stout siphonal canal. Protoconch multispiral, cyrtoconoid, of about 3 glossy whorls. Teleoconch of up to 10 gently convex whorls; suture distinct. Sculpture of rounded spiral cords, intersected by closely set, slightly prosocline axial riblets, forming fine beaded sculpture on early spire whorls. On last adult whorl cords more widely set, and beaded sculpture indistinct, as axial riblets are well pronounced in interspaces between spiral cords only. Siphonal canal stout, deeply notched, slightly recurved at its tip. Aperture elongate, its outer lip evenly convex throughout its length. Inner lip calloused, with four closely set fine columellar folds at its mid-height. Shell white with irregular yellow or brownish blotches. Distribution: Indo-Pacific, bathyal depths. Species included: Cancillopsis meimiaoae (Huang & Salisbury 2017) 1 comb. nov., †C. acuminata (Shuto, 1969) 3 comb. nov.

  The name Cancillopsis refers to the close resemblance between C. meimiaoae, the type species of the new genus, and species of Cancilla. Gender feminine.Remarks. Cancillopsis meimiaoae shows a notable resemblance to the species of the imbricariine genus Cancilla as well as to the species of Profundimitra n. gen. and Fusidomiporta n. gen., while showing no phylogenetic affinity to either of them. A visible concavity at the transition of shell base to the siphonal canal distinguishes the new genus from similarly sculptured Cancilla species. We hypothesize that Cancillopsis n. gen. is more closely related to the Domiporta group of Mitrinae, and so we provisionally place it in Mitrinae. However, a more comprehensive taxon sampling of the deep water Mitridae is obviously needed to delineate these convergent deep water lineages and to clarify affinities of Cancillopsis n. gen.GenusPseudonebularia Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Figs 19, 20 A -E) Type species: Mitra tornata Reeve, 1845; OD, herein. Diagnosis: Shell small (9-30 mm), fusiform or broadly fusiform to biconical, vividly coloured. Protoconch pointed, narrowly conical, of about 3 slightly convex, glossy whorls. Suture slightly impressed, often indistinct. Teleoconch whorls evenly convex or flattened, sometimes giving the spire a distinctly stepped profile. Sculpture of strong, rounded spiral cords, regularly interspaced and covering the entire shell or, rarely, limited to shell base. Interspaces between cords concave, smooth or bearing fine, dense riblets. Siphonal canal moderately long, stout, siphonal notch shallow or absent. Aperture rather narrow, elongate, with undulating outer lip, sometimes bearing rounded denticles on its inside. Inner lip calloused, often reflected, with 3-4 strong columellar folds, adapicalmost strongest. Radula with broad laterals, bearing equal, rather long, widely set cusps throughout their width; rachidian not exceeding half the width of the laterals, multi-or unicuspidate (Figs 20 A -E). Distribution: Indo-Pacific, intertidal to upper subtidal depths, rocky flats, reef crevices and coral rubble.

  Gemmulimitra Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Figs 20 F, G, 21) Type species: Gemmulimitra neocaledonica n. sp.; OD, herein. Diagnosis: As the genus is established primarily based on the molecular data, and its recognition based on the morphology is problematic, we do not provide a formal diagnosis. Description (based on the type species): Shell small (about 20-25 mm), elongate-fusiform to almost biconical. Protoconch multispiral, of 3+ slightly convex, glossy whorls. Suture

  ramp forming distinct shelf, giving spire a stepped appearance. Spire whorls sculptured with strong, closely set, spiral cords overriding low indistinct axial ribs to form series of prominent beads. On later teleoconch whorls, interspaces between gemmate spiral cords broadened, with axial ribs becoming more distinct and sculpture rather reticulate.Interspaces between axial ribs and spiral cords forming deep quadrangular depressions with microsculpture of fine axial growth lines. Shell base extended to rather long, tapering, slightly notched siphonal canal. Aperture narrow, elongated. Outer lip wavy, convex in adapical portion and straight below mid-height; inside of outer lip smooth. Inner lip strongly calloused, sometimes reflected, with four closely set fine columellar folds. Rachidian narrow, bearing 8 strong, moderately long, pointed cusps; laterals wide, bearing 16+ strong, pointed, rather widely set cusps.Distribution: Indo-Pacific, subtidal to bathyal depths.Species included: Gemmulimitra aliciae(Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., G. apprimapex[START_REF] Poppe | New species of Mitridae and Costellariidae from the Philippines with additional information on the Philippine species in these families[END_REF] 3 comb. nov., G. avenacea (Reeve, 1845) 1 comb. nov., G. boucheti(Cernohorsky, 1988) 2 comb. nov., G. duplilirata(Reeve, 1845) 3 comb. nov., G. edgari (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., G. gonatophora (Sturany, 1903) 3 comb. nov., G. hansturneri (E. Guillot de Suduiraut & E.G. Guillot de Suduiraut, 2009) 3 comb. nov., G. margaritata (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., G. neocaledonica n. sp. 1 , G. rubiginosa (Reeve, 1844) 3 comb. nov., G. strongae (Poppe, Tagaro & Salisbury, 2009) 1 comb. nov.Etymology:The genus name refers to the sculpture of gemmate cords that characterizes many species in this newly recognized lineage, including the type species G. neocaledonica.Remarks. Gemmulimitra neocaledonica n. sp. and G. avenacea that together constitute the Gemmulimitra clade on multi-gene trees have little in common in shell morphology, and thus this clade is at present defined solely based on the molecular characteristics. The third species confidently included in the genus is Gemmulimitra strongae that is close to G. neocaledonica n.sp. in the COI based tree. The other species here attributed to Gemmulimitra are tentatively classified in this genus based on their general resemblance to the type species or to Mitra avenacea, rather than on any specific diagnostic feature. Therefore, the species included in Gemmulimitra constitute two conchologically distinctive groups: species in the first group have a typical sculpture of gemmate cords, such as Gemmulimitra neocaledonica n. sp., G. boucheti 21 C), G. strongae (Fig.21 D), and G. gonatophora (holotype figured by[START_REF] Albano | An illustrated catalogue of Rudolf Sturany's type specimens in the Naturhistorisches Museum Wien, Austria (NHMW): Red Sea gastropods[END_REF], while species in the second group have continuous, smooth spiral cords, such as G. avenacea (Fig.21 E), G. edgari, and G. rubiginosa (Fig.21 F). Species of the first group closely resemble Fusidomiporta ponderi n. sp., whereas species of the second group resemble species of Pseudonebularia, from which, however, they can be distincuished by a more elongated shell with more flattened spire whorls. The radula of Gemmulimitra is plesiomorphic and phylogenetically uninformative. More studies are needed to better define the boundaries of Gemmulimitra n. gen. and elaborate its diagnosis. Subfamily Strigatellinae Troschel, 1869 (Figs 22, 23) Genus Strigatella Swainson, 1840 Type species: Mitra zebra Lamarck, 1811 [= Voluta paupercula Linnaeus, 1758]; SD, Gray (1847: 141). Synonyms: Chrysame H. Adams & A. Adams, 1853; Type species: Mitra coronata Lamarck, 1811; SD, Cox (1927: 91). Phaeomitra Martens, 1880; Type species: Mitra fulva Swainson, 1829 [= Mitra coffea Schubert & Wagner, 1829]; SD, Coan (1966: 131). Diagnosis: Shell small (12-40 mm), broadly fusiform, solid and stout in appearance, Protoconch pointed, narrowly-conical, of about 2.5-3 smooth, very slightly convex whorls. Suture deeply impressed. Spire whorls gently convex or flattened to subcylindrical, sometimes with narrow subsutural area, giving spire a somewhat telescopic outline. Sculpture typically not developed; sometimes shell sculptured with very faint to strong rounded spiral cords over all the shell surface or limited to shell base. Last adult whorl usually strongly convex, siphonal canal typically short and stout to moderately long, with distinct siphonal fasciole. Siphonal notch shallow or absent. Aperture elongate, narrow; outer lip straight adapically and strongly convex in its abapical portion, often thickened adapically, forming a callous inner varix, constricting aperture. Inner lip with 3-5 fine oblique columellar folds. Shell orange to dark brown, typically with a pattern of white bands and blotches. Radula tri-or monoseriate. In triseriate radulae, rachidian narrow, with 5-7 very short and robust cusps, unpaired central cusp always present (Fig 23); in uniseriate radulae, rachidian bearing 20-25 pointed cusps with unpaired central cusp distinctly exceeding others. Laterals broad, wavy, with their distal halves smooth and 7-10 rather short and robust cusps situated proximally.

  Indo-Pacific and Panamic provinces, intertidal and subtidal depths, mostly on hard substrates (rocky shores, reef crevices or coral rubble).

Genus

  Imbricaria Schumacher, 1817 (Figs 24, 25 A -H) Type species: Imbricaria conica Schumacher, 1817 [unnecessary substitute name for Mitra conularis Lamarck, 1811]; M. Synonym: Conoelix Swainson, 1821. Type species: Conoelix lineatus Swainson, 1821 [= Mitra conularis Lamarck, 1811]; OD. Diagnosis: Shell small to medium-sized (10-55 mm), fusiform, narrowly biconical to nearly conical, uniformly coloured or with variegated pattern. Protoconch narrowly conical, of 3+ slightly convex or flattened, smooth and glossy whorls. Early teleoconch whorls forming narrowly conical apex; later whorls widening at equal rate, resulting in fusiform or biconical shell, or (typically) at increasing rate, resulting in acuminated spire and slightly inflated or conical last adult whorl. Early spire whorls sculptured with rounded spiral cords; late whorls smooth or with spiral cords, the latter wide and rounded or fine and sharp, sometimes bearing regular tubercles. Interspaces between cords smooth or with fine, dense riblets. Siphonal canal tapering, with distinct siphonal notch. Aperture elongate. narrow to slit-like. Outer apertural lip convex in its adapical portion; inner lip with 4-5 fine columellar folds gradually diminishing towards siphonal canal. Rachidian with 4-7 cusps, with 1-2 marginal pairs of cusps notably weaker than the 2 or 3 medial cusps; unpaired central cusp when present equal or weaker than 2 flanking cusps (Fig 25 A -H). Lateral teeth broad, with 7-12 triangular or spindle-like, long and pointed subequal cusps, gradually increasing in size towards rachidian, with one cusp strongest, sometimes very long, greatly exceeding others, resembling a tin opener (Figs 25 D -F). Distribution: Indo-Pacific, intertidal to bathyal depths, mostly on soft bottoms, mud, silty or well washed sand. Species included: Imbricaria amoena (A. Adams, 1853) 3 comb. nov., I. annulata (Reeve, 1844) 3 comb. nov., I. armonica (T. Cossignani & V. Cossignani, 2005) 3 comb. nov., I. astyagis (Dohrn, 1860) 3 comb. nov., I. bacillum (Lamarck, 1811) 3 comb. nov., I. baisei (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., I. bantamensis (Oostingh, 1939) 1 comb. nov., I. bellulavaria (Dekkers, Herrmann, Poppe, Tagaro, 2014) 3 comb. nov., I. cernohorskyi (Rehder & Wilson, 1975) 3 comb. nov., I. cloveri (Cernohorsky, 1971) 3 comb. nov., I. conularis (Lamarck, 1811) 1 , I. flammea (Quoy & Gaimard, 1833) 3 comb. nov., I. flammigera (Reeve, 1844) 1 comb. nov., I. fulgetrum (Reeve, 1844) 1 comb. nov., I. hrdlickai (Salisbury, 1994) 1 comb. nov., I. insculpta (A.

  ) 1 comb. nov., I. interlirata(Reeve, 1844) 2 comb. nov., I. nadayaoi (Bozzetti, 1997) 3 comb. nov., I. philpoppei (Poppe, Tagaro & Salisbury, 2009) 3 comb. nov., I. pretiosa (Reeve, 1844) 2 comb. nov., I. pugnaxa[START_REF] Poppe | New species of Mitridae and Costellariidae from the Philippines with additional information on the Philippine species in these families[END_REF] 1 comb. nov., I. rufilirata (Adams & Reeve, 1850) 3 comb. nov., I. ruberorbis (Dekkers, Herrmann, Poppe, Tagaro, 2014) 3 comb. nov., I. rufogyrata (Poppe, Tagaro & Salisbury, 2009) 1 comb. nov., I. salisburyi (Drivas & Jay, 1990) 2 comb. nov., I. tahitiensis (Herrmann & Salisbury, 2012) 3 comb. nov., I. verrucosa (Reeve, 1845) 1 comb. nov., I. yagurai, (Kira, 1959) 1 comb. nov., I. zetema (Dekkers, Herrmann, Poppe & Tagaro, 2014) 3 comb. nov. Remarks. The genus Imbricaria is, in the present classification, undergoing a dramatic rearrangement: whereas species traditionally classified in Imbricaria, like I. punctata and I. olivaeformis, that are conchologically close to I. conularis, are excluded from the genus, numerous Indo-Pacific species earlier assigned to Ziba and Subcancilla are now transferred to Imbricaria.

