Alexander E Fedosov

Gavin Malcolm

Yves Terryn

Juliette Gorson

Maria-Vittoria Modica

Mande Holford

Nicolas Puillandre
email: nicolaspuillandre@gmail.com

Phylogenetic classification of the family Terebridae (Gastropoda: Mollusca)

The conoidean family Terebridae Mörch, 1852 is an intriguing lineage of marine gastropods that are of considerable interest due to their multifaceted anatomy and complex venoms. Terebrids are abundant easily recognizable and widely distributed in tropical and subtropical waters, however the systematics of this group requires revision.. Here we elaborate the classification of Terebridae based on a recently published molecular phylogeny including 154 species, shell and anterior alimentary system characters. The 407 living species of the family, including 7 species described herein, are assigned to three subfamilies: Pellifroniinae new subfamily, Pervicaciinae Rudman, 1969 and Terebrinae Mörch, 1852. The new subfamily Pellifroniinae includes five deep-water species in two genera Pellifronia Terryn & Holford, 2008 and Bathyterebra n. gen.. Pellifroniinae possess radulae of duplex marginal teeth, well developed proboscis and venom gland, and a very small rhynchodeal introvert. The subfamily Pervicaciinae encompasses ~ 50 species in the predominantly Indo-Pacific genera Duplicaria Dall, 1908 and Partecosta Dance & Earnes, 1966. Pervicaciinae possess salivary glands, a radula of solid recurved marginal teeth and weakly developed rhynchodeal introvert, but lack proboscis and venom gland. The remaining Terebridae species are classified into 15 genera in the subfamily Terebrinae (including four genera described herein); nine genera are defined based on the phylogenetic data, and sixsolely on shell morphology. The Indo-Pacific genera Profunditerebra n. gen., Maculauger n. gen. and Myurellopsis n. gen. each include about a dozen species. The first is restricted to the deep waters of the Indo-West Pacific, and the latter two range widely in both geographic and bathymetric distribution. The Neoterebra n. gen. encompasses about 65 species from a range of localities in the eastern Pacific, Caribbean and Atlantic, and from varying depths. The highly diversified genera Terebra Bruguière, 1789, Punctoterebra Bartsch, 1923, Myurella Hinds, 1845, and Duplicaria, each comprise several morphological clusters. To characterize these genera we propose DNA based diagnoses combined with more informative descriptions in defining most of the supraspecific taxa of Terebridae to provide a comprehensive revision of the group.

INTRODUCTION

The conoidean family Terebridae, or auger snails, is a diverse lineage of predatory marine gastropods with about 400 currently accepted recent species [START_REF] Bratcher | Living terebras of the world[END_REF][START_REF] Taylor | The anatomy of the foregut and relationships in the Terebridae[END_REF]WoRMS 2018). Terebrids are specialized sand and mud dwellers. Although many terebrid species thrive in deep sea in both Pacific and Atlantic, the family reaches its highest diversity in shallow coastal waters of the Indo-West Pacific [START_REF] Miller | Studies on the biology of Indo-Pacific Terebridae[END_REF][START_REF] Castelin | Macroevolution of venom apparatus innovations in auger snails (Gastropoda; Conoidea; Terebridae)[END_REF].

The local abundance of Terebridae can be remarkable: in some habitats terebrids by far outnumber other molluscan taxa, and can be collected by hundreds per dive [START_REF] Castelin | Macroevolution of venom apparatus innovations in auger snails (Gastropoda; Conoidea; Terebridae)[END_REF].

Being locally diverse and abundant, and furthermore easily recognizable among other molluscs due to their elongate, multi-whorled shells, Terebridae is an excellent model taxon for studying patterns of microevolution [START_REF] Castelin | Macroevolution of venom apparatus innovations in auger snails (Gastropoda; Conoidea; Terebridae)[END_REF][START_REF] Holford | Evolution of the Toxoglossa Venom Apparatus as Inferred by Molecular Phylogeny of the Terebridae[END_REF]Fedosov et al. 2014), or assessing magnitudes of local biodiversity [START_REF] Castelin | Macroevolution of venom apparatus innovations in auger snails (Gastropoda; Conoidea; Terebridae)[END_REF][START_REF] Modica | A Good Compromise: Rapid and Robust Species Proxies for Inventorying Biodiversity Hotspots Using the Terebridae (Gastropoda: Conoidea)[END_REF]. Furthermore, there is a growing interest in the Terebridae family as similar to cone-snails, terebrids employ complex venoms for hunting that can be investigated to understand venom evolution, function and diversification, and to discover novel potent bioactive compounds (Puillandre & Holford, 2010;[START_REF] Imperial | The augertoxins: biochemical characterization of venom components from the toxoglossate gastropod Terebra subulata[END_REF][START_REF] Imperial | Venomous Auger Snail Hastula (Impages) hectica (Linnaeus, 1758): Molecular Phylogeny, Foregut Anatomy and Comparative Toxinology[END_REF]Kendrel et al. 2013;[START_REF] Gorson | Molecular diversity and gene evolution of the venom arsenal of Terebridae predatory marine snails[END_REF][START_REF] Renner | A return to Linnaeus's focus on diagnosis, not description: the use of DNA characters in the formal naming of species[END_REF]Verdes et. al. 2017).

The placement of the Terebridae into Conoidea is widely accepted because of the presence of a toxoglossan envenomation apparatus in most terebrids [START_REF] Taylor | The anatomy of the foregut and relationships in the Terebridae[END_REF]. Affinities with other Conoidea and relationships within Terebridae have been addressed with both, traditional morphological [START_REF] Rudman | Observations on Pervicacia tristis (Deshayes, 1859) and comparison with other toxoglossan gastropods[END_REF][START_REF] Taylor | The anatomy of the foregut and relationships in the Terebridae[END_REF]Taylor et al. 1993), and molecular phylogenetic approaches [START_REF] Holford | Evolution of the Toxoglossa Venom Apparatus as Inferred by Molecular Phylogeny of the Terebridae[END_REF][START_REF] Puillandre | The dragon tamed? A molecular phylogeny of the Conoidea (Gastropoda)[END_REF][START_REF] Castelin | Macroevolution of venom apparatus innovations in auger snails (Gastropoda; Conoidea; Terebridae)[END_REF]. A striking diversity of anterior digestive system anatomies among Terebridae, has led [START_REF] Rudman | Observations on Pervicacia tristis (Deshayes, 1859) and comparison with other toxoglossan gastropods[END_REF] to question the monophyly of the family, and establish the family Pervicaciidae mainly for species of the genus Duplicaria lacking the venom gland, and possessing an ancestral radula type. The distinctive features of Pervicaciidae were further acknowledged by [START_REF] Taylor | The anatomy of the foregut and relationships in the Terebridae[END_REF], however, its independent origin from the rest of Terebridae was rejected based on an extensive morphological dataset (Taylor et al. 1993). The monophyly of Terebridae, including Duplicaria was further confirmed by molecular studies [START_REF] Holford | Evolution of the Toxoglossa Venom Apparatus as Inferred by Molecular Phylogeny of the Terebridae[END_REF][START_REF] Castelin | Macroevolution of venom apparatus innovations in auger snails (Gastropoda; Conoidea; Terebridae)[END_REF], which recognized Duplicaria as one of the early diverging lineages of the family, together with an enigmatic deep-water species 'Terebra' jungi Lai, 2001[START_REF] Holford | Evolution of the Toxoglossa Venom Apparatus as Inferred by Molecular Phylogeny of the Terebridae[END_REF][START_REF] Castelin | Macroevolution of venom apparatus innovations in auger snails (Gastropoda; Conoidea; Terebridae)[END_REF]. T. jungi was subsequently reassigned to a separate genus Pellifronia [START_REF] Terryn | [END_REF]. However, beyond these punctuated revisions, little has been done to reconcile the systematics of Terebridae with the emerging phylogenetic framework of the family.

The present day taxonomy of the Terebridae as published in Terryn (2007) is largely based on shell characters, and no new genus or family group taxa, except Pellifronia, have been proposed since 1969. As taxonomic sampling for phylogenetic analyses of Terebridae improved, culminating in the recently published updated phylogeny of the family (Gorson et al. submitted), the inconsistency between the evolutionary relationships and systematics of the Terebridae has further increased. Given the molecular efforts, it is now clear that the genus level classification of the family needs to be revised because many of the currently recognized genera were found non-monophyletic in prior published phylogenies [START_REF] Holford | Evolution of the Toxoglossa Venom Apparatus as Inferred by Molecular Phylogeny of the Terebridae[END_REF][START_REF] Castelin | Macroevolution of venom apparatus innovations in auger snails (Gastropoda; Conoidea; Terebridae)[END_REF], Modica et al. 2014, Gorson et al. submitted). Additionally, the growing multidisciplinary interest in the Terebridae warrants a revision of the family.

Here we present a revision of the Terebridae classification that is informed by recent terebrid phylogenetic analyses. We describe a new taxonomic arrangement of Terebridae, with most genera defined based on the inferred phylogenetic relationships. As the genus level taxonomy undergoes thorough revision, placement of all known terebrid species have been revisited, and complete species lists are provided for each genus. Transition from the traditional shell-based taxonomy to the new classification, which is based largely on the phylogenetic relationships, posed some methodological issues, in particular in the proposition of informative and useful diagnoses. The observed multiple cases of convergent evolution of shell morphology prompted us to introduce molecular character based diagnoses to the recognized taxa of the Terebridae. Although use of molecular characters to diagnose supraspecific taxa remains uncommon, when the molecular data is decisive it is viewed as a necessary step in transition to new classifications as applied here with the Terebridae.

MATERIAL AND METHODS

Rationales for name attribution and ranking of recognized phylogenetic clades

As detailed in Puillandre et al. 2014, transition from the phylogenetic tree to a classification includes several steps, namely: 1) delineating putative taxa based on the inferred monophyletic lineages, morphology and phylogeography; 2) attributing names to these groups; and 3) ranking the recognized taxa. Here we followed the same general strategy, using the recently published updated phylogeny of the family Terebridae as a source of phylogenetic information (Gorson et al. submitted). Only well-supported clades in the phylogenetic analysis of (Gorson et al. submitted) are named. When molecular clades to be named are identified, a name is pinpointed to a molecular clade primarily based on the inclusion of a type species of a nominative genus in this clade (when several type species are included, priority is given to the oldest genus name). If no such species is included, the name is applied based on concordance in morphological characteristics between the species in a clade and a type species of a nominative genus. If no available names are found for a molecular clade, a new nominative taxon is proposed. Decisions on naming clades are made aiming for minimal phenotypical variability in a candidate taxon, but avoiding unnecessary fragmentation of taxa. We thus followed a conservative approach; for instance, although the names Cinguloterebra [START_REF] Oyama | On some new facts of the taxonomy of Terebridae[END_REF], Dimidacus Iredale, 1929, Myurellina Bartsch, 1923and Triplostephanus Dall, 1908, are available for four well-supported subclades of the Gorson et al. (submitted) clade C, all the lineages in the clade C are here referred to Terebra, a decision substantiated by the observed homogeneity of anatomy and radular morphology across the clade C (Gorson et al. submitted) (Fig. 1). On the contrary, six subclades of the major clade E were each assigned genus rank, leading to naming four new genera, Profunditerebra n. gen. (Clade E3), Neoterebra n. gen. (Clade E4), Maculauger n. gen. (Clade E5A) and Myurellopsis n. gen. (Clade E5B), in addition to the names Myurella applied to the clade E1 and Punctoterebra -to the clade E2. Recognition of six separate genera within this well supported clade is justified by the observed substantial differences among subclades in foregut anatomy, radular morphology and distribution.

Two well-supported lineages of the Gorson et al. (submitted), referred to as F1 and F2 therein, share the characteristic "Duplicaria" arrangement of foregut structures (Fig. 1). However, they are distinctive in shell size, morphology and distribution. Duplicaria duplicata (Linnaeus, 1958), the type species of the genus, is included in the clade F1, therefore, the name can be allocated to either the clade F1 or the entire clade F (i.e. F1+F2). As proposing genera recognizable by shell morphology undoubtedly increases the operational ability of a classification, we allocate the name Duplicaria to the clade F1 of Gorson et al. (submitted) only. The name Partecosta Dance & Eames, 1966 is available ffor the phylogenetic lineage F2, based on the notable resemblance of sequenced species in this lineage to Terebra fuscobasis E. A. Smith, 1877, the type species of Partecosta, and is so applied here. Genetic and anatomical distinctiveness of the Gorson et al. (submitted) The median estimated times of diversification in each of the recognized genera range from 14 million years ago (Mya) (Maculauger) to 29 Mya (Terebra), which suggests that diversification of Fig. 1 terebrid genera has taken place in Oligocene-Miocene (Gorson et al. submitted). These estimates fall within the range estimated for the most recent common ancestors in other genera of neogastropods in the muricid subfamily Rapaninae [START_REF] Claremont | Global phylogeny and new classification of the Rapaninae (Gastropoda: Muricidae), dominant molluscan predators on tropical rocky seashores[END_REF]) and in the family Conidae [START_REF] Duda | Species-level phylogeography and evolutionary history of the hyperdiverse marine gastropod genus Conus[END_REF]Puillandre et al. 2014).

Allocation of species to genus level groups

In identifying the 1400 sequenced specimens analysed in Gorson et al (2018), we found that the species boundaries as predicted by shell morphology in many cases corresponds to complexes of closely related species (Gorson et al. submitted). In most cases, revealed molecular operational taxonomic units (MOTUs) were distinguishable by shell morphology, and in several cases we demonstrate that species previously considered as synonyms actually correspond to distinct species.

We thus raised to species level those that we can confidently identify. Phylogenetic analysis of Gorson et al. (submitted) also identified several MOTUs that undoubtedly correspond to undescribed species. Although massive description of new species is not the goal of the present study, those new species that are deemed important for understanding boundaries and identities of the newly introduced genera, are described herein. In only one case were specimens of a currently accepted species found to be junior synonyms of another species. Despite many striking parallelisms in shell morphology uncovered by the phylogenetic analysis (see Discussion), we found that in general species with similar morphology predictably grouped together in distinct clusters within a given clade. This confirmed that shell morphology is an overall reliable proxy that could be used to allocate species for which no sequence data is available for the proposed taxa (further discussed below).

Morphological and molecular circumscription of taxa

For the morphological circumscription of taxa we adopted a three-step approach. In a first "indicative" step, we identified species and groups of species based on their distinctive features in both shell morphology and anatomy within each genus-level clade identified in the phylogenetic tree (Gorson et al. submitted). The International Commission on Zoological Nomenclature (ICZN) allows two alternative options for establishing morphological identity of a taxon: (1) to propose a diagnosis -i.e. 'a summary of the characters that differentiate the new nominal taxon from related or similar taxa' (ICZN Recommendation 13A), or (2) to propose a description, in which taxonomically informative and uninformative characters are not separated [START_REF] Renner | A return to Linnaeus's focus on diagnosis, not description: the use of DNA characters in the formal naming of species[END_REF]. In a second "diagnostic" step, we were able to use the defined features to add other non-sequenced species to the proposed taxa and to formulate a synopsis of morphological characteristics for each generic taxon. As multiple cases of convergent evolution of shell features were demonstrated for Terebridae (Gorson et al. submitted), the synopsis content was usually not sufficient for unequivocal identification of a primarily phylogenetically defined genus. In such cases, a formal Description was provided, and additional information on how the members of each genus can be differentiated from similar but unrelated forms is to be found in our Remarks sections. In several genera, several distinctive subclades were recognized by the phylogenetic analysis, and they could have been considered as subgenera. Here we chose to not define subgenera to avoid introduction of multiple new names for lineages whose morphological identities are not entirely understood at present. However, we retained in the remarks, brief descriptions of the potential subgroups for reasons of clarity and usability. In a third step, we investigated the species that were difficult to allocate with confidence to the phylogenetically defined genera. We found that in many cases these species formed groups, which, when diagnosed for distinctive morphological features, were found to be attributable to currently accepted genera for which no DNA sequence data is available. As no phylogenetic data is available on these genera, we cannot resolve their status with certainty, and thus such genera are retained as valid. In the systematic section below, for each genus we propose a formal diagnosis or description, depending on our ability to identify the characteristics that differentiate a proposed genus from other groups in Terebridae and provide a list of member species.

We compiled molecular diagnoses to supplement traditional taxonomic diagnoses and descriptions in response to the fact that recognition of primarily phylogenetically defined genera by morphological means was often problematic. As the four-gene (COI, 18S, 12S, and 23S) phylogenetic analysis (Gorson et. al. submitted) was not comprehensive for all species and specimens per clade, the credibility of a molecular diagnosis derived from this dataset would have been questioned. To combat this we used only the COI dataset including only sequences with no more than 5 ambiguously called nucleotides (963 records), representing 179 putative species.

Initially, species were attributed to pre-defined clades based on the results of the four-gene analyses of Gorson et. al. (2018) and that was consistent with allocations to genera and subfamilies in a classification detailed below. The total length of the COI alignment was 658 base pairs (the "barcode" fragment defined by the Folmer et al (1994) primers), which were thus numbered from 1 to 658. First, an attempt was made to find pure diagnostic single-nucleotide positions (i.e. at a given position, all the members of the clade and none of the non-members of the clade share a given nucleotide) using the R package SPIDER 1.4-1 [START_REF] Brown | Spider: An R package for the analysis of species identity and evolution, with particular reference to DNA barcoding[END_REF]. However, only one diagnostic single nucleotide character was identified for each of the clades A1 and E5A, and none in the others. Thus we explored the possibility of providing molecular diagnoses based on diagnostic combinations of nucleotides (i.e. at two or more positions, all the members of the clade and none of the non-members of the clade share a combination of nucleotides). To our knowledge, only CAOS [START_REF] Sarkar | CAOS software for use in characterbased DNA barcoding[END_REF]) is capable of detecting such diagnostic combinations. However, as CAOS implements a tree-based algorithm, it poses certain requirements to the data to run efficiently, such as a pre-defined group is only provided diagnosis (referred to as characteristic attributes or CA's by [START_REF] Sarkar | CAOS software for use in characterbased DNA barcoding[END_REF][START_REF] Rach | Character-based DNA barcoding allows discrimination of genera, species and populations in Odonata[END_REF] if it corresponds to a clade in the provided tree. The tree reconstructed using all our COI sequences (Gorson et al. submitted) did not retrieve all the subfamilies and genera monophyletic, thus making CAOS not applicable. Consequently, we developed a custom Python script (available upon request) that is based on an alternative, purely statistical approach, and implements iterated random search to identify diagnostic combinations of nucleotides for each predefined group in an assemblage of aligned sequences. The in-house Python script provided the most comprehensive and reproducible results for our dataset.