  Cancilla Swainson, 1840 (Figs 25 I -M, 26)Type species: Mitra isabellaSwainson, 1831; SD, Herrmannsen (1846: 166).Diagnosis: Shell small to rather large (15-110 mm), fusiform, with high aperture and long, tapering siphonal canal. Protoconch conical, of about 3 very slightly convex smooth whorls.Spire whorls evenly convex to subcylindrical; suture distinct, impressed. Shell sculptured with spiral cords that are slightly gemmate or with finely dentate margins, very wide and flat, separated by narrow grooves, or with strong, narrow and elevated cords separated by broad depressions bearing regular, very fine riblets. Shell base gradually extended into rather long, tapering siphonal canal. Siphonal fasciole not pronounced, notch deep or shallow. Aperture elongate, narrow; outer aperture lip smooth, gently convex adapically, and flattened in its lower lip with four oblique columellar folds, adapicalmost strongest. Shell pale or cream, typically with multiple darker spiral strokes on crests of spiral cords. Radula with narrow rachidian, usually bearing four cusps, of which central pair notably stronger (Figs25 I -M).Lateral teeth with 7-10 pointed triangular cusps, second proximal cusp greatly exceeding others, and subsequently gradually weakening towards margins of radula.Distribution: Indo-Pacific, bathyal depths, mostly on mud.Species included: Cancilla baeri(Turner & Cernohorsky, 2003) 1 , C. fibula Poppe, Tagaro & Salisbury, 2009 1 , C. heinickei (Salisbury & Guillot de Suduiraut, 2003) 3 comb. nov., C. herrmanni Dekkers, 2014 3 , C. isabella (Swainson, 1831) 2 , C. liliformis Huang & Salisbury 2017 3 , C. rehderi (Webb, 1958) 1 , C. schepmani (Salisbury & Guillot de Suduiraut, 2003) 1 comb. nov., C. turneri Poppe, Tagaro & Salisbury, 2009 3 .

(

  Salisbury & Huang 2015: figs 1.1, 3.2), a pattern comparable to the laterals in Scabricola, which are however still more robust. Of all Cancilla species studied by us, C. baeri is probably closest to C. isabella: the specimen of C. baeri MNHN IM-2013-58853 (Fig. 26 A) resembles one of the two specimens of C. isabella studied by Salisbury and Huang (2015: fig. 3.2b) in shell proportions, whorl outline and shape of the siphonal canal. The other studied Cancilla species -the C. rehderi -C. fibula complex and C. schepmani -are generally smaller, with less convex whorls and stouter siphonal canal, which is not clearly delineated from the shell base. Therefore, the genus Cancilla, as we is rather heterogeneous, and can be subdivided by shell morphology into two conventional groups: 'robust' Cancilla with species, such as C. isabella, C. baeri, and C. chuoi, and 'slender' Cancilla, with species like C. fibula, C. rehderi, C. schepmani and C. liliformis. The relationships of these two conchological groups is supported by a) inclusion of C. baeri in the Cancilla clade (clade 24-7) in the COI based tree, and b) occurrences of very similar radular morphologies in 'slender' species of the C. fibula -C. rehderi complex (Figs 25 I, K) and in 'robust' C. baeri IM-2013-58853 (Fig. 25 L). It should be noted, however, that the radula of C. schepmani (Fig. 25 M), with a morphology closest to the radulae of Strigatella or of the mitrine Neotiara nodulosa (Fig. 10 E), is thoroughly different from the radulae of other studied Cancilla species. Therefore, Cancilla appears to be very heterogeneous in radular morphology as well.

  Imbricariopsis Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Figs 27, 28 A) Type species: Conoelix punctatus Swainson, 1821; OD, herein. Diagnosis: Shell small (10-20 mm), conical, uniformly pale or with dark tip of siphonal canal. Protoconch pointed multispiral, narrowly conical, dark. Suture distinct, impressed. Spire very short, less than 20% of total shell height, conical or distinctly acuminate, starting with several subcylindrical whorls. Last adult whorl very tall, sculptured with regular, distinctly punctate, spiral grooves. Siphonal notch shallow or moderately deep. Aperture very tall and narrow, slit-aperture lip thick, flattened, smooth inside; inner lip with five fine oblique folds situated on abapical portion of columella. Radula triserial; rachidian with six cusps, paired central cusps greatly exceeding those situated laterally (Fig. 28 A). Lateral teeth with five cusps, second cusp from rachidian very robust, notably exceeding the others. Distribution: Indo-Pacific, intertidal and upper subtidal depths, on sand patches and in reef crevices. Species included: Imbricariopsis conovula (Quoy & Gaimard, 1833) 2 comb. nov., I. punctata (Swainson, 1821) 1 comb. nov., I. vanikorensis (Quoy & Gaimard, 1833) 2 comb. nov., ?I. carbonacea (Hinds, 1844) 3 comb. nov. Etymology: The name Imbricariopsis reflects the resemblance of the species in this newly recognized lineage with the type species of Imbricaria, I. conularis, that has long distracted mitrid taxonomists. Gender feminine.

  highlighted the notable resemblance of the Upper Cretaceous "Imbricaria" conoidea (Matheron, 1843) with Recent Imbricaria species, in particular, I.

  Scabricola Swainson, 1840 (Figs 28 D -F, 29 A -F) Type species: Mitra serpentina Lamarck, 1811 [=Voluta variegata Gmelin, 1791]; SD, Gray (1847: 141). Diagnosis: see under Remarks. Distribution: Indo-Pacific, upper subtidal depths, on sand. Species included: Scabricola bicolor (Swainson, 1824) 3 , S. desetangsii (Kiener, 1838) 1 , S. olivaeformis (Swainson, 1821) 1 , S. padangensis (Thiele, 1925) 2 , S. variegata (Gmelin, 1791) 1 , ?Scabricola barrywilsoni (J. Cate, 1968) 3 , ?S. albina A. Adams, 1853 3 , ?S. caerulea (Reeve, 1844) 3 , ?S. condei Guillot de Suduiraut, 2001 3 , S. coriacea (Reeve, 1845) 2 , ?S. dampierensis Salisbury & Heinicke, 1998 3 , S. dianneae (Salisbury & Guillot de Suduiraut, 2003) 3 comb. nov., ?S. eximia (A. Adams, 1853) 3 , ?S. geigeri Poppe, Tagaro & Salisbury, 2009 3 , ?S. gilbertsoni (Cate, 1968) 3 comb. nov., ?S. guttata (Swainson, 1824) 3 comb. nov., ?S. hayashii (Kira, 1959) 3 , ?S. ivanmarrowi Marrow, 2016 3 , ?S. lavoisieri Guillot de Suduiraut, 2002 3 , ? S. lorenzi (Poppe & Tagaro, 2006) 3 comb. nov., ?S. mariae (A. Adams, 1853) 3 , ?S. martini Poppe & Tagaro, 2006 3 , ?S. petiti Poppe & Tagaro, 2006 3 , ?S. parkinsoni Salisbury & Wolff, 2005 3 , ?S. potensis (Montrouzier, 1858) 3 , ?S. sowerbyi d'Orbigny, 1852 3 , ?S. splendidula Salisbury & Guillot de Suduiraut, 2003 3 , ?S. vicdani Cernohorsky, 1981 3 , ?S. yaekoae (Habe & Kosuge, 1966) 3 . The genus Scabricola as commonly understood is extremely heterogeneous. It combines such dissimilar forms as Scabricola yaekoae, which in shell outline is close to Mitramitra, and S. potensis (Montrouzier, 1858), with its ovate shell resembling species of Pterygia.

  Swainsonia H. Adams & A. Adams, 1853 (Figs 28 B, C, 29 G -I) Type species: Mitra fissurata Lamarck, 1811; by typification of replaced name [Mitrella Swainson, 1831 non Risso, 1826].Synonym: Mitrella Swainson, 1831 (not Risso, 1826); Type species: Mitra fissurata Lamarck, 1811; SD,Cossmann (1899: 152).Diagnosis: Shell small to medium-sized (15-65 mm), fusiform or elongate biconical, weakly sculptured and often polished in appearance. Protoconch narrowly-conical, of about three smooth, weakly convex, whorls. Spire ortho-or cyrtoconoid, spire whorls evenly convex to flattened; suture distinct, impressed. Sculpture of regular, shallow, and widely spaced punctate spiral grooves, either covering the entire shell surface, or limited to the adapical whorl portion, . Axial sculpture usually not developed or, when present, consisting of numerous, fine, and low, rounded folds, developed on the spire but indistinct on the periphery of the last adult whorl. Shell base typically with convex outline, continuing into short and stout, deeply notched, siphonal canal. Siphonal fasciole indistinct or moderately developed. Aperture elongate, narrow to moderately wide. Outer aperture lip straight or gently evenly convex adapically; anterior edge of outer lip strongly convex, sometimes slightly drawn abapically.Inner lip with 4-5 fine or strong, subequal columellar folds. Shell light brown, orange, or white, typically with intricate scale-like pattern of fine light or white lines. Scabricola-type radula with comb-like rachidian bearing six long, slender cusps, the four central ones notably exceeding the two marginal ones (Figs28 B, C). Lateral teeth with very strong, robust cusp, giving tooth the appearance of a tin-opener, the serrate inner surface of tooth bearing 3-4 weaker cusps. Base of laterals with curved upper margin, forming a socket interlocking laterals of adjacent rows.Distribution: Indo-Pacific, upper subtidal depths, on sand. Species included: Swainsonia biconica Bozzetti, 2011 3 , S. casta (Gmelin, 1791) 1 , S. fissurata (Lamarck, 1811) 2 , S. fusca (Swainson, 1824) 1 , S. newcombii (Pease, 1869) 3 , S. ocellata (Swainson, 1831) 3 . Remarks. Many contemporary authors have treated Swainsonia as a subgenus of Scabricola (Cernohorsky 1970; 1991; Pechar, Prior & Parkinson 1980; but see Poppe & Tagaro 2008), but our results indicate that it should be elevated to full genus. This treatment of Scabricola and Swainsonia, however, contradicts the phylogenetic hypothesis implied by radular morphology. The typical Scabricola radula (as described by Cernohorsky 1970) is present in Scabricola variegata, Swainsonia fusca and Swainsonia casta, while the radulae of Scabricola desetangsii and S. olivaeformis show a divergent morphology. Genus Neocancilla Cernohorsky, 1966 (Figs 28 G -I, 30) Type species: Voluta papilio Link, 1807; OD. Diagnosis: Shell small to medium-sized (10-65 mm), fusiform, with short and stout siphonal canal. Protoconch pointed, narrowly conical, of about 3 slightly convex glossy whorls. Suture deeply impressed to canaliculate. Spire orthoconoid, teleoconch whorls evenly convex. Sculpture of wide spiral cords, low or slightly elevated adapically, or bearing rows of fine beads. Spiral cords separated by narrow deep groves and intersected by axial riblets of similar strength, sculpture. Siphonal canal short and robust, gradually merging to shell base, although separated from it by narrow depression at about the level of the second columellar fold. Siphonal notch deep. Aperture elongate; outer aperture lip evenly convex along most of its length. Inner aperture lip with four strong columellar folds, adapicalmost strongest. Shell cream, pale or orange, usually with contrasting darker bands. Radula with narrow rachidian bearing six blunt, very robust cusps; paired central cusps greatly exceeding the others (Figs 28 G -I). Lateral teeth with four or five very short and blunt cusps situated proximally, distal portions of teeth without cusp. Distribution: Indo-Pacific, intertidal and upper subtidal depths, mostly on sand. Species included: Neocancilla antoniae (H. Adams, 1870) 3 , N. arenacea (Dunker, 1852) 3 , N. clathrus (Gmelin, 1791) 1 , N. hartorum Poppe, Salisbury, Tagaro, 2015 3 , N. hemmenae (Salisbury & Heinicke, 1993) 3 , N. kayae Cernohorsky, 1978 3 , N. madagaskarensis Herrmann, 2017 3 , N. papilio (Link, 1807) 1 , N. rufescens (A. Adams, 1853) 1 , N. takiisaoi (Kuroda, 1959) 2 , N. waikikiensis (Pilsbry, 1921) 2 .

Genus

  PterygiaRöding, 1798 (Figs 31, 32 A -C) Type species: Voluta dactylus Linnaeus, 1767; SD,Dall (1915: 52).Synonym: Cylindromitra P. Fischer, 1884. Type species: Cylindra coronata Schumacher, 1817[objective synonym of Voluta crenulata Gmelin, 1791]; by typification of replaced name[Cylindra Schumacher, 1817, non Illiger, 1802; itself by M].Diagnosis: Shell small to medium-sized (20-70 mm), ovate or ovate-elongate, with very low spire and tall aperture. Protoconch cyrtoconoid, with about 3 smooth evenly convex whorls.Spire height attaining only about 10-15% of total shell height, with only 3-4 whorls exposed, and a proportionally very high last adult whorl. Suture adpressed. Last adult whorl convex or strongly convex, smooth or with dominating spiral or cancellate or crenulate sculpture. Spiral elements represented by fine grooves or cords that can be fine or wide, undulating, rounded or resembling scales tiling in posterior direction. Siphonal notch distinct. Aperture elongate, usually narrow. Outer aperture lip convex, smooth or denticulate, inner lip calloused, bearing 6-8 subequal columellar folds. Shell off-white, whitish or brownish, or with a pattern of red or orange bands. Radula very narrow, consisting of rachidian only, bow shaped, bearing multiple cusps, or with a single blunt medial cusp and lateral serration (Figs32 A -C).