RESULTS

Levels of confidence in attribution to genera

Valid recent species entries (401) from current WoRMS database (as per February 1 2018) were used in our analysis. DNA sequence data were available for 135 currently accepted species (Gorson et al. submitted) that constitute 134 species entries (except Oxymeris troendlei (Bratcher, 1981), which was found to be a junior synonym of O. maculata (Gmelin, 1791)).

These 134 species as well as seven new species revealed by the phylogenetic analysis of Gorson and coauthors (2018) and described herein are allocated to genera with a superscript 1 ( 1 ), indicating highest degree of confidence.

The use of morphological characters in combination with geographic distribution resulted in a non-ambiguous placement into one of the groups recognized phylogenetically of 166 more species; these are marked with superscript 2 ( 2 ).

In multiple cases, shell morphology initially appeared to be inconclusive for confident attribution of a species to a genus, as more than one hypothesis on its placement was proposed; in such cases we suggested a tentative allocation taking into account all available morphological and distribution data. This is the case for 59 species entries marked with superscript 3 ( 3 ).

Of the remaining species, 39 showed no clearly definable affinity to any single molecularly defined genus. However, based on the shell morphology alone, each species could be allocated to one of six terebrid genera Granuliterebra [START_REF] Oyama | On some new facts of the taxonomy of Terebridae[END_REF], Hastulopsis Oyama, 1961, Pristiterebra Oyama, 1961, Gradaterebra Cotton & Godfrey, 1932, Perirhoe Dall, 1908, Microtrypetes Pilsbry & Lowe, 1932. These six genera remain solely defined by shell characters pending further molecular data.

Six species showed no definite resemblance to any terebrid genus, these are here treated as incertae sedis and more data are needed to suggest credible genus allocations.

Identification of COI DNA based diagnoses

Numerous diagnostic combinations were returned for all but three genera, and were sorted by length with priority given to short combinations. The diagnostic nucleotides identified for the clades A1 and E5A by SPIDER were also recovered by our in-house Python script. However, we failed to identify diagnostic combinations of nucleotides for the genera Terebra and Punctoterebra, and only found diagnostic combinations for Hastula H. Adams & A. Adams, 1853, when four sequences with a substitution at the position 181 of the COI fragment were removed from the dataset. As a result, for Terebra, Punctoterebra, and Hastula taxa we propose separate diagnoses for the inferred phylogenetic clusters within each genus.

SYSTEMATICS

Superfamily Conoidea Fleming, 1822

Family Terebridae Mörch 1852 [Synonyms: Acusidae Gray, 1853] Description: Shell elongate or conical, multi-whorled, with very high spire and relatively small last whorl. Aperture small, with anterior siphonal notch, or very short straight siphonal canal. Anal sinus indistinct.

Anatomy. Operculum always present, horny, rounded or leaf-shaped, with terminal nucleus. Head with two very short closely set eye tentacles with eye situated near their tips. Rhynchodaeal introvert always present, small to very large. Proboscis, radula, salivary gland, accessory salivary gland, venom gland and accessory proboscis structure present or absent. Radula, when present, composed of marginal teeth onlyvarying in shape from flat, solid and recurved, semi-enrolled or duplex to hollow hypodermic. Subfamily Pellifroniinae n. subfam.

(Fig. 2) Type genus: Pellifronia [START_REF] Terryn | [END_REF] Zoobank registration: urn:lsid:zoobank.org:act:E827742C-E1B3-42DD-B5EB-5D7EAA6E2C95

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 1.

Description: Shell small to medium-sized (12-50 mm), with predominant axial sculpture. Early teleoconch whorls often with angulated profile, later -flattened or even slightly concave to convex. Sculpture of numerous closely set ribs, even, or forming raised nodules bordering upper or lower suture. Spiral sculpture absent or of very shallow and wide subsutural depression.

Anatomy. Rhynchodeal introvert small, proboscis, radular sac, odontophore, salivary and venom glands well developed. Radula with well-developed membrane, bearing two rows of wide marinal teeth of duplex type. Accessory proboscis structure absent.

Remarks: The subfamily comprises five described species in two genera, Bathyterebra and Pellifronia that occur in Indo-Pacific and Caribbean at bathyal depths. Anatomy of the studied species of the subfamily demonstrates characters that are believed to be ancestral for Terebridae in general -i.e. weakly developed rhynchodeal introvert and radular teeth of duplex type [START_REF] Castelin | Macroevolution of venom apparatus innovations in auger snails (Gastropoda; Conoidea; Terebridae)[END_REF]. Definition This genus includes species closely related to Pellifronia jungi (Lai, 2001) based on the molecular phylogenetic analysis (Gorson et al. submitted) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable to Pellifronia jungi or any genetically proven member of the genus.

Genus

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 1.

Description: Shell medium-sized (25-50 mm), elongate, light brown. Protoconch brown, paucispiral, cyrthoconoid, of 1.5-2.5 whorls. Coarse sculpture of pronounced ribs, orthocline or prosocline, sometimes arcuate on late teleoconch whorls. Axial ribs often thickened in their adapical and/or abapical part(s), giving slightly concave outline to spire whorls. Last adult whorl with flattened adapical portion, and short somewhat angulate transition to shell base. Siphonal canal moderately long, recurved at its tip, shallowly notched; siphonal fasciole weak or indistinct. Aperture elongate, curved. Fig. 2 Table 1 Anatomy. Rhynchodeal introvert small, proboscis, radular sac, odontophore, salivary and venom glands well developed. Acessory proboscis structure absent. Radula with well-developed membrane, bearing two rows of wide marginal teeth of duplex type.

Distribution: Indo-Pacific, bathyal depths Remarks: The genus differs from the closely related Bathyterebra in having stronger axial sculpture, more flattened whorl outline and last adult whorl constricted to siphonal canal with visible angulation. Species of Pellifronia can be distinguished by their coarse sculpture, swollen ribs ends with slightly raised subsutural area. Similar species in other genera, e.g. Neoterebra puncturosa (S.S. Berry, 1959), N. crenifera (Deshayes, 1859) and Granuliterebra bathyrhaphe (E.A. Smith, 1875), normally have a sharper nodular structure on the subsutural band or ribs' endings, or can be separated by the lack of radula (species of Neoterebra).

Included species:

Pellifronia brianhayesi (Terryn & Sprague, 2008) Definition: This genus includes species closely related to Bathyterebra benthalis (Dall, 1889) based on the molecular phylogenetic analysis (Gorson et al. submitted), and those that show a combination of conchological, anatomical and distribution characteristics closely comparable to Bathyterebra benthalis or any genetically proven member of the genus.

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 1.

Description: Shell small, of 12-30 mm, elongate, white to tan. Protoconch paucispiral, cyrthoconoid, of about 1.5 glossy whorls. Sculpture of fine dense orthocline ribs, interrupted by depression in adapical portion of whorl. Axial ribs weakening throughout their height on mature whorls, often forming weak nodules bordering the adapical depression, giving distinctly angulated outline to early spire whorls; late teleoconch whorls gently convex. Last adult whorl evenly convex, transiting to shell base without visible angulation. Siphonal canal stout, slightly recurved at its tip, shallowly notched; siphonal fasciole weak, or rather strong. Aperture elongate, rather wide.

Anatomy. Rhynchodeal introvert small, proboscis, radular sac, salivary and venom glands well developed. Acessory proboscis structure absent. Radula with well-developed membrane, bearing two rows of wide marinal teeth of duplex type.

Distribution: South China Sea, New Caledonia, Caribbean, bathyal depths Etymology: The genus name refers to the bathyal depths, at which three known species of the genus were sampled. Gender feminine.

Remarks: Bathyterebra spp. can be differentiated from most other genera of Terebridae by the combination of relatively faint sculpture lacking spiral elements, with weakly defined subsutural band and relatively high last adult whorl with convex outline. In shell proportions the genus somehow resembles Oxymeris Dall, 1903; however, the latter genus has more flattened whorls and typically can be easily differentiated by a notably larger size at maturity.

Included species:

Bathyterebra benthalis (Dall, 1889) 1 comb. nov.;

B. coriolisi (Aubry, 1999) (Fig. 3) Type genus: Pervicacia Iredale, 1924(=Duplicaria Dall, 1908) Description: Shells elongate ranging from 10 to 100 mm, with high spire, predominant axial sculpture and wide aperture.

Anatomy. Rhynchodeal introvert rather small, salivary gland and radular sac present, radula of solid curved marginal teeth attached to strong membrane. Venom gland, proboscis and accessory proboscis structure absent. [START_REF] Dall | Subdivisions of the Terebridae[END_REF] (Fig. 3 A -E) [Synonyms: Diplomeriza Dall, 1919(Unnecessary nom. nov. pro Duplicaria Dall, 1908, by Dall treated as a homonym of the non-existent "Duplicaria Rafinesque, 1833"); type species: Buccinum duplicatum Linnaeus, 1758; by typification of replaced name; Myurellisca Bartsch 1923; type species: Myurella duplicatoides Bartsch, 1923;OD;Pervicacia Iredale, 1924; type species: Terebra ustulata Deshayes, 1857; OD] Type species: Buccinum duplicatum Linnaeus, 1758; OD.

Genus Duplicaria

Definition:

The genus is primarily defined based on the results of the phylogenetic analysis (Gorson et al. submitted), and comprises species which have been demonstrated to form a well-supported clade with the type species Duplicaria duplicata (Linnaeus, 1758), and additionally species which show shell morphology, anatomical or genetic characteristics closely comparable to Duplicaria duplicata or any genetically proven member of the genus.

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 2. Table 2 13 Description: Shell elongate, small to large-sized, adult size 20-100mm, with wide aperture and predominant axial sculpture, either continuous from suture to suture or interrupted by a shallow to deep subsutural indentation. Spiral structure obsolete, sometimes, microscopic striae or weak band joining nodes.

Anatomy. Rhynchodeal introvert rather small, salivary gland and radular sac present, radula of solid curved marginal teeth attached to strong membrane. Venom gland proboscis and accessory proboscis structure absent.

Distribution: Tropical and subtropical Indo-Pacific and West Africa.

Remarks: By shell morphology 3 distinct clusters can be delineated within Duplicaria: Duplicaria duplicata group. Small to large shells, adult size 25-100 mm, with paucispiral protoconch and sculpture of strong axial ribs. Deep narrow spiral groove delineating distinct flattened subsutural band and no spiral structure. Whorl outline flattened, or shouldered due to elevated ribs below the subsutural band.

Duplicaria tristis (Deshayes, 1859) group. Small shells, adult size 10-30 mm, paucispiral protoconch with sculpture of strong axial ribs, subsutural area weakly impressed.

Duplicaria tricincta (E.A. Smith, 1877) group. Small shells, adult size 10-25 mm, paucispiral protoconch. Strong spiral ribs. A more convex shape with outline flattened due to elevation of ribs below the subsutural area. Ribs weakening on mature whorls.

Included species:

Duplicaria albozonata (E. A. Smith, 1875) (Aubry, 1999) Definition: This genus includes species that show a combination of conchological, anatomical and distribution characteristics closely comparable to Strioterebrum wilkinsi Dance & Eames, 1966, including all species in the clade F2 of the (Gorson et al. submitted), or any species proven phylogenetically close to them.

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 2.

Description: Shell small, adult size 10-30 mm, pale or tan, often with spiral bands and dark basal area. Protoconch from 1.5 to 3 whorls. Dominant sculpture of axial ribs spanning across whorl's height, resulting in even, slightly convex whorl outline, or (typically) interrupted by depression or narrow groove forming subsutural row of nodules. Spiral sculpture indistinct or represented by fine regular striae. Siphonal canal very short, stout, indistinctly or very shallowly notched, with strong fasciole. Aperture wide, pyriform.

Anatomy. Rhynchodeal introvert rather small; salivary gland and radular sac present, radula of solid curved marginal teeth attached to strong membrane. Venom gland proboscis and accessory proboscis structure absent.

Distribution: Indian Ocean, intertidal and shallow subtidal depths.

Remarks: Three clusters with varying sculpture patterns can be recognized:

Partecosta fuscobasis group with sculpture of strong ribs and clearly delineated subsutural band forming distinct nodules at intersections with ribs (Figs 3F -I). It is closely comparable to species of the New World genus Neoterebra, however, can be differentiated by smaller size at maturity, wider aperture, usually with wide anterior notch, and solid curved radular teeth.

Partecosta trilineata (Bozzetti, 2008) group with glossy shells sculptured by strong continuous ribs, no distinct spiral sculpture or subsutural band (Figs 3K,L). Very similar sympatric forms are known in the genus Punctoterebra (i.e. P. solangeae Bozzetti, 2015 group), that can only be reliably differentiated by anatomical characters (radula of solid curved teeth in Partecosta spp. and lacking in P. solangeae).

Partecosta macleani (Bratcher, 1988) group with flattened whorl outline, very short and robust siphonal canal, and very weak sculpture represented by rounded, almost indistinct subsutural nodules extended into even less distinct axial folds on whorl periphery (Fig 3M). Some species of Hastula are close to Partecosta macleani, however, they are commonly larger, have polished surface and retain distinct ribbing at least on the adapical portions of the whorls.

Included species:

Partecosta albofuscata (Bozzetti, 2008) (Aubry, 1991) 3 comb. nov. 517

Subfamily Terebrinae Mörch 1852

Type genus: Terebra Bruguière, 1789 Description: Shell elongate, multi-whorled, with varying whorl outline, sculpture and colour pattern. Aperture very small relative to shell height, moderately wide to very wide, with short, straight or very weakly recurved, widely open siphonal canal.

Anatomy. Rhynchodeal introvert large, occupying most of the rhynchocoel cavity. Radula, proboscis, venom gland, salivary gland and accessory proboscis structure present or absent. Radula when presentconsisting of hypodermic, flat, semi-enrolled or very rarely -duplex marginal teeth, never of solid type.

Remarks: Although the subfamily is consistent with a highly supported clade in the phylogenetic tree of Gorson et al. (submitted), its conceptual circumscription (i.e. based on the shared synapomorphic characters) is difficult. Among such characters only the large rhynchodeal introvert that occupies most of the rhynchocoel cavity can be mentioned. Foregut anatomy offers some other characters that can practically facilitate recognition of Terebrinae. Until proven otherwise, all the radula-less terebrids are in the Terebrinae, as well, as all species with flat, semi-enrolled or hypodermic radular teeth. Practically, recognition of the terebrines can be aided by the fact that the two other terebrid subfamilies are notably more compact and are generally clearly delineated; therefore, everything that is definitively not a member of neither Pellifroniinae nor Pervicaciinae is here considered in Terebrinae. Definition: The genus is primarily defined based on results of the phylogenetic analysis (Gorson et al. submitted), and includes species clustering into a well-supported clade of Terebra subulata (Linnaeus, 1767), and species, showing resemblance in shell morphology, anatomy and distribution to any genetically proven member of the genus.

Genus

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided separately for the seven clades of the genus inferred by Gorson et al. (submitted) in the Table 3. Table 3 Description: Shell elongate, medium-sized to large, reaching 192 mm, with truncated base shortly constricted to siphonal canal, sometimes with distinct angulation. Sculpture from very weak, limited to often obsolete subsutural groove, to very strong. Subsutural band simple, or consisting of two closely set cords, often gemmate. Lower whorl portion smooth or with varying sculpture pattern, with spiral elements being stronger or at least pronounced equally with axials. Shell base separated from siphonal canal by concave waist. Aperture elongate to quadrangular.

Anatomy. Rhynchodeal introvert, proboscis, venom gland, salivary gland and radula well-developed. Radula consisting of hypodermic marginal teeth, with only the bases attached to strongly reduced membrane. Accessory proboscis structure present or absent.

Distribution: Indo Pacific, West Africa, Panamic, Caribbean and Western Atlantic Remarks: The genus Terebra is here established in the boundaries consistent with the clade C of Gorson et al. (submitted). This clade is in turn subdivided into eight lineages, referred to as C1 -C7 (see figure 4), their relationships being not resolved. The clades C1, C2, C3-2, C4, and C7 are well supported; clade C5 comprising one species only -Terebra argus. Because of the extreme heterogeneity of the clade C our attempts to provide a molecular diagnosis failed, and therefore, we compiled separate diagnoses for the inferred lineages within C (Table 3). Although lineages C3-1 and C3-2 do not constitute a clade, their affinity is not rejected by the phylogenetic analysis, and here they are grouped together based on the biogeographic considerations. The genus Terebra is enormously heterogeneous in regard of shell morphology as well (Fig. 5); some common morphotypes can be delineated based on shell morphology, but their correspondence with inferred phylogenetic groupings is far from being perfect. Below we provide brief overview of these morphotypes: Typical Terebra are characterized by a weakly sculptured or smooth medium sized shell, with gently convex or subcylindrical whorls, often bearing pattern of large spots. Species referable to this morphotype are included in clades C1 (T. subulata and T. guttata (Röding, 1798)), C4 (T. ornata Gray, 1834) and C6 (T. argus Hinds, 1844). This morphotype is probably closest to Oxymeris, and differentiation of the two lineages is tricky. In most cases Terebra spp. can be recognized by the narrower anterior notch; however, examination of the foregut to determine whether proboscis and radula are present is advised for definite allocation.