Genus

  Nebularia Swainson, 1840 33) Type species: Mitra contractaSwainson, 1820; SD, Herrmannsen (1847: 110). Synonyms: Dibaphus Philippi, 1847. Type species: Mitra edentula Swainson, 1823; M. Mutyca H. Adams & A. Adams, 1853. Type species: Mitra ancillides Broderip, 1836; SD, Wenz (1943: 1296). Mitroidea Pease, 1865. Type species: Mitroidea multiplicata Pease, 1865; M. ?Mauritia H. Adams, 1869. Type species: Mauritia barclayi H. Adams, 1869; M. Diagnosis: Shell small to large (15-150 mm), fusiform to elongate fusiform. Protoconch pointed, narrowly-conical, of 3+ slightly convex, smooth and glossy whorls. Teleoconch whorls usually flattened in outline, with orthoconoid or acuminate spire. Sculpture variable, from indistinct to strong and deep groves separating flattened or rounded, sometimes undulating, cords. Siphonal canal robust, short to moderately long, sometimes with distinct fasciole, giving siphonal canal a recurved appearance. Aperture elongate, its outer lip often thickened in its adapical portion. Margin of outer lip smooth or bearing fine denticles throughout its length. Inner lip with 4-5 delicate folds of subequal strength. Shell pale, usually with a pattern of brown bands and blotches. Radula triserial, with narrow rachidian and laterals of varying width. Rachidian with five cusps of which a central unpaired cusp greatly exceeds the others in strength (Figs 32 D -F). Laterals bearing 6-12 cusps, 2nd-4th from rachidian cusp being strongest, and succeeding ones gradually diminishing. Distribution: Indo-Pacific, intertidal and upper subtidal depths, rocky shores and reef crevices.Species included: Nebularia acuminata (Swainson, 1824) 1 comb. nov., N. aegra (Reeve, 1845) 3 comb. nov., N. ancillides (Broderip, 1836) 3 comb. nov., N. baerorum(Poppe & Tagaro, 2010) 3 , N. bellula (A. Adams, 1853) 3 comb. nov., N. chrysostoma (Broderip, 1836)3 , N. coarctata(Reeve, 1844) 3 , N. contracta (Swainson, 1820) 1 , N. deynzeri(Cernohorsky, 1980) 3 comb. nov., N. dondani (Cernohorsky, 1985) 3 comb. nov., N. edentula (Swainson, 1823) 3 comb. nov., N. eremitarum (Röding, 1798) 1 comb. nov., N. fastigium (Reeve, 1845) 3 comb. nov., N. ferruginea (Lamarck, 1811) 2 , N. gourgueti(Poppe, Salisbury, Tagaro, 2015) comb. nov., N. guidopoppei (Thach, 2016) 3 comb. nov., N. incompta (Lightfoot, 1786) 1 comb. nov., N. inquinata(Reeve, 1844) 3 , N. multiplicata (Pease, 1865)3 comb. nov., N. nebulosa (Broderip, 1836) 3 comb. nov., N. nivea (Broderip, 1836) 3 comb. nov., N. pellisserpentis (Reeve, 1844) 3 comb. nov., N.

Genus

  Isara H. Adams & A. Adams, 1853 (Figs 34, 35 A -C) Type species: Mitra bulimoides Reeve, 1845 [= M. glabra Swainson, 1821, after Cernohorsky (1976)]; SD, Cossmann (1899: 153). Synonym: Fuscomitra Pallary, 1900. Type species: Mitra fusca Pallary, 1900 [= Mitra cornea Lamarck, 1811]; SD

  Indo-Pacific, West Africa and Mediterranean, Caribbean, from upper subtidal to upper bathyal depths.

Genus

  Subcancilla Olsson & Harbison, 1953 (Figs 35 D -F, 36) Type species: Mitra sulcata Swainson, 1825; OD. Diagnosis: Shell small to large (15-125 mm), fusiform, with white or light background colour and contrasting brown lines on crests of spiral cords. Protoconch smooth, with slightly convex or flattened whorls, either pointed, narrowly-conical, of about 3 whorls, or bulbous, of about 2 whorls. Suture indistinct. Spire moderately high, spire whorls evenly convex, sometimes shouldered in appearance due to strong elevated spiral cords that are rather sharp and widely set. Interspaces between spiral cords sculptured with fine, dense riblets, or smooth with microsculpture of very fine growth lines. Shell base triangular with straight, tapering siphonal canal not delineated from shell base. Siphonal notch deep. Aperture high, about half of shell height, narrow, parallel-sided. Outer aperture lip very gently convex, smooth. Inner lip with three fine, oblique columellar folds. Thin light-brown periostracum sometimes present. Radula with narrow rachidian and laterals 2-4 times wider than rachidian (Figs 35 D -F), both multicuspidate, either with dense pointed cusps of subequal strength or with one cusp on laterals notably stronger than others and rachidian bearing symmetrical paired cusps also differentiated in length and strength. Distribution: Tropical East Pacific, Caribbean, subtidal and bathyal depths.Species included: Subcancilla attenuata (Broderip, 1836) 1 , S. belcheri (Hinds, 1843) 3 comb. nov., S. calodinota (S. S. Berry, 1960) 3 , S. candida(Reeve, 1845) 3 , S. directa (Berry, 1960) 3 , S. edithrexae (Sphon, 1976) 3 , S. erythrogramma (Tomlin, 1931) 1 , S. funiculata(Reeve, 1844) 3 , S. gigantea(Reeve, 1844) 3 , S. haneti (Petit, 1852) 3 , S. hindsii(Reeve, 1844) 3 , S. joapyra Simone & Cunha, 2012 3 , S. leonardi (Petuch, 1990) 1 comb. nov., S. leonardhilli Petuch, 1987 2 , Subcancilla lindae Petuch, 1987 3 , S. lopesi Matthews & Coelho, 1969 3 , S. malleti (Petit de la Saussaye, 1852) 3 , S. phorminx (S. S. Berry, 1969) 3 , S. sulcata (Swainson, 1825) 2 .

  Carinomitra Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Figs 37, 38 A -C) Mitra peculiaris Reeve, 1845; OD, herein.Diagnosis: Shell small(7-20 mm), elongate-fusiform, with characteristic pattern formed by light adapical portions of teleoconch whorls, gradually transiting to yellow or orange peripheries.Protoconch narrowly conical or cyrtoconoid, of 3 to almost 5, evenly convex, smooth and glossy whorls. Suture distinct, slightly impressed. Early spire whorls with flattened or slightly convex outlines, later gently convex or distinctly carinated at the suture. Except for subsutural carina, sculpture limited to very weak spiral grooves on shell base and siphonal canal, thus the shell appears smooth. Siphonal canal tapering, siphonal notch shallow or indistinct. Aperture elongate, rather narrow; outer lip smooth, convex. Inner lip with 4-5 fine oblique columellar folds.Rachidian narrow, bearing either six cusps of comparable size, or four cusps with two medial notably stronger than two laterals. Laterals wide, bearing multiple cusps, of which 2nd and 3rd from rachidian are strongest.Distribution: Indo-Pacific, upper subtidal depths, reef crevices and coral rubble. Species included: Carinomitra peculiaris (Reeve, 1845) 1 comb. nov., C. saltata (Peace, 1865) 2 comb. nov., C. typha (Reeve, 1845) 1 comb. nov. Etymology: The name Carinomitra refers to the presence of the distinct keel (Latin carina) in at least two species of this newly recognized lineage, including the type species C. peculiaris. Remarks. The genus Carinomitra is conchologically distinctive due to the carinated or adapically inflated adult whorls and the characteristic colour pattern. The radulae of the three studied species exhibit notable disparity. In M. peculiaris, the rachidian bears multiple cusps with a central unpaired cusp; conversely, in C. saltata and C. typha the unpaired central cusp of the rachidian is absent, and the paired central cusps greatly exceed the paired lateral cusps. This variation in radula morphology parallels that observed in Imbricaria and Cancilla (see discussion below). Genus Condylomitra Fedosov, Herrmann, Kantor & Bouchet, n. gen. (Figs 38 D, E, 39) Type species: Mitra tuberosa Reeve, 1845; OD, herein. Diagnosis: Shell small (10-25 mm) , ovate to almost conical, heavily sculptured, with pointed conical apex. Protoconch pointed, narrowly-conical, of about 3 very slightly convex, smooth whorls. Spire whorls convex; last adult whorl proportionally very high, sculptured with densely spiral cords, 1 to 3 adapical cords very strong, forming prominent, spirally elongated rounded tubercles, directed slightly upwards. Siphonal canal very short, with well developed fasciole, siphonal notch distinct. Aperture elongate, rather narrow, widening anteriorly. Outer apertural lip gently convex or straight, inner lip with four closely set columellar folds. Radula very narrow, monoserial, with rachidian bearing strong, unpaired medial cusp, flanked with 2-8 smaller pointed cusps on each side. Proboscis dorso-ventrally flattened. Distribution: Indo-Pacific, intertidal and upper subtidal depths, mostly on hard substrates. Species included: Condylomitra bernhardina (Roding, 1798) 1 comb. nov., C. tuberosa (Reeve, 1845) 1 comb. nov.

  Species included: Magnamitra sandrogorii Huang & Salisbury, 2017 2 . Distribution: Indo-Pacific, bathyal depths. Remarks. See remarks under Calcimitra. Atrimitra Dall, 1918 (Figs 40 A -C) Mitra idae Melvill, 1893; OD.Diagnosis: As no material on Atrimitra was available for the present study, we do not provide a diagnosis to this poorly studied genus.Species included: Atrimitra caliginosa (Reeve, 1844)3 comb. nov., A. catalinae (Dall, 1920) 2 comb. nov., A. effusa (Broderip, 1836) 3 , A. idae (Melvill, 1893) 3 , A. orientalis (Griffith & Pidgeon, 1834) 3 comb. nov., A. semigranosa (Martens, 1897) 3 comb. nov. Distribution: Tropical East Pacific, subtidal depths. Remarks: see remarks under Isara. Vicimitra Iredale, 1929 (Fig. 40 D) Type species: Vicimitra prosphora Iredale, 1929; M.Diagnosis: As no material on Vicimitra prosphora was available for the present study, we do not provide a diagnosis to this genus.Species included: Vicimitra prosphora Iredale, 19293 (Fig.40 D).Distribution: New South Wales, Australia.Remarks. See remarks under Quasimitra.Genus ProbataSarasúa, 1989 (Figs 38 E, 40 E, F) Type species: Mitra espinosai Sarasúa, 1978; OD.Diagnosis: Shell small to medium-sized (15-45 mm), broadly fusiform, light brown, sometimes with occasional white flakes. Protoconch white, pointed, of about 2.5 smooth glossy whorls.Suture distinct, impressed. Teleoconch whorls slightly convex, sculptured with fine, low, evenly interspaced spiral cords. Siphonal canal stout, tapering, siphonal notch shallow. Aperture elongate, outer lip convex, thickened adapically. Inner apertural lip with five fine columellar folds, weakening gradually towards siphonal canal. Radula triserial, rachidian narrow, with eight pointed cusps, two medial longest, those situated laterally becoming gradually weaker. Laterals with 15+ cusps, strong and long compared to length of tooth base.

  shell proportions and number of columellar folds. The overall plesiomorphic radula of Probata barbadensis (Fig.38 F) resembles the radula of many other lineages of Mitridae and gives no clue as to where the genus should be assigned.DibaphimitraCernohorsky, 1970 (Fig. 40 G) Type species: Mitra florida Gould, 1856; OD.

  Species included: Dibaphimitra florida (Gould, 1856) 2 .Distribution: Caribbean, subtidal to upper bathyal depths.Remarks: The radula of D. florida pictured by[START_REF] Cernohorsky | Systematics of the families Mitridae and Volutomitridae[END_REF] is overall a typical mitrid radula, with a narrow rachidian and very wide nulticuspidate laterals. Although this morphology unequivocally places Dibaphimitra in Mitridae, it is inconclusive regarding the subfamilial placement of D. florida. More studies are needed to characterize the affinities of this enigmatic Caribbean lineage.

  IM-2013-40651, Papua New Guinea, North of Rabaul, BIOPAPUA Stn CP3671, 04°04'S, 151°56'E,[585][586][587][588][589][590][591][592][593][594][595][596][597][598][599][600][601] lv, 24.15 mm (Figs 17G; 18 A, B).Paratype.MNHN IM-2013-58429, KAVIENG 2014 Stn CP4449, 02°10'S, 150°11'E, 623-908 m, lv, 35.5 mm. Other material examined: Papua New Guinea, BIOPAPUA Stn 3736, off Lancasay, 08°14'S 150°32'E, 760-769 m, 1 lv (21.5 mm); PAPUA NUIGINI Stn CP3979, North of Bagabag Is, 04°44'S 146°11'E, 540-580 m, 1lv (MNHN IM-2013-19698, 25.75 mm); Stn CP3994, North from Cape King William, 06°00'S 147°35'E, 648-652 m, 1 lv (MNHN IM-2013-19833, 33.6 mm); MADEEP Stn DW4321 08°39'S 151°47'E, 612 m, 1 lv (MNHN IM-2013-45873).? NEW CALEDONIA, BATHUS 3 Stn DW790, 23°49'S 169°48'E, 685-715 m, 17.9 mm; Norfolk Ridge, BATHUS 4 Stn DW920 18°45'S 163°17'E,[610][611][612][613][614][615][616][617][618][619][620] 15.1 mm (Fig. 18 C).

  pale-yellowish; microsculpture of very fine collabral growth lines.Radula (of holotype) 0.7-0.75 mm long, 0.14 mm wide, consisting of about 85 rows.Rachidian about 40 µm and laterals 45 µm wide, all three similar, with evenly convex anterior margin, bearing 12-13 moderately long, pointed subequal cusps, those at teeth margins smaller than others. : BIOPAPUA Stn DW3735, 08°15'S, 150°34'E, 450-500 m, 21.4 mm; Stn data lost, 28.5 mm; Stn DW3771, 05°34'S, 151°33'E, 295-422 m, 9.2 mm.DescriptionShell medium-sized (holotype 25.2 mm), fusiform, heavily sculptured, with rather high spire and elongated aperture. Protoconch pointed, cyrtoconoid, of 3+ glossy, convex whorls (usually strongly corroded or missing). Teleoconch of 6+ slightly convex whorls, suture distinctly canaliculated. Subsutural shelf well developed, rather narrow, slightly inclined towards axis, bordered by projecting margin of adapical spiral cord. Early teleoconch whorls corroded, third and subsequent teleoconch whorls with five very strong, elevated, gemmate spiral cords separated by deep interspaces. Spiral cords subequal, adapical and abapical cords slightly weaker than intermediate ones, microsculpture of fine growth lines in interspaces between spiral cords; abapical cord of previous whorl partly superposed by adapical cord of succeeding whorl.Gemmules on spiral cords axially aligned, connected by low and weak axial riblets in interspaces between spiral cords. Spiral cords become distinctly beaded on penultimate and last adult whorls, forming rows of closely set, rounded, glossy granules. Five spiral cords on adapical portion of last adult whorl succeeded by six cords on shell base and five cords on siphonal canal, of which last three oblique, weaker than preceding, irregularly gemmate. Siphonal canal delineated from shell base by slightly widened interspace with low wavy spiral thread. Siphonal canal rather long and stout, slightly recurved.Aperture elongate, its outer lip gently convex throughout its height, wavy, reflecting arrangement of spiral cords. Inside of outer lip with 8 strong lirae. Inner lip strongly calloused, reflected, retaining weak reticulate sculpture adapically, with three strong oblique columellar folds at mid-height.Shell uniformly white or pale.Radula of holotype 0.76-0.8 mm long, 0.19-0.20 mm wide, composed of about 65 transverse rows of uniform, delicate, multicuspidate teeth. Rachidian about 60 µm wide, with 17 cusps, of which two marginals very weak, and the others subequal, long and pointed. Laterals about 50 µm wide, with slightly convex anterior margin, bearing 14-15 pointed, rather long cusps.Distribution: Papua New Guinea, North of main island, 269-450 m.

  The species is named after another giant in malacology, Winston Ponder of the AMS, whose early work, including his studies on mitriform families, set a milestone in our understanding of neogastropod evolution.

  Fedosov, Herrmann, Kantor & Bouchet, n. sp.

  m, 1 lv (Paratype 2, 26.9 mm); Stn 3810, East coast off Toupeti, 21°44'S, 166°38'E, 384-385 m, 2 lv (Paratype 1, 27.35 mm; Paratype 4, 24.0 mm); Stn 3814, Passe du Solitaire, 21°49'S, 166°44'E, 331-344 m, 1 lv (Paratype 5, 20.4 mm).

  Aperture narrow, elongated; outer lip wavy, convex in adapical portion and straight below mid-height. Inside of outer lip smooth. Inner lip strongly calloused, sometimes reflected, retaining weak reticulate sculpture adapically, with four closely set fine columellar folds at mid height. Coloration uniformly cream, protoconch white. Radula (of paratype 5) 0.62 mm long, 0.13 mm wide, of about 75 rows. Rachidian narrow, about 20 µm wide, bearing 8 strong, moderately long, pointed cusps. Laterals attaining 47 µm in width, with straight anterior margin bearing 16+ strong, pointed, rather widely set cusps.

  New Caledonia, Papua New Guinea, Etymology: The species epithet refers to the type locality.Remarks: While the specimens from New Caledonia show a fine reticulate sculpture pattern on the late teleoconch whorls, the specimen from Papua New Guinea is characterized by denser axial ribs and in overall shell morphology is somewhat intermediate between such typical G. neocaledonica n. sp. and Fusidomiporta ponderi n. sp. The latter species is the one that resembles most G. neocaledonica n. sp., but it can be differentiated by its notably stronger spiral cords and rather fusiform shell. The combination of elongate-biconical shell with subcylindrical whorl profile and characteristic reticulate pattern on late whorls allows for easy recognition of G. neocaledonica n. sp. Other species of Gemmulimitra n. gen. differ either in shell shape or in sculpture pattern, and G. avenacea, despite being the closest to G. neocaledonica n. sp. in our multi-gene analyses, shows no morphological resemblance to it whatsoever.

  morphological studies for the taxonomy of the Mitridae

5 -

 5 Neocancilla-type: radula characterized by a rachidian with a pair of very robust and blunt central cusps, and laterals bearing few short and robust cusps in their medial portions. This type of radula is known in all species of Neocancilla. The radulae of Scabricola olivaeformis and S. coriacea have morphologies somewhat intermediate between the Scabricola-type and the Neocancilla-type. 6 -Pterygia-type: very narrow uniserial radulae remarkable by the complete loss of laterals. Rachidians of varying morphology, but usually flattened, with serrated margins. Radulae of this type are found in all species of the genera Pterygia and Condylomitra, and in the Strigatella lugubris -Strigatella coronata clade of Strigatella.

  , as there are no data on the possible adaptive value of different morphologies and selection pressures that have led to the emergence of the observed diversity. Further studies on the functional morphology and biochemistry of mitrid secretions would in this respect open new perspectives.

(

  Domiporta, Dibaphimitra, Neocancilla and Sohlia). In the present study we have re-assessed the genus-level systematics of Recent Mitridae based on a combination of molecular and morphological data. The genus Charitodoron is segregated in the newly established family Charitodoronidae. The 26 genera of the revised Mitridae comprise six subfamilies: Mitrinae (with 14 genera), Strigatellinae (with the single genus Strigatella), Imbricariinae (with six genera), Cylindromitrinae (with the genera Pterygia and Nebularia), Isarinae new subfamily genera Isara and Subcancilla) and Pleioptygmatinae (with the single genus Pleioptygma); six genera, Atrimitra Dall, 1918, Carinomitra gen. nov., Condylomitra gen. nov., Dibaphimitra Cernohorsky, 1970, Magnamitra Huang & Salisbury, 2017 and Probata Sarasúa, 1989 are treated as incertae sedis. Undoubtedly the most revolutionary change in the taxonomy of the Mitridae is the falling apart of Mitra with the genus in its former taxonomic extension now reassigned to 14 genera, of which 6 are new. Other noteworthy changes are the transfer of the formerly mitrine Nebularia to Cylindromitrinae; of Strigatella to its own, newly recognized, subfamily Strigatellinae; of the formerly imbricariine genera Ziba and Domiporta to Mitrinae; and of Subcancilla to the newly established subfamily Isarinae. The genera Mitra and Ziba now include a much reduced species diversity, as all the Indo-Pacific species earlier placed in Ziba are now transferred to Imbricaria. The contents of Imbricaria is also expanded as a result of the transfer of the Indo-Pacific species of Subcancilla, which appear unrelated to the New World species of that genus. The 32 genera now recognized in the family Mitridae increase considerably the previously accepted genus-level diversity (21 genera, including Charitodoron, indexed in WORMS as of May 2017). Still, the placement of several sequenced species remains uncertain. Lineage 3 represented by MNHN IM-2007-35623 and lineage 16 represented by MNHN IM-2007-30270

  Indo-Pacific, of which 19 do not occur outside the Indo-Pacific according to our data. Moreover, the subfamilies Imbricariinae and Cylindromitrinae, as circumscribed herein, are represented solely by Indo-Pacific forms. At least three genera (Episcomitra, Isara and Ziba) are found in West Africa, with the first two also inhabiting the Mediterranean. The New World fauna of Mitridae includes at least eight genera in four subfamilies: Mitrinae (Neotiara), Strigatellinae (Strigatella), Isarinae (Isara, Subcancilla), and Pleioptygmatinae (Pleioptygma), plus the genera Atrimitra, Dibaphimitra and Probata that we treat as incertae sedis. The six genera Atrimitra, , Pleioptygma, Probata, and Subcancilla are currently thought to be endemic to Panamic and/or the Caribbean, although a close affinity of Panamic and Caribbean species was herein confirmed only in the genera Neotiara and Subcancilla.

  taken place in i) Late Tethys -Paratethys -Indo-Pacific, ii) West Africa and iii) the New World.

  Etude zoologique et anatomique de la Mitra zonata, Marryat. Journal de Conchyliologie, 49: 77-95. West TL. 1990. Feeding behavior and functional morphology of the proboscis of Mitra idae (Mollusca: Gastropoda: Mitridae). Bulletin of Marine Science 46(3): 761-779. West TL. 1991. Functional morphology of the proboscis of Mitra catalinae Dall, 1920 (Mollusca: Gastropoda: Mitridae) and the evolution of the mitrid proboscis. Bulletin of Marine Science 48: 702-718. WoRMS. 2017. World Register of Marine Species, http://www.marinespecies.org/ on 2017-05-31.

Figure 1 .

 1 Figure 1. Bayesian phylogenetic tree of Mitroidea based on the analysis of cytochrome oxydase subunit I (COI) gene. Branch support as Maximum likelihood bootstrap values (when >50) / Bayesian PP (when >0.7).

Figure 2 .

 2 Figure 2. Examples of conflicting species hypotheses; with morphospecies delineated based on the shell morphology not consistent with the phylogenetic groupings inferred from the analysis of COI. A. Strigatella aff. paupercula (in COI clade 11); B, C. Strigatella paupercula (COI clade 7); D, E. Mitridae gen. sp. (in COI clade 15); F. Mitridae gen. sp. (in COI clade 18); G -L. "Mitra" glaphyria (in COI clade 13); M -R. Cancilla fibula / rehderi (in COI clade 24-7).

Figure 3 .

 3 Figure 3. Bayesian phylogenetic tree of Mitridae, Charitodoronidae new family and Pyramimitridae based on the analysis of 3-mit dataset. Branch support as Maximum likelihood bootstrap values (when >70) / Bayesian PP (when >0.75). Type species of nominal genera marked with asterisks. Inferred phylogenetic lineages that are given a taxonomic status labelled correspondingly.

Figure 4 .

 4 Figure 4. Bayesian phylogenetic tree of Mitridae, Charitodoronidae new family and Pyramimitridae based on the analysis of 4G dataset. Support values and labelling as in Figure 3.

Figure 5 .

 5 Figure 5. Bayesian phylogenetic tree of Mitridae, Charitodoronidae new family and Pyramimitridae based on the analysis of 4G-red dataset. Support values and labelling as inFigures3-4.

Figure 6 .

 6 Figure 6. Histological sections of the foregut of Charitodoron veneris. A. Section through the tip of the proboscis and salivary gland, general view; B. Close up of the salivary gland. C. Close up of the proboscis tip; D. Section through the buccal mass, mid-and posterior oesophagus, general view. E. Close up of the odontophore, radula and salivary ducts; F. Same area as previous, few sections further -opening of salivary ducts; G. Close up of posterior oesophagus and valve-like structure.

Figure 7 .

 7 Figure 7. Transversal histological sections of the proboscis of Condylomitra tuberosa. A. Section through the mouth opening; B. Section through the buccal cavity; C. Close up of the buccal mass and epiproboscis tip; D. Section through the buccal mass at the level of the odontophore; E. Close up of the buccal mass and epiproboscis complex; F. Section through the

Figure

  Figure 8. Charitodoron species. A. Charitodoron sp.,MNHN IM-2007-38306, off

Figure 41 .

 41 Figure 41. Distribution of different radula types throughout the Mitridae tree. Dark grey -Mitra type; yellow -Strigatella type; brown -Nebularia type; red -Profundimitra type; dark-green -Imbricaria type; light-green -Cancilla type; light-blue -Scabricola type; dark blue -Neocancilla type; purple -Pterygia type; light-grey -no data. The topology corresponds to the 4G tree on Figure 4.

Figure 42 .

 42 Figure 42. Taxonomic composition (subfamilies) and geographical coverage of our molecular sampling. Diameter of the circles proportional to the number of species in each principal location, which is indicated for each circle with more than one species sampled.

  ; Cylindromitrinae -blue; Imbricariinae -green; Isarinae -purple; Mitrinae -orange; Strigatellinae -yellow; Incertae sedis -grey.

Figure 43 .

 43 Figure 43. Bathymetric distribution of the genera of Mitridae. Black bars -based on our sampling; grey bars -based on published data[START_REF] Lozouet | Eumitra récentes de la région néo-calédonienne et Charitodoron fossiles de l'Oligocène supérieur d'Aquitaine (Mitridae)[END_REF][START_REF] Cernohorsky | The Mitridae of the World. Part 1. The Subfamily Mitrinae[END_REF] 1991;[START_REF] Huang | Calcimitra, a new genus of deep-water Mitridae (Gastropoda: Mitridae) with the description of five new species from Taiwan and the Philippines[END_REF]

Figure 1 .Figure 1 .Figure 2 .Figure 3 .Figure 4 .Figure 5 .Figure 6 .Figure 7 .Figure 8 .Figure 9 .Figure 10 .Figure 11 .Figure 12 .Figure 13 .Figure 14 .Figure 15 .Figure 16 .Figure 17 .Figure 18 .Figure 19 .Figure 20 .Figure 21 .Figure 22 .