The fenestrata morphotype comprises most heavily sculptured species, which are distinctive because of strong, often elevated gemmate subsutural band which is closely followed by second spiral band of similar morphology, but usually weaker and/or narrower. The lower portion of the whorl is sculptured with spiral cords, or have cancellate pattern. Species exhibiting this morphotype comprise the clade C2, and strikingly similar species can be also found in the clades C7 (T. aff. adamsii (E. A. Smith, 1873), T. aff. rubrobrunnea Preston, 1908 MNHN IM-2013-14908, Terebra anilis (Röding, 1798)) and C8 (Terebra cumingi Deshayes, 1857 species complex). Strong, often elevated, gemmate or beaded spiral elements readily differentiate this morphotype from most terebrid genera. The similar pattern however is present in Granuliterebra spp., the species of which being differentiated by the single subsutural band and usually by a prevailing axial sculpture on the lower portion of the whorl.

Triplostephanus morphotype unifies species with very high spire and very small, quadrate aperture. Whorls flattened or slightly concave due to raised subsutural band, and lower part of whorl sculptured by regular cords, while axial sculpture is not developed. The clade C5 is mostly represented by species exhibiting this morphotype; however, species with sub-cylindrical whorls and stepped outline of spire (such as Terebra laevigata Gray, 1834 and Terebra funiculata Hinds, 1844) 6 Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided for the entire clade with the exception of four specimens of H. albula (see Materials and Methods) and separately, for the subclade H. albula -H. natalensis -H. aff. casta of the clade D (Table 4).

Description: Shell small to medium-sized, shiny, adult size (12-90 mm). Protoconch pauci-or multispiral of about 1 to 5 whorls. Axial sculpture represented by crenulations on whorl's subsutural portion, or by fine ribs on part or throughout the whorls' height; spiral sculpture represented by single subsutural groove, seldom with additional row of punctations. Siphonal canal stout, very short, with well-developed fasciole. Aperture elongate to rounded.

Anatomy. Rhynchodeal introvert, proboscis, venom gland, salivary gland and radula well-developed. Radula consisting of hypodermic marginal teeth attached to thin, reduced membrane. Walls of marginal teeth often with multiple perforations (Figs 6 E,F) Distribution: Throughout Indo-Pacific, West Africa, Caribbean.

Remarks: Species of Hastula are rather easy to recognize amongst Terebridae, by their typically shiny shells, with fine closely set of axial ribs and usually by the lack of clearly defined subsutural band. The shell morphology in the Partecosta macleani group closely resembles species of Hastula; however, the latter genus is usually notably larger at maturity, and shells of comparable size can be differentiated by texture -polished surface in Hastula versus dull in Partecosta. Furthermore, species of Hastula possess venom gland and hypodermic radular teeth, whereas Partecosta spp. lack venom gland and their radulae consist of solid teeth.

Included species:

Hastula aciculina (Lamarck, 1822) 2 ; 769 H. acumen (Deshayes, 1859) 1 ; 770 alboflava Bratcher, 1988 Definition: The genus includes species closely related to Oxymeris maculata (Linneaus, 1758), based on the molecular phylogenetic analysis (Gorson et al. submitted) and those that show a combination of conchological, anatomical and distribution features closely comparable to Oxymeris maculata (Linnaeus, 1758) or any genetically proven member of the genus.

H.
Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 5.

Description: Shell medium-sized to large, adult size reaching 273 mm, weakly sculptured, typically with orthoconoid spire and rather wide aperture. Protoconch multispiral of 3 -3.75 whorls. Whorls flattened in outline. Sculpture of subsutural band, smooth or with regular crenulations; juvenile sculpture often consisting of axial ribs. Last adult whorl inflated; siphonal canal short and stout. Aperture wide, elliptic to rounded.

Anatomy. Rhynchodeal introvert very large, proboscis absent or vestigial, salivary gland absent or very small, radula venom gland and accessory proboscis structure absent.

Distribution: Indo-Pacific, tropical East Pacific, West Africa.

Remarks: Species of Oxymeris are among the largest terebrids, and can usually be easily identified by their large, shiny shells, with an orthoconoid or even slightly acuminate spire, often with a bulbous last whorl, a very wide aperture and a very short siphonal canal, separated from the shell base by a deep groove. Nevertheless, some smaller species (i.e. O. cerithina) have spire with convex sides, and superficially resemble medium-sized Punctoterebra and Myurella. However, in most cases (except O. crenulata) the Oxymeris species can be recognized by the complete lack of axial sculpture other than growth lines, while axial elements of varying strength and shape are present in Punctoterebra, Myurella Definition: The genus is primarily defined based on results of the phylogenetic analysis (Gorson et al. submitted), and comprises species which have been demonstrated to form a well-supported clade with the type species Myurella affinis (Gray, 1834), and additionally species which show shell morphology, anatomical or genetic characteristics closely comparable to Myurella affinis or any genetically proven member of the genus.

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 6.

Description: Shell very small to medium-sized, with elongated, with high orthoconoid spire and sculpture of arcuate or undulating ribs fine and narrow to flat and wide.

Anatomy. Rhynchodeal introvert very large, proboscis, radula and venom gland absent, salivary gland and accessory proboscis structure present in few species.

Distribution: Indo-Pacific, at intertidal to upper bathyal depths.

Remarks:

In regard of shell morphology, four distinct clusters can be recognized: Fig. 7 Table 6 22 Myurella affinis group. Shell medium-sized; axial sculpture of low and wide, flattened ribs, subsutural band with elongate nodules delineated by punctations, which are also pronounced in interspaces between axial ribs. Similar species that earlier were classified to Myurella, but that were found not immediately related to T. affinis (i. e. M. undulata, M. columellaris and M. kilburni) are here reclassified to the genus Myurellopsis. The typical Myurella can be recognized by the lower axials and shallower interstices between them. Because of the overall flatter axials, the subsutural band is visually continuous in Myurella; on the contrary, due to the more elevated ribs in Myurellopsis spp., the subsutural band is represented by a row of nodules.

Myurella pseudofortunei group. Shell elongate of variable size (17-60 mm), with very high and slender spire, evenly convex whorls, indistinct subsutural band and narrow, clearly arcuate axial ribs; interspaces with regular striae. Species of this group exhibit a very distinctive morphotype, which has little in common with other species here attributed to Myurella. Terebrids with this morphology were previously placed in the genus Clathroterebra Oyama, 1961, of which some smaller species were found to be unrelated lineages, the differences between which are still to be elaborated.

Myurella pygmaea group. Shell small, circa 10 mm, with curving outline, straight sides to whorls and very short, tapering siphonal canal. Axial sculpture of numerous fine oblique ribs; subsutural band, or other spiral elements absent. The group includes one species, the only representative of the former genus Terenolla. The very small but rather solid, finely ribbed shell, of Myurella pygmaea is easily recognizable amongst terebrids; its recognition is further aided by the color pattern with flesh background and a contrasting dark-brown spiral line just below the suture.

Myurella amoena group. Shell medium-sized, sculptured with fine to strong ribs, subsutural band defined by line of punctuations, spiral structure of narrow grooves or cords. The species of this group form a wellsupported clade and exhibit a shallow water morphotype that was formerly classified within Hastulopsis [START_REF] Oyama | On some new facts of the taxonomy of Terebridae[END_REF]. Similar deeper water species are found in the genus Maculauger n. gen., most of which can be recognized by having a row of large spots on the periphery. The other similar species remain within the genus Hastulopsis, comprising a number of Indian Ocean and Japanese endemic species. Species of Myurella amoena group have strong but narrow ribs that are, however, notably stronger than spiral elements. In Myurellopsis spp. the ribs are about as strong, but are wider and rounded, with interstices being narrower than ribs themselves. Both Hastulopsis and Maculauger comprise species with axial and spiral elements of comparable strength, resulting in a rather cancellate pattern, and a continuous subsutural band.

Genus Punctoterebra Bartsch, 1923 (Fig. 8) Type species: Terebra nitida Hinds, 1844; OD.

Definition:

The genus is primarily defined based on results of the phylogenetic analysis (Gorson et al. submitted), and comprises species which have been demonstrated to form a well-supported clade with Punctoterebra nitida (Hinds, 1844), and additionally species which show shell morphology, anatomical or genetic characteristics closely comparable to Punctoterebra nitida or any genetically proven member of the genus.

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided separately for the four phylogenetic clusters of the Punctoterebra inferred by Gorson et al. (submitted) in the Table 7 (the deep water lineage composed by two currently undescribed species represented by MNHN IM-2013-52264 and MNHN IM-2013-52293 treated separately from the Punctoterebra textilis group).

Description: Shell elongate, small to medium sized (adult size below 50 mm), with high spire and prevailing axial sculpture. Protoconch paucispiral or multispiral, of varying morphology. Axial ribs strong on spire whorls, sometimes becoming obsolete on last whorl. Spiral sculpture limited to a weak punctuate groove delineating subsutural band, or well-developed represented by regular striae in interspaces between axial ribs.Siphonal canal delineated from shell base by distinct concavity, often slightly recurved.