 112345678910111213141516171819202122 Figure 1. Bayesian phylogenetic tree of Mitroidea based on the analysis of cytochrome oxydase subunit I (COI) gene. Branch support as Maximum likelihood bootstrap values (when >50) / Bayesian PP (when >0.7).229x314mm (300 x 300 DPI)

Figure 23 .Figure 24 .Figure 25 .Figure 26 .Figure 27 .Figure 28 .Figure 29 .Figure 30 .Figure 31 .Figure 32 .Figure 33 .Figure 34 .Figure 35 .Figure 36 .Figure 37 .Figure 38 .Figure 39 .Figure 40 .

 232425262728293031323334353637383940 Figure 23. Radulae in species of Strigatella. A. Strigatella paupercula, MNHN IM-2013-15188, PAPUA NIUGINI Stn PM39, 05°12.1'S, 145°48.4'E, intertidal, 22.2 mm; B. S. luctuosa, MNHN IM-2013-11682, PAPUA NIUGINI Stn PR14, 05°12'S, 145°48.1'E, 2-3 m, 20.9 mm; C. S. decurtata, MNHN IM-2013-15843, PAPUA NIUGINI Stn PM41, 05°08.1'S, 145°49.3'E, 0-1 m, 22.1 mm; D. S. vexillum, MNHN IM-2013-12570, PAPUA NIUGINI Stn PR24, 05°12.3'S 145°48.8'E, 24.0 mm; E. S. coronata, MNHN, Society Is, off Moorea, 9.0 mm; F. S. lugubris, MNHN IM-2013-40678, Austral Is, TUHAA PAE Stn AT07, 23°19.7'S 149°29.3'W, 14 m, 13.7 mm; G. S. tristis, BAU 2534-2, Panama, Rio Mar, 8°27.5'N, 79°58.0'W, intertidal, 20.3 mm. 167x86mm (300 x 300 DPI)

Figure 41 .

 41 Figure 40. Genera Atrimitra, Vicimitra, Probata, and Dibaphimitra. A. Atrimitra idae, holotype, NMW.1955.158.00066, Point Loma, Lower California, Mexico, 60.0 mm; B. Atrimitra effusa, lectotype, NHMUK 1966415, Guacomayo, Central America, 31.6 mm; C. Atrimitra caliginosa, holotype, NHMUK 1966718, locality unknown, 34.9 mm; D. Probata espinosai, paratype, MNHN-IM-2012-37719, in front of Marianao, Havana, Cuba, 10 mm; E. Probata barbadensis, MNHN IM-2013-7772, Guadeloupe, KARUBENTHOS Stn GR36, 16°30.6'N, 61°28.4'W, 12 m, 20.6 mm; F. Dibaphimitra florida, holotype of Mitra fergusoni G.B. Sowerby II, 1874, NHMUK 1879.2.26.133, Panama, 42.7 mm. 167x149mm (300 x 300 DPI)

Figure 42 .Figure 43 .

 4243 Figure 42. Taxonomic composition (subfamilies) and geographical coverage of our molecular sampling. Diameter of the circles proportional to the number of species in each principal location, which is indicated for each circle with more than one species sampled. Charitodoronidae -brown; Cylindromitrinae -blue; Imbricariinae -green; Isarinae -purple; Mitrinae -orange; Strigatellinae -yellow; Incertae sedis -grey. 81x39mm (300 x 300 DPI)

Subfamily Isarinae subfam. nov.

  ., Cancillopsis n. gen., Neotiara n. gen., Profundimitra n. gen., Pseudonebularia n. gen., Quasimitra n. gen.,

	F o
	r
	R e v i e w
	O n
	Roseomitra n. gen.
	Subfamily Strigatellinae Troschel, 1869
	Genus Strigatella Swainson, 1840.
	Subfamily Imbricariinae Troschel, 1867
	Genera Imbricaria Schumacher, 1817, Cancilla Swainson, 1840, Scabricola Swainson, 1840,
	Neocancilla Cernohorsky, 1966, Swainsonia H. Adams & A. Adams, 1853, Imbricariopsis
	n. gen.
	Subfamily Cylindromitrinae Cossmann, 1899
	Genera Pterygia Röding, 1798, Nebularia Swainson, 1840.
	Genera Isara H. Adams & A. Adams, 1853, Subcancilla Olsson & Harbison, 1953.

l y 22 Subfamily Pleioptygmatinae Quinn, 1989 Genus Pleioptygma Conrad, 1863

y 23 Taxonomical part Superfamily Mitroidea Swainson, 1831 Family Charitodoronidae Fedosov, Herrmann, Kantor & Bouchet, new family

		(Fig. 8)	., Condylomitra
	n. gen. Diagnosis: Shell small (13-35 mm), fusiform, white or pale, sometimes with dark bands, or
	covered with brown periostracum. Protoconch white, bulbous, of about 1.5 smooth glossy whorls
	and with large initial nucleus. Suture distinct, impressed. Spire tall, orthoconoid; spire whorls
	evenly convex, smooth or sculptured with weak spiral grooves or cords. Sometimes distinct
	cancellate sculpture developed. Siphonal canal moderately long, robust or tapering, sometimes
	slightly recurved at its tip; siphonal notch shallow. Aperture wide, elongate; outer aperture lip F smooth, evenly convex. Inner lip with no columellar folds. o Head-foot pale, with sometimes nebulose dark bands. Tentacles moderately long with eyes r situated laterally at about mid-length. Siphon long. Osphradium as long as gill, attaining about
	2/3 of the length of the latter. Proboscis well developed, without distinct proboscis rim; R epiproboscis absent. Radula with narrow rachidian, bearing five subequal cusps, and wide, e multicuspidate laterals, with cusps being more dense and stout proximally and becoming more v slender towards radula margins; lateral 1/5 portion of teeth smooth. Mid-oesophagus muscular; i mid-gut gland absent. Salivary gland single with paired ducts opening in buccal tube ventrally e w and anterior to radular diverticulum.
	Remarks.	n O
		l

The newly proposed family Charitodoronidae is distinctive in many ways. The shell bears no columellar folds and rather resembles a buccinoid shell; for this reason many Charitodoron species were originally erroneously placed in Columbella. The bulbous protoconch is unusual for Mitridae and suggests a non-planctotrophic development, consistent with the narrow

  distribution range. Remarkably, despite in morphology of the radula Charitodoron is close to mitrids, it does not possess the key morphological innovation of the Mitridae, the epiproboscis. The obvious morphological distinctiveness of Charitodoron, and its position in the molecular tree as a sister group to all other Mitridae justify its recognition as a separate family.

Genus Charitodoron Tomlin, 1932

	Species included: Charitodoron agulhasensis (Thiele, 1925) 3 , C. alcyone Lussi, 2009 3 ,
	F o
	r
	R e v i e w
	O n
	l y
	Type species: Charitodoron euphrosyne Tomlin, 1932 [junior subjective synonym of Columbella
	barbara Thiele, 1925]; OD.
	24

C. barbara (Thiele, 1925) 3

  , C. bathybius (Barnard, 1959)1 ; C. rosadoi Kilburn, 19953 , C. thalia

	Tomlin, 1932 3 , C. veneris (Barnard, 1964) 2 .
	Family Mitridae Swainson, 1831
	Subfamily Mitrinae Swainson, 1831
	(Figs 9-21)
	Genus Mitra Lamarck 1798 (Figs 9, 10 A)
	Type species: Voluta mitra Linnaeus, 1758, by tautonymy.
	Synonym: Tiarella Swainson, 1840. Type species: Voluta papalis Linnaeus, 1758; SD (Gray,
	1847).

C. kingtsio Huang, 2011 3

  C. chiangfucius[START_REF] Huang | Magnamitra n. gen. and nomenclatural remarks on large Cancilla and Mitra from Taiwan and the Philippines (Gastropoda: Mitridae)[END_REF] 2 comb. nov., C. christinae (Poppe, 2008) 1 comb. nov., C. glaphyria (Turner, 2001) 1 comb. nov., C. hilli (Cernohorsky, 1976) 3 comb. nov., C. invicta (Huang & Salisbury, 2017) 1 comb. nov., , C. labecula (Herrmann & Dekkers, 2009) 1 comb. nov., C. lussii

	comb. nov.,
	F o
	r
	R e v i e w
	O n
	l y
	3
	26

  . 'Eumitra' suduirauti Bozzetti, 19973 .

	Remarks. Eumitra was not included in our molecular analysis, and our understanding of this
	genus is largely based on published data
	F o
	r
	R e v i e w
	O n
	l y
	28

  The name refers to the now disused name Tiara Swainson, 1831, one of the genera first established in what later became the family Costellariidae.

	straight, and abapical half strongly convex. Inner aperture lip with 3-4 strong columellar folds,
	adapicalmost strongest. Shell dull, light to dark brown, sometimes with well developed
	periostracum. Radula with broad laterals, bearing robust, short and blunt cusps on their inner
	portion, turning gradually to fine serration on outer half (Figs 10 D, E). Rachidian with robust,
	very short and deeply rooted cusps, their tips merely reaching anterior margin of adjacent tooth
	base.
	Species included: Neotiara crenata (Broderip, 1836) 3 comb. nov., N. fultoni (E. A. Smith, 1892)
	3 comb. nov., N. gausapata (Reeve, 1845) 3 comb. nov., N. inca (d'Orbigny, 1841) 3 comb. nov.,
	N. lens (Wood, 1828) 1 comb. nov., N. muricata (Broderip, 1836) 3 comb. nov., N. nodulosa
	(Gmelin, 1791) 1 comb. nov., N. pallida (Nowell-Usticke, 1959) 3 comb. nov., N. rupicola F o (Reeve, 1844) 3 comb. nov., N. sphoni (Shasky & Campbell, 1964) 3 comb. nov.
	Etymology:	r
		R e v i e w
		O n
		l y
	Siphonal notch shallow or indistinct. Outer aperture lip evenly convex, or its adapical half about
		29

  Quoy & Gaimard, 1833) 2 comb. nov., P. gracilefragum[START_REF] Turner | Six new species of Mitridae from the Indian and Pacific Oceans, with remarks on Mitra abacophora Melvill, 1888 (Neogastropoda: Muricoidea)[END_REF] 2 comb. nov., P. kantori[START_REF] Poppe | New species of Mitridae and Costellariidae from the Philippines with additional information on the Philippine species in these families[END_REF] 3 comb. nov., P. kilburni[START_REF] Poppe | New species of Mitridae and Costellariidae from the Philippines with additional information on the Philippine species in these families[END_REF] 3 comb. nov., P. lienardi (G.B. Sowerby II, 1874) 3 comb. nov., P. maesta (Reeve, 1845) 1 comb. nov., P. oliverai(Poppe, 2008) 3 comb. nov., P. pediculus (Lamarck, 1811) 1 comb. nov., P. perdulca[START_REF] Poppe | New species of Mitridae and Costellariidae from the Philippines with additional information on the Philippine species in these families[END_REF] 3 comb. nov., P. proscissa (Reeve, 1844) 3 comb. nov., P. rubritincta (Reeve, 1844) 3 comb. nov., P. ruepellii (Reeve, 1844) 3 comb. nov., P. semiferruginea (Reeve, 1845) 3 comb. nov., P. silviae(Turner, 2007) 3 comb. nov., P. tabanula (Lamarck, 1811) 1 comb. nov., P. tornata (Reeve, 1845) 1 comb. nov., P. wareni[START_REF] Poppe | New species of Mitridae and Costellariidae from the Philippines with additional information on the Philippine species in these families[END_REF] 3 comb. nov., P. willani[START_REF] Poppe | New species of Mitridae and Costellariidae from the Philippines with additional information on the Philippine species in these families[END_REF] 3 comb. nov., P. yayanae(Huang, 2011) 3 comb. nov.

	P. fraga (
	F o
	r
	R e v i e w
	O n
	l y
	40

3

comb. nov., P. cingulata (A. Adams, 1853) 3 comb. nov., P. connectens (Dautzenberg & Bouge, 1923) 1 comb. nov., P. cucumerina (Lamarck, 1811) 3 comb. nov., P. cuyosae

(Poppe, 2008) 3

comb. nov., P. doliolum (Küster, 1839) 3 comb. nov., P. dovpeledi

(Turner, 1997) 3 comb. nov.,

. paupercula (Linnaeus, 1758) 1

  3 , S. aurora (Dohrn, 1861) 2 comb. nov., S. colombelliformis (Kiener, 1838) 2 , S. coronata (Lamarck, 1811) 1 comb. nov., S. crassicostata (G. B. Sowerby II, 1874) 3 comb. nov., S. decurtata (Reeve, 1844) 1 , S. fasciolaris (Deshayes, 1834) 3 comb. nov., S. flavocingulata (Lamy, 1938) 3 , S. fulvescens (Broderip, 1836) 1 comb. nov., S. holkosa (Li, Zhang & Li, 2005) 3 comb. nov., S. imperialis (Röding, 1798) 1 comb. nov., S. litterata (Lamarck, 1811) 1 , S. luctuosa (A. Adams, 1853) 1 , S. lugubris (Swainson, 1821) 1 comb. nov., S. nanus (Reeve, 1844) 3 comb. nov., S, S. pica (Dillwyn, 1817) 3 , S. retusa (Lamarck, 1811) 3 , S. scutulata (Gmelin, 1791) 1 comb. nov., S. subruppeli (Finlay, 1927) 3 comb. nov., S. telescopium (Reeve, 1844) 3 , Mitra testacea Broderip, 1836 3 comb. nov., S. ticaonica (Reeve, 1844) 3 comb. nov., S. tristis (Broderip, 1836) 1 , S. vexillum (Reeve, 1844) 1 , S. vultuosa (Reeve, 1845) 3 comb. nov., ?S. coffea (Schubert & J. A. Wagner, 1829) 2 comb. nov., ?S. pudica (Pease, 1860) 2 , ?S. tabida (Herrmann & Salisbury, 2013) 3 comb. nov.