Anatomy. Proboscis, venom gland, radula and salivary gland typically present, radula, when present, composed of flat marginal teeth. Accessory proboscis structure absent.

Distribution: Throughout Indo-Pacific, intertidal to upper bathyal depths (to 405 m), typically shallow.

Remarks: With regard to shell morphology, species of the genus form three distinct clusters that are consistent with three phylogenetic lineages inferred by (Gorson et al. submitted).

Punctoterebra nitida group. Shell small to medium-sized (<50mm), polished in appearance, with subsutural band delineated by deep punctures or short grooves; sculpture predominantly represented by axial ribs. The small species of the group (i.e. P. plumbea or P. solangeae) are conchologically very close to some species of Partecosta (see Remarks section under latter genus).

Punctoterebra teramachii (Burch, 1965) group. Shells medium sized (<45mm) with dominant structure of orthocline axial ribs; spiral structure absent. Subsutural band clearly defined. Species of this group are conchologically close to the genus Duplicaria, from which they can be differentiated by the punctate subsutural groove and narrower aperture, and by morphology of radular teeth that are solid and recurved in Duplicaria, and flat (or absent) in Punctoterebra spp.

Punctoterebra textilis (Hinds, 1844) group. Shell medium sized (<50mm), heavily sculptured, with gently convex whorls. Sculpture of arcuate ribs, interspaces bearing regular distinct striae; subsutural band delineated by a punctuate groove of varying strength. Syphonal canal typically recurved, delineated from shell base by distinct waist of deep groove. This group is rather heterogeneous in shell morphology, with the core formed by heavily sculptured species of the Punctoterebra textilis -P. succincta (Gmelin, 1891) complex. Similar heavily sculptured shells characterize typical Profunditerebra; however, these can be distinguished by more rounded ribs, a row of nodules on the subsutural band, and by straight siphonal Fig. 8 Table 7 Genus Profunditerebra n. gen.

(Fig. 9 A -E) Type species: Profunditerebra papuaprofundi n. sp.; OD herein.

Zoobank registration: urn:lsid:zoobank.org:act:A5A78A3D-AB11-41C3-86F0-7C1451512FF4 Definition: This genus includes species closely related to Profunditerebra papuaprofundi n. sp., based on the molecular phylogenetic analysis (Gorson et al. submitted) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable to Profunditerebra papuaprofundi n. sp. or any genetically proven member of the genus.

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 8.

Description: Shell small to medium-sized (<45 mm), with multispiral protoconch and slender siphonal canal. Whorl subcylindrical or flattened. Sculpture pattern varying greatly with axial sculpture of rounded ribs, weak to very strong and elevated. Spiral sculpture typically limited to striae in interstices and a subsutural groove, absent or represented by strong cords generating overall cancellate pattern.

Anatomy. Proboscis, radula, venom gland and salivary gland present. Radula of duplex marginal teeth (Fig. 9E). Accessory proboscis structure absent. Distribution: Almost exclusively in deep water (typically 150-600 m) of tropical Indo-West Pacific from Taiwan to New Caledonia; one species is known from shallow water of South Australia.

Remarks: Profunditerebra is the only terebrine genus with duplex radular teeth similar to those in Pellifroniinae; members of the two groups can be easily differentiated by shell characters. Species of Profunditerebra can be separated into four clusters based on their shell morphology; however, only the first cluster corresponds to a monophyletic lineage.

Profunditerebra papuaprofundi n. sp. et gen. group. Shell uniformly coloured white to dark brown, heavily sculptured, with distinct subsutural groove and subcylindrical whorls. Sculpture of strong rounded ribs forming a row of prominent nodules on subsutural band. Spiral sculpture of fine striae limited to interstices between axials, or overriding them to form fine continuous cords (including on subsutural band), or coarse cancellate pattern. Species in Neoterebra (i.e. N. armillata) exhibit overall similar sculpture pattern, but can be differentiated from Profunditerebra by their flattened whorls. The Punctoterebra textilis group also includes some similar forms (see Remarks section under Punctoterebra).

Profunditerebra orientalis (Aubry, 1999) group. Shells with flattened whorl profile, and variegated colour pattern. Sculpture of fine orthocline ribs intersected by cords of about equal strength or slightly weaker than axials to form delicate cancellate pattern. Subsutural band delineated by shallow punctuate groove. Species of this group can be easily differentiated from congeners that either lack spiral elements, or display an overall much coarser sculpture. Both Profunditerebra orientalis and Profunditerebra hiscocki closely resemble species in the genus Maculauger from which they can be confidently differentiated only by anatomy (all studied species of Maculauger are radula-less) or molecular characters.

Profunditerebra anseeuwi (Terryn, 2005) group. Shells with flattened whorl outline, subsutural band represented by short ribs or indentation. Sculpture of axial ribs pronounced throughout whorl's height, but weakening and becoming obsolete on later whorls; spiral sculpture absent. Species of this group are conchologically close to Duplicaria and to the Punctoterebra teramachii group; however, they can be differentiated from both by the overall weaker sculpture. Besides, these lineages can be confidently differentiated by the radular teeth morphology, with Duplicaria having solid and recurved teeth, Profunditerebra -duplex teeth, and Punctoterebra -flat teeth.

Profunditerebra brazieri (Angas, 1871) stands apart from other species in the clade, being an endemic of subtropical shallow waters off Tasmania. Morphologically, it is close to the Profunditerebra anseeuwi group, but has a very weakly defined subsutural band.

Etymology:

The genus name refers to the considerable depths, at which all known species of the genus occur. Gender feminine.

Included species:

Profunditerebra anseeuwi (Terryn, 2005) (Fig. 9 F -H) Type species: Terebra specillata Hinds, 1844; OD herein.

Zoobank registration: urn:lsid:zoobank.org:act:68993F36-2AD1-4DDE-A853-8832B4A9EE30 Definition: The genus is primarily defined based on results of the phylogenetic analysis (Gorson et al. submitted), and comprises species which have been demonstrated to form a well-supported clade with Neoterebra specillata (Hinds, 1844), and additionally species which show shell morphology, anatomical or genetic characteristics closely comparable to Neoterebra specillata or any genetically proven member of the genus.

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 9.

Description: Shell small to medium-sized (<60 mm) uniformly creamy-white to light brown or bearing pattern of brown blotches, with pointed multi-or paucispiral protoconch of 1.5 -about 3 whorls. Axial sculpture of orthocline or arcuate ribs, forming axially elongated subsutural nodules on well-defined band. Spiral sculpture from weak striae in interspaces between ribs to distinct cords, overriding ribs, often forming nodes at intersection or on lower portion of ribs. Last whorl typically shortly constricted to siphonal canal. Canal clearly separated from shell base, bearing well-pronounced fasciole.

Anatomy. Radula absent; proboscis, venom gland and accessory proboscis structure typically absent; salivary gland typically present.

Distribution: Panamic and Caribbean, off Brazil, intertidal to upper bathyal depths, typically shallow.

Remarks: Shallow-water species of the Neoterebra differ notably from their congeners described from deep water of Brazil and Caribbean -the former being larger at maturity and sharing characteristic sculpture of strong axial ribs forming elevated, sometimes axially elongated nodules on subsutural band. Caribbean species have notably weaker subsutural band, and species from deep water commonly lack a recognizable subsutural band and display a fine cancellate pattern of narrow axial riblets, intersected by spiral cords of equal strength. Although the affinity of Panamic and Caribbean species is suggested by the analysis of COI, further studies are needed to document relationships and delineate major groupings within the genus.

Etymology:

The name is a contraction of neo (latin: new) and Terebra, referring to the fact that species of this new genus only occur in East Pacific, Caribbean and off Brazil, constituting the majority of the New World terebrid fauna.

Included species:

Neoterebra acrior (Dall, 1889) Zoobank registration: urn:lsid:zoobank.org:act:3565DB17-E4EF-478B-8757-3A682BB8F6CF Definition: This genus includes species closely related to: Maculauger pseudopertusus (Bratcher & Cernohorsky, 1985), based on the molecular phylogenetic analysis (Gorson et al. submitted) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable to Maculauger pseudopertusus or any genetically proven member of the genus.

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 10.

Description: Shell small to medium-sized (<60mm), typically, orange-brown with irregular maculations on subsutural band. Whorl outline flattened, or gently convex. Subsutural band lightly ribbed or flattened, separated from lower portion of the whorl by narrow groove of row of punctations. Axial and spiral sculpture elements of about equal strength producing cancellate sculpture below subsutural band.

Anatomy. Proboscis, radula, venom gland and accessory proboscis structure absent, salivary gland present or absent.

Distribution: Indo-Pacific, subtidal and bathyal depths down to about 600 m.

Remarks:

The newly established genus Maculauger comprises several species that were formerly placed in Hastulopsis, and can be differentiated by the equal pronunciation of spiral and axial spiral elements, producing a fine cancellate sculpture when compared to similar species in Myurella. The species in Hastulopsis have very weak sculpture and are found in the Indian Ocean or endemic to Japan. Some species of Maculauger, such as Maculauger campbelli, are conchologically very close to Profunditerebra orientalis -a case where only anatomical and molecular characters can ensure correct identification (see Remarks section under Profunditerebra).

Etymology: The name of this new genus combines 'macula' referring to commonly maculated subsutural band in the species of the genus, with 'auger' -a common English epithet for Terebridae (auger shells). Gender masculine.

Included species:

Maculauger alveolatus (Hinds, 1844) 1 (Fig. 7 G -I) Type species: Terebra undulata Gray, 1834; OD herein.

Zoobank registration: urn:lsid:zoobank.org:act:EC3F14B2-3595-482B-9890-399BA78A7631 Table 10 29 Definition: This genus includes species closely related to: Myurellopsis undulata (Gray, 1834), based on the molecular phylogenetic analysis (Gorson et al. submitted) and those that show a combination of conchological, anatomical and distribution characteristics closely comparable to Myurellopsis undulata or any genetically proven member of the genus.

Diagnosis. Diagnostic nucleotide combinations in the COI barcode region provided in the Table 10.

Description: Shell small to medium-sized (<50mm), pinkish or orange, often with white subsutural band. Protoconch multispiral. Sculpture of strong rounded slightly undulating ribs, polished in appearance. Row of punctuations with groove partially cutting ribs forming subsutural band. Interstices between ribs deep and narrow, bearing regular striae and often darkly colored. Adapical parts of ribs forming one row of rounded nodules, separated from their lower portions by shallow depression or partial groove. Siphonal canal moderately long, tapering, with weakly developed fasciole.

Anatomy. Two distinct states found in species of the genus: either all typical foregut structures present, or all are lacking; in the latter case accessory proboscis structure present.

Distribution: Throughout Indo-Pacific, intertidal to upper bathyal depths (to 358 m), typically shallow.

Remarks: Species in genus Myurellopsis n. gen. are closest to the Myurella spp., but can be differentiated by the sharper more elevated ribs often with nodes on the subsutural band or top of the whorls.

Etymology:

The genus name refers to the close resemblance between its members and species in the genus Myurella. Gender feminine.

Included species:

Myurellopsis alisi (Aubry, 1999) (Aubry, 1999) Diagnosis: Shells of small size (<36mm), slender, sculptured by strong ribs, bearing nodules on subsutural band and on bottom of ribs giving concave outline to whorls. Protoconch 2.5 to 4 whorls. Remarks: Species in the genus Pellifronia are similar in having concave whorl outline but have notably weaker nodules and less clearly defined subsutural band. Species with similar nodulose sculpture exist in the genus Neoterebra but generally these have a wider apical angle to their outline or a very shiny texture. The species Duplicaria tricincta, with similar sculpture but including two raised thin spiral cords on whorls, was found to be in the genus Duplicaria.

Included species:

Granuliterebra bathyrhaphe (E. A. Smith, 1351 1875) 4 ; 1352 G. constricta [START_REF] Thiele | comb[END_REF] 4 ; 1353 G. eddunhami Terryn & Holford, 2008 4 ; 1354 G. oliverai Terryn & Holford, 2008 4 ; 1355 G. palawanensis (Aubry & Picardal, 2011) 4 ; 1356 G. persica (E. A. Smith, 1877) 4 . 1357

Genus Hastulopsis Oyama, 1961 (Fig. 10 B)

Type species: Terebra melanacme E. A. Smith, 1873; OD.

Diagnosis: Shells small to medium-sized (up to 60 mm), shiny faintly axially ribbed, with numerous fine spiral incisions. Subsutural band generally not spirally decorated, and bordered by relatively deep incision (Terryn, 2007).

Distribution: Indian Ocean and western Pacific

Remarks: The genus comprises two groups of shallow water species found along eastern coast of Africa to the Arabian Gulf and a group found in mainland Japan waters to Australia. Similar deeper water species are found in the genus Maculauger n. gen., most of which can be recognized by having a row of large spots on the periphery. The species within Myurella have a much stronger and more distinct sculpture features, and generally a brown basal area.

Included species:

Hastulopsis baliensis Terryn & Dekker, 2017 4 ; 1375 H. blanda (Deshayes, 1859) 4 ; 1376 H. cebuensis Gargiulo, 2014 4 ; 1377 H. elialae (Aubry, 1994) 4 ; 1378 H. gotoensis (E. A. Smith, 1879) 4 ; 1379 H. loisae (E. A. Smith, 1903) 4 ; 1380 H. maestratii Terryn & Rosado, 2011 4 ; 1381 H. marmorata (Deshayes, 1859) 4 ; 1382 H. masirahensis Terryn & Rosado, 2016 4 ; 1383 H. melanacme (E. A. Smith, 1873) 4 ; 1384 H. mirbatensis Terryn & Rosado, 2016 4 ; 1385 H. suspensa (E. A. Smith, 1904) 4 ; 1386 H. whiteheadae (Aubry & Marquet, 1995) 4 . 1387 [Synonym: None.] Diagnosis: Shell small to medium-sized (< 50mm), solid, axial structure weak to obsolete, spiral structure of irregular punctate groves; subsutural band weakly defined.

Genus

Distribution: Southern Pacific, particularly New Caledonia.

Remarks: Perirhoe is close to the genus Oxymeris in terms of shell proportions; however, it can be readily differentiated from species of the latter by the distinct spiral grooves throughout the whorl's height. The sculpture pattern of widely set spiral grooves in Perirhoe resembles some species of Terebra, but the latter group has notably more slender shells with higher spire and less robust siphonal canal.

Included species:

Perirhoe circumcincta (Deshayes, 1857) 4 ;

Perirhoe valentinae (Aubry, 1999) [START_REF] Bratcher | [END_REF]-Aubry 1992: fig. 9;Aubry et al. 2006: pl. 16. 11.

Description: Shell of medium size, protoconch paucispiral consisting of about 1.0-1.5 broad whorls. Teleconch of holotype with 11 whorls. Outline of teleoconch whorls straight. Spiral sculpture absent. Subsutural band beset with axially elongated nodes corresponding to axial ribs on the remainder of the whorl. Subsutural band bordered by deep and wide depression lacking punctations. Straight axial ribs stretching across whorl, sharply angular adapically and becoming more rounded abapically. Axial ribs half as wide as interspaces; 17 ribs on penultimate whorl of holotype. Axial growth lines pronounced throughout whorl height. Aperture wide, somewhat quadrate, brown with white band; columella straight with visible fold. Shell colour dark blackish brown, lighter coloured early whorls and a spiraling whitish line at the periphery.

Habitat: Retrieved from depths between 0 and 55 m. Remarks: Shell colour varies: juveniles have shiny beige to fawn colour; also lighter brown coloured specimens are known. This species has always been confused with Duplicaria mozambiquensis [START_REF] Bratcher | [END_REF], even by its authors. The holotype of Duplicaria mozambiquensis (NMSA H7843/T2541) is a small, slender shell of 22.3 mm. The type series of Duplicaria mozambiquensis shows a large discrepancy in many features between the holotype and all the paratypes which are specimens of Duplicaria herberti n. sp. Subsequent authors have to our knowledge always featured specimens of Duplicaria herberti n. sp. as Duplicaria mozambiquensis which should be considered a rarely encountered species.

Fig. 11

The whorls of Duplicaria mozambiquensis (Fig. 11D) have a narrower apical angle, its convex axial ribs giving the outline an extremely rounded convex impression, the subsutural band comprises round nodes, compared to elongated nodes on D. herberti n. sp. The axial ribs of Duplicaria herberti n. sp. become straight creating an angular projection posteriorly. The peripheral white band in Duplicaria mozambiquensis is wider and more clearly defined. Both have a subsutural furrow with minute axial growth striae, appearing as a punctate groove in D. mozambiquensis, while this feature is not present in Duplicaria herberti n. sp. We have no confirmation that Duplicaria mozambiquensis has ever been found in the south of Mozambique while specimens of D. herberti n. sp. are found extensively in the south of Mozambique and South Africa. Within Duplicaria, several species change the sculpture of their whorls as they grow and do so at variable rates of change. However a comparison of the early whorls of the two species highlights differences in the early whorls and protoconch of Duplicaria mozambiquensis and D. herberti. The protoconch of Duplicaria mozambiquensis is about 30% less wide than that of D. herberti n. sp. and the latter has a broader inflated shape. The early whorls of D. mozambiquensis have distinctive round nodes compared to convex ribs on D. herberti n. sp. Zoobank registration: urn:lsid:zoobank.org:act:6E3E67B5-1B55-4A99-8D39-6517DA4D065C Diagnostic nucleotide positions in COI alignment: see Table 11. Description: Shell small; protoconch paucispiral of a single whorl with broad nucleus. Holotype with 9 teleoconch whorls; outline of teleoconch whorls weakly convex. No subsutural band, or demarcation thereof, except for minor indentation of ribs; no visible spiral sculpture. Axial sculpture consists of almost straight, thin ribs, 14 on penultimate whorl, stretching across whorl height; interspaces notably wider than ribs. Last whorl elongate, tapering towards base. Columella straight, aperture elongate. Shell color offwhite with darker tinged spiral band adapically on the whorls and below the periphery; brownish blotches between the axial ribs abapically on the whorls; dark brown/purplish inner columella.