  3 comb. nov., I. antillensis (Dall, 1889) 3 comb. nov., I. badia (Reeve, 1844) 3 comb. nov., I. beui (Thach, 2016) 3 comb. nov., I. carbonaria (Swainson, 1822) 1 comb. nov., I. chalybeia (Reeve, 1844) 3 comb. nov., I. chinensis (Gray, 1834) 3 comb. nov., I. cookii (G. B. Sowerby II, 1874) 3 comb. nov., I. cornea (Lamarck, 1811) 1 comb. nov., I. declivis (Reeve, 1844) 3 comb. nov., I.

	gabonensis (Biraghi, 1984) 3 comb. nov., I. glabra (Swainson, 1821) 1 comb. nov., I. joostei
	(Lussi, 2009) 3 comb. nov., I. lenhilli (Petuch, 1988) 3 comb. nov., I. midwayensis (Kosuge,
	1979) 3 comb. nov., I. nigra (Gmelin, 1791) 1 comb. nov., I. pele (Cernohorsky, 1970) 3 comb.
	nov., I. peterclarksoni (Marrow, 2013) 3 comb. nov., I. picta (Reeve, 1844) 3 comb. nov., I.

slacksmithae

(Marrow, 2013)

3 comb. nov., I. straminea (A. Adams, 1853) 1 comb. nov. I. swainsonii (Broderip, 1836) 3 comb. nov., I. turtoni (E. A. Smith, 1890) 3 comb. nov., I. ulala (Garcia, 2011) 3 comb. nov.

Table 1 .

 1

	84

Table 2 .

 2

				annealing	amplification	alignment	total for	analysed in 4G	substitution		gamm
	Gene	Primers	Reference	temperature	length bp	length bp	Mitridae	total (Mitridae)	model (AIC1) I	a
	COI HCO-1490 GGTCAACAAATCATAAAGAYATGYG	Folmer et al. (1994)	48-50	658	658	149	148 (74)	TIM+I+G	0.42 0.48
		LCO-2198 TAAACTTCAGGGTGACCAAARAAYCA	Folmer et al. (1994)
	16S 16SH CCGGTCTGAACTCAGATCACG	Palumbi (1996)	48-50	605		79	165 (86)	TIM+I+G	0.37 0.48
		16LC GTTTACCAAAAACATGGCTTC	Palumbi (1996)
	12S 12SI TGCCAGCAGCCGCGGTTA	Oliverio, Mariottini, 2001	57*	~540		76	148 (72)	TrN+I+G	0.23 0.54
		12SIII AGACGACGGGCGRTTWGTAC	Oliverio, Mariottini, 2002
	H3 H3F ATGGCTCGTACCAAGCAGACVGC	Colgan et al. (2000)	55	328	328	59	126 (67)	TrN+I+G	0.63 0.78
		H3R ATATCCTTRGGCATRATRGTGAC	Colgan et al. (2000)

  Species of Profundimitra n. gen., Fusidomiporta n. gen. and Cancillopsis n. sp. A -Radulae of Pseudonebularia n. gen. and Gemmulimitra n. gen. A, B. Cancilla cf. fibula, MNHN IM-2013-40646, New Caledonia, EXBODI Stn CP3822, 21°52'S, 166°51'E, 341-506 m, 37.6 mm; F. C. schepmani, MNHN IM-2013-40645, New Caledonia, EXBODI Stn DW3926, 18°35'S, 164°20'E, 364-473 m, 15.6 mm. Radulae in the genera Imbricariopsis, Swainsonia, Scabricola and Neocancilla. A. Radulae in the genera Carinomitra n. gen., Condylomitra n. gen. and Probata. A. Carinomitra typha var. micans, MNHN, New Caledonia, Noumea area, LAGON Stn 1359, 22°19.9'S, 166°13.2'E, 25-30 m, 9.5 mm; C. Carinomitra saltata, MNHN,

	8. Charitodoron species. A. Charitodoron sp., MNHN IM-2007-38306, off Mozambique, MAINBAZA Stn 3157, 21°46'S, 36°25'E, 1410-1416 m; B, C. C. bathybius, Figure 11. Genus Calcimitra. A, B. C. glaphyria; A. MNHN IM-2007-30066, Philippines, PANGLAO 2005 Stn CP2342, 9°24'N 123°53'E, 1240-1258 m; B. MNHN IM-2013-40650, Papua New Guinea, BIOPAPUA Stn CP3728, 07°52'S, 148°01'E, 498-501 m, 31.9 mm; C. C. triplicata, MNHN IM-2007-38186, Madagascar, MIRIKY Stn CP3253, 15°25'S, 45°55'E, 943-950 m,; D. C. labecula, MNHN IM-2007-30127, Coral Sea, EBISCO Stn DW2515, 24°04'S, 159°41'E, 330-370 m; E, F. C. invicta; E. MNHN IM-2007-32130, Philippines, PANGLAO Greece, Astypalea Is, Agrilidi, 36°35.0'N, 26°25.4'E, 2-7 m, 12.1 mm; G, H. MNHN IM-2013-40661, Italy, Sicily, Syracuse, 37°00.5'N, 15°18.6'E, 6-12 m, 9.0 mm; G. Shell; H. Protoconch. Figure 15. Quasimitra species. A. Quasimitra sanguinolenta, MNHN IM-2013-40677, Mozambique, INHACA 2011 Stn MS02, 26°06.3'S, 32°58.0'E, 17-19 m, 28.7 mm; B. Q. cardinalis, MNHN IM-2013-40681, Papua New Guinea, KAVIENG 2014 Stn KR178, 02°45.2'S 2013-40665, Papua New Guinea, BIOPAPUA Stn CP3692, 02°10'S, 147°19'E, 408-448 m, 25.2 mm. Figure 18. C. Profundimitra taylori n. sp. A, B. Holotype, MNHN IM-2013-40651, Papua New Guinea, BIOPAPUA Stn CP3671, 04°04'S, 151°56'E, 585-601 m, 24.2 mm; C. New Caledonia, Figure 24. Imbricaria species. A. Imbricaria conularis, MNHN IM-2013-18043, PAPUA NIUGINI Stn PR203, 05°10.3'S 145°48.5'E, 1-19 m, 16.8 mm; B. I. bantamensis, MNHN IM-2013-16688, PAPUA NIUGINI Stn PD67, 05°15.5'S, 145°46.8'E, 2-6 m, 21.8 mm; C. I. insculpta, MNHN IM-2013-40657, Vietnam, Nha-Trang Bay, South Mot Is, 12°10.4'N 109°16.3'E, 15-18 m, 18.3 mm; D. I. flammigera, MNHN IM-2013-40656, Vietnam, Nha-Trang 23.75 mm; D. MNHN IM-2007-38688, Society Is, TARASOC Stn DW3441, 16°41.7'S, Figure 30. Neocancilla species. A. Neocancilla papilio, MNHN IM-2013-12560, PAPUA NIUGINI Stn PR24, 05°12.3'S, 145°48.8'E, 28.2 mm; B. N. clathrus, MNHN IM-2007-30178, Vanuatu, SANTO 2006 Stn DR73, 15°22.5'S, 167°11.3'E, 10-25 m; C. N. rufescens, MNHN IM-2013-40644, Mozambique, INHACA 2011 Stn MD22, 25°59.7'S, 32°46.8'E, 22 m, 16.4 mm; D. N. hemmenae, holotype, NHMUK 1992088, off Somalia, 40.8 mm. 1967709, locality not known, 51.5 mm; D. I. carbonaria, molecular voucher, AMS C.335424, Australia, North Sydney, Collaroy, Long Reef, 33°44.7'S , 151°19' E, intertidal; E. I. chinensis, lectotype, NHMUK 1967708, "China", 58.6 mm; F. I. nigra, MNHN IM-2013-40676, Congo, Mission ZANAGA Stn DV08, 04°42.9'S, 11°47.5'E, 14-15 m, 45.2 mm; G. I. cornea, BAU2537.1, Italy, Giannutri Is., Punta San Francesco, 42°15.1'N, 11°07'E, 10-20 m, 36.75 mm; Figure 38. Carinomitra peculiaris, MNHN IM-2013-6168, PAPUA NIUGINI Stn PR218, 05°07.3'S, 145°49.4'E, 11.1 mm; B. Society Is, off Moorea, 5.4 mm; D. Condylomitra tuberosa, MNHN, Philippines, PANGLAO 151°26.1'W, 350-360 m; E. Figure 27. Species of Imbricariopsis n. gen. A. Imbricariopsis punctata, MNHN IM-2013-H. Isara turtoni, lectotype of Mitra turtoni, NHMUK 1889.1.10.-97, St. Helena Is., 26.2 mm; I. 2004 Stn S12, 9°29.4'N, 123°56.0'E, 6-8 m, 14.4 mm; E. Condylomitra bernhardina New
	MNHN IM-2013-40658, off Mozambique, MAINBAZA, Stn 3171, 25°59'S, 34°42'E, 771-776 2005 Stn CP2383, 08°45'N, 123°18'E, 338-351 m, *** mm; F. MNHN IM-2013-15373, PAPUA 150°43.1'E, 2-46 m, 58.8 mm; C. Q. sophiae, MNHN IM-2007-30056, Vanuatu, SANTO 2006 BATHUS 4 Stn DW920, 18°45'S, 163°17'E, 610-620 m, MNHN, 15.1 mm; D. Profundimitra n. Bay, South Mot I., 12°10.4'N 109°16.3'E, 15-18 m, 14.8 mm; E. I. fulgetrum, MNHN IM-2013-11604, PAPUA NIUGINI Stn PM12, 05°00.2'S, 145°47.6'E, 0-1 m, 17.2 mm; B. I. conovula, Figure 31. Pterygia species. A. Pterygia dactylus, MNHN IM-2013-14989, PAPUA NIUGINI I. straminea, MNHN IM-2013-56423, French Guiana, GUYANE Stn CP4375, 06°39'N, Caledonia, Noumea area, LAGON Stn 1343, 22°17.8'S, 166°19.9'E, 7 m, 16.1 mm; F. Probata
	m, 12.0 mm; B. Shell; C. Radula; D, E. C. veneris; D. NMSA D4172, off Durban 29°50.4'S, NIUGINI Stn CP3949, 05°12'S, 145°51'E, 380-407 m, 61.6 mm; G, H. C. christinae, G. MNHN Stn DR09, 15°34.6'S, 167°13.8'E, 12 m, 25.1 mm; M. Q. puncticulata, MNHN IM-2013-40685, sp. aff. taylori, Papua New Guinea, BIOPAPUA Stn CP3636, 07°27'S, 147°31'E, 462-495 m, 18112, PAPUA NIUGINI Stn PR240, 05°08.2'S, 145°48.7'E, 3-20 m, 16.8 mm; F. I. verrucosa, lectotype of Mitra virgo, NHMUK 1967912, Society Is,14 mm; C. I. vanikorensis, lectotype, Stn PM39, 05°12.1'S, 145°48.4'E, intertidal, 25.1 mm; B. P. sinensis, MNHN IM-2009-15439, 52°30'W, 195-200 m, 33.5 mm. barbadensis, MNHN IM-2013-7772, Guadeloupe, KARUBENTHOS Stn GR36, 16°30.6'N,
	31°14.0'E, 150 m, 14.1 mm (voucher of sectioned specimen); E. NMSA E. 9237, northern Natal, IM-2007-32131, Philippines, PANGLAO 2005 Stn CP2383, 08°45'N, 123°18'E, 338-351 m; H. Philippines, off Noc-Nocan Is, 10°15.1'N 124°25.1'E, 4-12 m, 40.6 mm. 26.8 mm; E. Profundimitra planofilum, MNHN IM-2007-32149, Philippines, PANGLAO 2005 MNHN IM-2013-14284, PAPUA NIUGINI Stn PD39, 04°59.3'S, 145°47.5'E, 2-4 m, 11.6 mm; MNHN IM-2000-30312, Vanikoro, Solomon Is, 22.5 mm; D. Mitra carbonacea, Senegal, off Madagascar, ATIMO VATAE Stn CP3568, 25°04.7'S, 47°03.4'E, 64-65 m; C. P. fenestrata, 61°28.4'W, 12 m, 20.6 mm.
	off Shellfield Beach, 29°39.8'S, 31°30.2'E, 150 m, radula. MNHN IM-2007-32133, Philippines, PANGLAO 2005 Stn CP2359, 8°50'N 123°35'E, 437-476 Stn CP2398, 9°33'N, 123°40'E, 713-731 m; F. Profundimitra n. sp. aff. planofilum, MNHN, IM-G. I. salisburyi, Papua New Guinea, Hansa Bay, 8-18 m, 15.3 mm; H. I. hrdlickae, MNHN IM-Gorée I., 10-15 m, 20.5 mm. MNHN IM-2007-30039, Vanuatu, SANTO 2006 Stn DR09, 15°34.5'S, 167°13.5'E, 12 m; D. P. Figure 35. Radulae in the subfamily Isarinae. A. Isara glabra, MNHN IM-2013-40640,
	Figure 9. Mitra species. A. Mitra mitra, MNHN IM-2013-40639, Mozambique, INHACA 2011 Stn MR15, 26°00.0'S, 32°54.4'E, 4 m, 89.1 mm; B. M. papalis, MNHN IM-2013-50981, Papua New Guinea, KAVIENG Stn KR34, 02°34,6'S, 150°46,3'E, 10-22 m, 70.7 mm; C. M. stictica, Society Islands, off Moorea, 61.0 mm, MNHN; D. M. deprofundis, holotype, MNHN IM-2000-30197, New Caledonia, Antigonia Bank, SMIB 4, Stn DW57, 23°21'S, 168°04'E, 210-260 m, 67.5 mm. Figure 10. Radulae of Mitrinae species. A. Mitra mitra, MNHN IM-2013-40639, Mozambique, F o r R e v i e w O n l y m. Figure 12. Eumitra species. A. Eumitra apheles, MNHN, New Caledonia, BATHUS 4 Stn 918, 18°49'S, 163°16'E, 613-637 m, 18.6 mm; B. E. caledonica, holotype, MNHN-IM-2000-30181, Norfolk Ridge, BIOCAL Stn CP61, 24°11'S, 167°32'E, 1070 m, 29.9 mm; C. E. imbricata, holotype, MNHN-IM-2000-30242, Coral Sea, Lansdowne-Fairway, CORAIL 2 Stn DE14, 21°01'S, 160°57'E, 650-660 m, 15.5 mm; D, E. Eumitra n. sp.; D. Morphology voucher, MNHN, KARUBAR Stn CC56, 08°16'S, 131°59'E, 549-552 m, 48.75mm; E. 'Eumitra' suduirauti holotype, MNHN IM-2000-30306, Philippines, Mindanao, off Tanala, 500-600 m, 51.7 mm. Figure 13. Neotiara species. A, B. Neotiara lens A. Neotype, MNHN IM-2000-33145, Panama, Panama City, 08°95'N, 79°53'W, intertidal, 40.0 mm; B. BAU 2532, same locality as previous, 33.6 mm; C. N. lignaria, holotype, NHMUK 1967798, Sta Elena, Ecuador, 47.6 mm; D. N. rupicola, syntype, NHMUK 1964460, Sta Elena, Ecuador, 37.2 mm; E. N. gausapata, lectotype, NHMUK 1967769/1-2, Galapagos Is, 11.7 mm; F. N. nodulosa, MNHN IM-2013-9546, Guadeloupe, KARUBENTHOS Stn GM10, 16°12,53'N, 61°25,55'W, 1 m, 27.8 mm; G. N. muricata, lectotype, NHMUK 1967610/1-3, Galapagos Is, 29 mm; H. N. sp. cf. inca, BAU 0279-1, Panama, Venado Beach, 8°53.5'N, 79°36.4'W, intertidal, 11 mm. Figure 14. Ziba and Episcomitra species. A, B. Ziba carinata; A. lectotype of Mitra senegalensis, NHMUK 1967881, Senegal, 27.9 mm; B. MNHN IM-2013-40647, Congo, Mission ZANAGA, Stn 531DW, 04°43.0'S 11°47.0'E, 17 m, 24.5 mm; C, D. Z. gambiana, syntype, NHMUK 1967768, "Senegambia", 28 mm; E. Episcomitra zonata, BAU2538, Italy, off F o r R e v i e w O n l y Figure 16. Species of Domiporta and Roseomitra. A. Domiporta filaris, MNHN IM-2013-12956, Papua New Guinea, PAPUA NIUGINI Stn PD 23, 05°06'S, 145°49.2'E, 3-7 m, 15.8 mm; B. D. praestantissima, MNHN IM-2013-40642, Vietnam, Nha-Trang Bay, West of Mun Is, 12°10.1'N, 109°17.8'E, 6-18 m, 25.6 mm; C. D. circula, MNHN IM-2007-30018, Vanuatu, SANTO 2006 Stn LD24, 15°31.4'S, 167°10.0'E, 47 m; D. D. cf. carnicolor, MNHN IM-2013-40684, Philippines, off Noc-Nocan Is, 10°15.1'N, 124°25.1'E, 4-12 m, 23.9 mm; E. D. granatina, MNHN IM-2013-40641, Mozambique, INHACA 2011 Stn MR15, 26°00.0'S, 32°54.4'E, 4 m, 34.2 mm; F. Roseomitra millepunctata, neotype, MNHN IM-2013-40686, Philippines, off Noc-Nocan Is, 10°15.1'N, 124°25.1'E, 4-12 m, 29.8 mm; G. R. incarnata, lectotype, NHMUK 196779/1-3, Misamis Is, Mindanao, Philippines, 21.6 mm; H. R. tagaroae, Philippines, Olango Channel, dredged from 80 m, 9.5 mm; I. R. rosacea, Mozambique, INHACA 2011 Stn MD13, 26°03.1'S 33°01.0'E, 50-53 m, 7.9 mm. Figure 17. Radulae in the Domiporta clade. A. Domiporta cf. carnicolor, MNHN IM-2013-40684, Philippines, off Noc-Nocan Is, 10°15.1'N, 124°25.1'E, 4-12 m, 23.9 mm; B. D. granatina, MNHN IM-2013-40641, Mozambique, INHACA 2011 Stn MR15, 26°00.0'S, 32°54.4'E, 4 m, 34.2 mm; C. D. praestantissima, MNHN IM-2013-40642, Vietnam, Nha-Trang Bay, West of Mun Is, 12°10.1'N, 109°17.8'E, 6-18 m, 25.6 mm; D. D. filaris, MNHN, Mozambique, INHACA 2011 Stn MR13, 25°59.7'S, 32°54.5'E, 2-5 m, 17.4 mm; E. Roseomitra rosacea, MNHN, Mozambique, INHACA 2011 Stn MD13, 26°03.1'S, 33°01.0'E, 50-53 m, 7.9 mm; F. R. millepunctata, MNHN, IM-2013-40686, Philippines, off Noc-Nocan Is, 10°15.1'N, 124°25.1'E, 4-12 m, 29.8 mm; G. Profundimitra taylori n. sp., holotype, MNHN, IM-2013-40651, Papua New Guinea, BIOPAPUA Stn CP3671, 04°04'S, 151°56'E, 585-601 m, 24.2 mm; F o r R e v i e w O n l y 2013-40654, New Caledonia, EXBODI Stn CP3829, 22°02'S, 167°05'E, 350-360 m, 16.9 mm; G. Profundimitra n. sp., MNHN IM-2007-35711, Philippines, AURORA 2007 Stn CP2658, 15°58'N, 121°49'E, 422-431 m; H. Profundimitra n. sp., MNHN IM-2007-36757, Madagascar, MIRIKY Stn CP3289, 14°29'S, 47°26'E, 332-379 m; I, K. Fusidomiporta ponderi n. sp., holotype, MNHN, IM-2013-40665, Papua New Guinea, BIOPAPUA Stn CP3692, 02°10'S, 147°19'E, 408-448 m, 25.2 mm; L. Fusidomiporta n. sp., MNHN IM-2007-32153, Philippines, PANGLAO 2005 Stn DW2364, 09°01'N, 123°25'E, 427 m; M. Cancillopsis meimiaoae MNHN IM-2007-30011, Vanuatu, SANTO' 2006, Stn AT19, 15°40'48''S 167°00'30''E, 503-600 m. Figure 19. Species of Pseudonebularia n. gen. A, B. Pseudonebularia tornata, A. Lectotype of Mitra tornata, NHMUK 1967897, Philippines, 18.1 mm; B. MNHN IM-2013-12538, PAPUA NIUGINI Stn PB14, 05°13.8'S, 145°48'E, 15 m, 16.5 mm; C. P. atjehensis, MNHN IM-2007-30264, Vanuatu, SANTO 2006 Stn EP01, 15°32.5'S, 167°09'E, 46-47 m; D. P. maesta, MNHN IM-2013-40648, Madagascar, MIRIKY Stn CP3288, 14°31.9' S, 47°26.5' E, 46-54 m, 15.6 mm; E. P. connectens, MNHN IM-2013-2342, PAPUA NIUGINI Stn PB16, 05°10.7'S, 145°47.7'E, intertidal, 10.4 mm; F. P. pediculus, MNHN IM-2013-12705, PAPUA NIUGINI Stn PB15, 05°04.7'S, 145°48.9'E, 5 m, 11.6 mm; G. P. gracilefragum, Philippines, Olango I., Caw Oy, 10-25 m, 20.1 mm. Figure 20. Pseudonebularia pediculus, MNHN IM-2013-12705, PAPUA NIUGINI Stn PB15, 05°04.7'S, 145°48.9'E, 5 m, 11.6 mm; C. P. connectens, MNHN IM-2013-2342, PAPUA NIUGINI Stn F o r R e v i e w O n l y F o r R e v i e w O n l y 2013-18020, PAPUA NIUGINI Stn PD75, 05°14.2'S, 145°47.8'E, 10-25 m, 16.5 mm; I. I. rufogyrata, MNHN IM-2007-36131, Solomon Is, SALOMONBOA 3 Stn DW2852, 09°46'S, 160°51'E, 220 m; K. I. pugnaxa, MNHN IM-2007-32124, Philippines, PANGLAO 2005 Stn CP2378, 08°38.7'N, 123°20.1'E, 65 m. BIOPAPUA Stn DW3641, 06°45'S, 148°01'E, 380-476 m, 20.8 mm; K. C. cf. fibula, MNHN IM-2013-40646, New Caledonia, EXBODI Stn CP3822, 21°52'S, 166°51'E, 341-506 m, 37.6 mm; L. C. baueri, MNHN IM-2013-58853, Papua New Guinea, KAVIENG 2014 Stn DW4484, 02°26'S, 149°55'E, 229 m, 34.2 mm; M. C. schepmani, MNHN IM-2013-40645, New Caledonia, EXBODI Stn DW3926, 18°35'S, 164°20'E, 364-473 m, 15.6 mm. Figure 26. Cancilla species. A. Cancilla baeri, IM-2013-58853, Papua New Guinea, KAVIENG F o r R e v i e w O n l y Figure 28. Imbricariopsis punctata, Mozambique, INHACA 2011 Stn MM3, 26°03.9'S, 32°57.3'E, 0-1 m, Beach, 12°10.1'N, 109°17.8'E, 6-18 m, 13.2 mm; G. Neocancilla papilio, Society Is, off Moorea, 36.6 mm; H. N. clathrus, Mozambique, INHACA 2011 Stn MR13, 25°59.7'S, 32°54.5'E, 2-5 m, 14.9 mm; I. N. rufescens, MNHN IM-2013-40644, Mozambique, INHACA 2011 Stn MD22, 25°59.7'S, 32°46.8'E, 22 m, 16.4 mm. Figure 29. Genera Scabricola and Swainsonia. A. Scabricola variegata, MNHN IM-2013-40683, Philippines, off Noc-Nocan Is, 10°15.1'N, 124°25.1'E, 4-12 m, 45.5 mm; B. Scabricola desetangsii, MNHN IM-2013-50703, Papua New Guinea, KAVIENG 2014 Stn KR14, 02°42.3'S 150°39.1'E, 7-10 m, 24.6 mm; C. Scabricola olivaeformis, MNHN IM-2013-18062, PAPUA NIUGINI Stn PR214, 05°10.2'S, 145°50.4'E, 1-8 m, 13 mm; D. Scabricola coriacea, syntype, NHMUK 1967728, Corrigidor I., Philippines, 11.1 mm; E. Scabricola albina, holotype, NHMUK 1966663, Batangas, Luzon I., Philippines, 26.7 mm; F. Scabricola potensis, lectotype, MNHN IM-2000-30283, New Caledonia, Belep Is, 24.5 mm; G. Swainsonia fissurata, Israel, off Eilat, 2-4 m, 63.1 mm,; H. Swainsonia fusca, MNHN IM-2013-40643, Mozambique, INHACA o r O n l y nucea, MNHN IM-2013-50702, Papua New Guinea, KAVIENG 2014 Stn KM04, 02°42'S, 150°44'E, 0-1 m, 42 mm. eremitarum, MNHN IN-2013-16323, PAPUA NIUGINI Stn PM51, 05°05.5'S, 145°48.6'E, 0-1 m, 48.4 mm; D. N. incompta, Philippines, Olango I., Cow-Oy, 10-25 m, 48.0 mm; E. N. edentula, 26.7 mm, Mozambique, Matibane Bay, Choca, 2-4 m; F. N. ustulata, Syntype of Mitra ustulata, NHMUK 1967918, ?Viti Is [Fiji], 34.2 mm; G. N. ancillides, holotype, NHMUK 1967712, "I. of Annaa" [Anaa, Tuamotu Is], 23.6 mm; H. N. acuminata, MNHN IM-2013-13288, PAPUA NIUGINI Stn PM25, 05°01.1'S, 145°47.9'E, intertidal, 23 mm; I. N. coarctata, syntype, NHMUK 1967722, locality unknown, 17.5 mm. F o r O n l y Esperance, WESTERN AUSTRALIA Stn WE01, 15°16.94'S, 124°06.3'E, 11 m, 62.7 mm; B. I. nigra, MNHN IM-2013-40676, Congo, Mission ZANAGA Stn DV08, 04°42.9'S, 11°47.5'E, 14-15 m, 45.2 mm; C. I. straminea, MNHN IM-2013-56423, French Guiana, GUYANE Stn CP4375, 06°39'N, 52°30'W, 195-200 m, 33.5 mm; D. Subcancilla attenuata, BAU392, Panama, MNHN IM-2013-61266, Guadeloupe, KARUBENTHOS 2 Stn DW4626, 15°57'N, 61°37'W, 210-233 m, 23 mm; D. Subcancilla attenuata, BAU 0392, Panama, Las Perlas, Contadora I., 08°36.2'N, 79°00.2'W, 22 m, 24 mm; E. Subcancilla gigantea, syntype, NHMUK 1964463, Xipixapi, Pacific coast of Columbia, 69.7 mm. Figure 37. Species of Carinomitra n. gen. A, B. Carinomitra peculiaris; A. syntype, NHMUK 1967833, Puerto Galera, Mindanao, Philippines, 10 mm; B. MNHN IM-2013-6168, PAPUA NIUGINI Stn PR218, 05°07.3'S, 145°49.4'E, 11.1 mm; C -E. C. typha; C. syntype, NHMUK 1967904, Philippines, 9.5 mm; D. C. typha var. micans, MNHN, New Caledonia, Noumea area, LAGON Stn 1359, 22°19.9'S, 166°13.2'E, 25-30 m, 9.5 mm; E. MNHN IM-2013-4030, PAPUA NIUGINI, Stn PB28, 05°11.9'S, 145°49.6'E, 10 m, 8.6 mm. F o r O n l y Figure 39. Species of Condylomitra n. gen. A, B. Condylomitra tuberosa; A. Syntype, NHMUK 1967899, Philippines, 14.4 mm; B. MNHN IM-2007-30311, Philippines, PANGLAO 2004 Stn R75, 09°32.8'N, 123°42.1'E, 3-35 m; C, D. C. bernhardina; C. MNHN IM-2013-54115, Papua New Guinea, KAVIENG 2014 Stn KB40, 02°36.6'S, 150°32.9'E, 2-7 m; D. MNHN, New 2011 Stn MS08, 25°59.F R e v i e w 2011 Stn MR24, 25°54.5'S, 33°02.8'E, 23-26 m, 29.9 mm; F. N. acuminata, MNHN IM-2013-13288, PAPUA NIUGINI Stn PM25, 05°01.1'S, 145°47.9'E, intertidal, 23 mm. R e v i e w Las Perlas, Contadora I., 08°36.2'N, 79°00.2'W, 22 m, 24 mm; E. Subcancilla cf. leonardhilli, MNHN IM-2013-61266, Guadeloupe, KARUBENTHOS 2 Stn DW4637, 15°52'N, 61°20'W, 217-225 m; F. Subcancilla erythrogramma, BAU424, Panama, Las Perlas, Contadora I., 08°36.6'N, 79°00.2'W, 25 m, 24.1 mm. w e i v e R Caledonia, Noumea area, LAGON Stn 1343, 22°17.8'S, 166°19.9'E, 7 m, 16.1 mm.
	Chioggia, 45°12'N, 12°40'E, 40 m, 96.2 mm; F -H. Episcomitra cornicula F. BAU1578.3, H. Profundimitra n. sp., MNHN, IM-2013-40654, New Caledonia, EXBODI Stn CP3829,
	22°02'S, 167°05'E, 350-360 m, 16.9 mm; I. Fusidomiporta ponderi n. sp., holotype, MNHN IM-
	92 93 94 96 97 98 99 100

PB16, 05°10.7'S, 145°47.7

'E, intertidal, 10.4 mm; D, E. P. maesta, MNHN IM-2013-40648,

Madagascar, MIRIKY Stn CP3288, 14°31.9' S, 47°26.5

' E, 46-54 m, 15.6

mm; F. Gemmulimitra neocaledonica, MNHN, Paratype 5, New Caledonia, EXBODI Stn CP3814, 21°49'S, 166°44'E, Figure 25. Radulae in the genera Imbricaria and Cancilla. A. I. pugnaxa, MNHN, New Caledonia, EXBODI Stn CP3795, 21°32'S, 166°21'E, 240-245 m, 31.9 mm; B. I. conularis, MNHN, Society Is, Moorea, 13.3 mm; C. I. fulgetrum, MNHN IM-2013-18112, PAPUA NIUGINI Stn PR240, 05°08.2'S, 145°48.7'E, 3-20 m, 16.8 mm; D. I. flammigera, MNHN IM-2013-40656, Vietnam, Nha-Trang Bay, South Mot I., 12°10.4'N 109°16.3'E, 15-18 m, 14.8 mm; E. I. insculpta, MNHN IM-2013-40657, Vietnam, Nha-Trang Bay, South Mot Is, 12°10.4'N 109°16.

3

'E, 15-18 m, 18.3 mm; F. I. verrucosa, MNHN, Papua New Guinea, KAVIENG 2014

Stn KR54, 02°42.3'S, 150°39.1

'E,[7][START_REF] Sanguinolenta | MNHN IM-2013-40677, Mozambique[END_REF][9][10] 12.3 mm; G. I. salisburyi, Papua New Guinea, Hansa Bay,[START_REF] Sanguinolenta | MNHN IM-2013-40677, Mozambique[END_REF][9][10][11][12][13][14][15][16][17][18] 15.3 mm; H. I. interlirata, Vietnam, Murray Beach, 12°10.1'N, 109°17.8'E,[START_REF] Neotiara | BAU1578.3, Greece, Astypalea Is[END_REF][7][START_REF] Sanguinolenta | MNHN IM-2013-40677, Mozambique[END_REF][9][10][11][12][13][14][15][16][17][18] 28.6

mm; I. Cancilla fibula / rehderi, MNHN, Papua New Guinea, 2014 Stn DW4484, 02°26'S, 149°55'E, 229 m, 34.2 mm; B -D. Cancilla fibula / rehderi; B. MNHN, Papua New Guinea, BIOPAPUA Stn DW3641, 06°45'S, 148°01'E, 380-476 m, 20.8 mm; C. MNHN IM-2013-19073, PAPUA NIUGINI Stn CP4059, 02°38'S, 141°18'E, 335 m, 14.7 mm; B. Swainsonia fusca, MNHN IM-2013-40643; C. Swainsonia casta, Papua New Guinea, Hansa Bay, 8-18 m, 25.6 mm; D. Scabricola desetangsii, MNHN IM-2013-50703, Papua New Guinea, KAVIENG 2014 Stn KR14, 02°42.3'S 150°39.1'E, 7-10 m, 24.6 mm; E. Scabricola variegata, MNHN IM-2013-40683, Philippines, off Noc-Nocan Is, 10°15.1'N, 124°25.1'E, 4-12 m, 45.5 mm; F. Scabricola olivaeformis, Vietnam, Nha-Trang Bay, Murray 5'S, 32°52.9'E, 9-17 m, 18.2 mm; I. Swainsonia casta, MNHN IM-2013-48000, Papua New Guinea, KAVIENG 2014 Stn KS17, 02°36.2'S, 150°46.3'E, 3 m, 30.5 mm.

Figure 32. Radulae in the subfamily Cylindromitrinae. A. Pterygia dactylus, MNHN IM-2013-14989, PAPUA NIUGINI Stn PM39, 05°12.1'S, 145°48.4'E, intertidal, 25.1 mm; B. P. fenestrata, Society Is, off Moorea, 11.5 mm; C. P. nucea, MNHN IM-2013-50702, Papua New Guinea, KAVIENG 2014 Stn KM04, 02°42'S, 150°44'E, 0-1 m, 42 mm; D. Nebularia incompta, Philippines, Olango I. Cow-Oy, 10-25 m, 48.0 mm; E. N. contracta, Mozambique, INHACA Figure 33. Nebularia species. A. Nebularia contracta, MNHN IM-2013-10345, PAPUA NIUGINI Stn PR04, 05°10.1'S, 145°50.5'E, 30m, 19.7 mm; B. N. chrysostoma, syntype, NHMUK 1967715, "Isle of Annaa, South Seas" [Anaa, Tuamotu Is], 51.3 mm; C. N.

Figure 34. Isara species. A. Isara glabra, MNHN IM-2013-40640, Esperance, WESTERN AUSTRALIA Stn WE01, 15°16.94'S, 124°06.3'E, 11 m, 62.7 mm; B. Holotype of Mitra bulimoides, NHMUK 1966656, locality not known, 33.2 mm; C. I. chalybeia, syntype, NHMUK Figure 36. Subcancilla species. A. Subcancilla erythrogramma, BAU 0424, Panama, Las Perlas, Contadora I., 08°36.6'N, 79°00.2'W, 25 m, 24.1 mm; B. Subcancilla funiculata, lectotype, NHMUK 1967767, Plata I., Pacific coast of Columbia, 27.2 mm; C. Subcancilla cf. leonardhilli, Figure 40. Genera Atrimitra, Vicimitra, Probata, and Dibaphimitra. A. Atrimitra idae, holotype, NMW.1955.158.00066, Point Loma, Lower California, Mexico, 60.0 mm; B. Atrimitra effusa, lectotype, NHMUK 1966415, Guacomayo, Central America, 31.6 mm; C. Atrimitra caliginosa, holotype, NHMUK 1966718, locality unknown, 34.9 mm; D. Probata espinosai, paratype, MNHN-IM-2012-37719, in front of Marianao, Havana, Cuba, 10 mm; E. Probata barbadensis, MNHN IM-2013-7772, Guadeloupe, KARUBENTHOS Stn GR36, 16°30.6'N, 61°28.4'W, 12 m, 20.6 mm; F. Dibaphimitra florida, holotype of Mitra fergusoni G.B. Sowerby II, 1874, NHMUK

1879.2.26.133, Panama, 42.7 mm.

Zoological Journal of the Linnean Society

Acknowledgements

The molecular material in this paper originates from numerous shore-based expeditions and deep sea cruises, conducted respectively by MNHN and Pro-Natura International (PNI) as part of the Our Planet Reviewed programme, by MNHN and Institut de Recherche pour le Développement (IRD) as part of the Tropical Deep-Sea Benthos programme, and by the Department of Biology and Biotechnology 'Charles Darwin', Sapienza University, Rome. Funders and sponsors include the Total Foundation, Prince Albert II of Monaco Foundation, Stavros Niarchos Foundation, Richard Lounsbery Foundation, the French Ministry of Foreign Affairs, and the Philippines Bureau of Fisheries and Aquatic Research (BFAR). Sampling in Western Australia was arranged by Hugh Morrison, with support of the Western Australian Museum. We are also grateful to the staff of the joint Russian-Vietnamese Tropical Center for the opportunity to collect specimens in Nha-Trang Bay. We thank, among others, Virginie Héros, Philippe Maestrati, Pierre Lozouet, Barbara Buge, Ellen Strong, Laurent Charles and Julien Brisset for their role in specimens processing during the expeditions and curation. We thank Catherine Rausch (MNHN) and Nadezda Surovenkova (IPEE) for access to SEM facilities, and Ilya Ryazanov for help with molecular facilities at SIEE RAS. We are grateful to Guido Poppe and Sheila Tagaro (Conchology Inc.), Mandy Reid and Alison Miller (AMS), who provided photos of mitrid types. We are thankful to Andreia Salvador for access to the NHMUK type collection and Manuel Caballier project E-RECOLNAT (ANR-11-INBS-0004) for photos of the MNHN types. The present study was largely accomplished during visiting curatorships of the first author to MNHN in 2013, 2014, 2015 and 2016, supported also by a 2014 Metchnikov Postdoctoral Fellowship granted by the French Embassy in Russia. The molecular phylogenetic studies were supported by the "Service de Systématique Moléculaire" (UMS 2700 CNRS-MNHN), the ATM Barcode (PIs: Sarah Samadi and Jean-Noël Labat / MNHN) and by grant from the Russian Science Foundation RSF 16-14-10118.

Distribution: Bismarck Sea off mainland New Guinea and New Ireland; East of New Caledonia, Norfolk Ridge, 580-760m.

Etymology: The species named after our friend and colleague John D. Taylor of the NHMUK in recognition of his immense contribution to malacology, and in particular to our knowledge of miter feeding biology.

Remarks: Profundimitra taylori n. sp. can be readily distinguished from its congeners by its almost smooth shell with faint columellar folds; both features are rather characteristic of Eumitra. Members of the latter genus, however, are characterized by more convex whorls, often with subcylindrical early teleoconch whorls. Smooth species of Calcimitra, such as C. labecula, resemble P. taylori n. sp. in shell proportions and sculpture; however, Calcimitra species can be recognized by their generally larger shells with strong columellar folds and, typically, by the presence of a siphonal notch. Finally, the smooth shallow water Quasimitra sarmientoi, although close to Profundimitra taylori n. sp. in shell proportions and lack of sculpture, can be differentiated by the intensely coloured shell and distinct siphonal notch in adult specimens.

Apart from the mentioned shell characters, the characteristic radula with similar laterals and rachidian differentiates Profundimitra taylori n. sp. and its congeners from most other mitrids.

Fusidomiporta ponderi Fedosov, Herrmann, Kantor & Bouchet, n. sp.

Type data

Holotype. MNHN IM-2013-40665, Papua New Guinea, BIOPAPUA Stn CP3692, SE of Manus I., 02°10'S, 147°19 'E, lv, 25.2 mm (Figs 18 I, K).

Paratype. BIOPAPUA Stn CP3629, Gulf of Huon, South of Lae, 06°57'S, 147°08 'E, 240-269 m, dd, 26.15 mm.