Partecosta bozzettii

Habitat: In sand at 3-4 m.

Distribution: Known only from the type locality.

Etymology: The species honours Luigi Bozzetti (Italy), who has contributed tremendously to the knowledge of the S Madagascar molluscan fauna.

Remarks:

The species resembles Partecosta trilineata (Bozzetti, 2008) (Fig. 11F) but differs from the latter by its more globose and convex whorls and by having a single row of brown blotches instead of two. Moreover, the protoconch of Partecosta trilineata is differently shaped, has 2 whorls with a medium nucleus. The phylogenetic analysis shows significant genetic distance between the two species. Partecosta bozzettii n. sp. shares some resemblance to Partecosta albofuscata (Bozzetti, 2008) but again differs from the latter by its more convex whorls, distinct axial ribbing and shell color pattern. Partecosta bozzettii n. sp. is similar in general morphology and shell colour pattern to Hastula daniae (Aubry, 2008) (Fig. 2G) described from West Africa but the latter species has a multispiral protoconch and more globose mature whorls. Zoobank registration: urn:lsid:zoobank.org:act:ABAFB4A7-FCD6-4632-A07C-E9945D21EA7A Diagnostic nucleotide positions in COI alignment: see Table 11.

Profunditerebra papuaprofundi

Description (holotype): Shell of moderate size to 42 mm. Protoconch of 4 narrow, conical whorls with small nucleus. Teleoconch of 12 whorls; teleoconch whorl outline convex constricted abapically at the suture. Subsutural groove represented by weak indentation on early whorls becoming sharper groove cutting through the ribs on mature whorls. Sculpture of strong slightly convex narrow axial ribs on subsutural band aligned with those on succeeding portion of whorls, varying significantly between specimens in spacing, and in number from 18 to 25 (20 ribs on penultimate whorl). No spiral sculpture in interspaces except for striae on base below periphery. Last whorl cup-shaped, becoming more elongate on mature specimens. Aperture wide with slight callus; columella straight with 2 folds. Colour off-white with tinges of yellow or pink with shiny white columella.

Habitat: Found at depths from 350-600 m.

Distribution: Known from Papua New Guinea, the Solomon Islands and Taiwan.

Etymology: Named after its deep water habitat in Papua New Guinea.

Remarks:

The only sequenced specimen of Profunditerebra papuaprofundi n. sp. with intact protoconch was designated as a holotype. It is notably smaller than other sequenced type specimens that reach shell length of 42 mm. The larger specimens demonstrate the increased curvature of ribs and increased number and density of ribs to 25 on penultimate whorl. Profunditerebra papuaprofundi n. sp. is phylogenetically closest to Profunditerebra orientalis (Aubry, 1999) and Profunditerebra anseeuwi (Terryn, 2005). Profunditerebra anseeuwi has a weaker subsutural groove, straight and oblique ribs and a mottled colour pattern. Profunditerebra orientalis has more axial ribs, a mottled colour pattern and a sculpture of ribs and spiral cords creating nodes at intersections. Specimens of Profunditerebra papuaprofundi n. sp. with more numerous curving ribs are similar to Maculauger cinctella (Deshayes, 1859) and Terebra textilis Hinds, 1844 [START_REF] Salvador | Type catalogue of Terebridae (Mollusca, Gastropoda, Conoidea) in the Natural History Museum[END_REF]. However M. cinctella (Fig. 12C) has a shorter protoconch of 2.5 whorls, a much narrower nodded subsutural band and many spiral striae within the interspaces between ribs on its whorls. In turn, Terebra textilis (Fig. 12D) differs in having a spiral groove defined by deep punctuations or partially cutting ribs and has several spiral striae in the interspaces between ribs. 11.

Profunditerebra macclesfieldensis

Description: Shell sized up to 30 mm. 12 teleoconch whorls on holotype. Protoconch multispiral with 4 translucent amber whorls and small nucleus. Teleoconch whorls with straight outline. Subsutural band defined by continuous punctuate groove. Axial ribs thin compared to interspaces gradually becoming more numerous resulting in about 25 ribs on penultimate whorl of holotype. Ribs only intersected by punctuate groove, extended onto flat subsutural band. On last whorl, the ribs fade at the periphery. No spiral sculpture visible between the ribs. Aperture long, creating elongate rounded shape to the last whorl. Columella straight with callus in mature specimens. Colour from brown to white with some brown markings.

Habitat: Found at 160-240 m.

Distribution: Known from the South China Sea, Philippines and Papua New Guinea.

Etymology: Named after the type locality, the Macclesfield bank in the South China Sea.

Remarks: In terms of shell morphology Profunditerebra macclesfieldensis n.sp. is most similar to Profunditerebra anseeuwi (Terryn, 2005) (Fig. 12G), Duplicaria duplicata (Linneaus, 1758) (Fig. 3A), and Punctoterebra teramachii (R.D. Burch, 1965) (Fig. 8B). Profunditerebra anseeuwi is the most similar in shell morphology; it has a much shorter columella creating a cup shape to the last whorl. Its ribs are set at an oblique angle and its colour is white, with broad brown spiral bands between the ribs and dark spots on the subsutural band. Duplicaria duplicata differs in having a paucispiral protoconch of 1.5 whorls, a shorter aperture and its ribs are broad and closely packed. Punctoterebra teramachii has a much narrower apical angle, the profile of its whorls is more rounded and the subsutural band narrower. Etymology: Named after the South China Sea.

Neoterebra guadeloupensis

Remarks: Maculauger campbelli (R.D. Burch, 1965) has a similar protoconch and slender overall shape with slightly convex whorls but in comparison with Maculauger sudchinensis n. sp. has a more continuous subsutural groove, less pronounced sculpture, with 5-7 spiral grooves and a distinctive pattern of straw colour with random square spots. Maculauger sudchinensis n. sp. is similar to Terebra helichrysum (Melvill & Standen, 1903) (Fig. 13F), which however has a much shorter curved columella, a multispiral protoconch of 3.5 whorls, its whorls are shorter and it has an mottled orange colour pattern. Terebra levantina (Aubry, 1999) (Fig. 13G) has similar sculpture, but its whorls are much shorter, there are 4-6 spiral grooves on its whorls, and it has a mottled colour pattern.

DISCUSSION

Terebridae systematics: a case study for integratived taxonomy This work describes the systematic revision of the venomous conoidean marine snail family Terebridae using an integrated molecular, morphological, and anatomical approach. With the increasing role of DNA sequence analysis in systematic studies, a critical step is the transition from inferred clades to formally described taxa. A phylogenetic tree allows recognition of clusters of closely related species, thus defining content of future taxa, but provides no means of ICZN valid circumscription of these taxa. Additionally, existing taxonomic practices are still deeply rooted in traditional morphology based alpha-taxonomy. Therefore, an informative morphology based diagnosis and / or description remains central for the formal establishment of new taxa.

Similarly, when contents of previously known taxa are redefined based on a phylogenetic analysis, provision of reliable diagnostic morphological characters consistent with inferred phylogenetic relationships is challenging, but notably increases the operationability of the revised taxonomic arrangement. Thus, knowing the phylogenetic relationships provides necessary insight, as it enables a 'guided' detection of inconspicuous and previously likely ignored features that may prove to be very important for understanding the morphological identity of a taxon. In this vain we applied the recent 154 species molecular phylogenetic analyses of the Terebridae (Gorson et. al. submitted) to elaborate the classification of Terebridae in combintation with shell and anterior alimentary system characters. Our results placed the 407 living species of the family, including 7 species described herein, into three subfamilies: Pellifroniinae new subfamily, Pervicaciinae [START_REF] Rudman | Observations on Pervicacia tristis (Deshayes, 1859) and comparison with other toxoglossan gastropods[END_REF]Terebrinae Mörch, 1852. Additionally, we devised an in-house computational method for analyzing COI fragments to propose DNA based diagnoses to define most of the supraspecific taxa of Terebridae. Using this approach we propose separate diagnoses for the inferred phylogenetic clusters within Terebra, Punctoterebra, and Hastula genera.

Challenges of transition from shell-based to DNA based taxonomy

Traditional molluscan shell-based taxonomy is increasingly being challenged by DNA based methods. When these two methods are in agreement it adds a significant amount of rigor and validation to the process of species delimitation. However, when shell-based and DNA findings are incongruent, deciphering a robust diagnoses is an arduous task. The notable discrepancies between morphology-based and DNA-based hypotheses of terebrid genera challenged our ability to formally establish these taxa following traditional taxonomic practices. Morphological variation in some phylogenetic lineages of Terebridae were so notable that it was extremely difficult to propose an inclusive diagnosis that would still be informative. A perfect example of such distinct conchological clusters can be found in the genus Punctoterebra, where P. solangeae is hardly distinguishable from the sympatric species of Partecosta. Species of the Punctoterebra teramachii complex demonstrate shell characteristics typical of Duplicaria, and the Punctoterebra textilis group would perfectly fit in a cluster of heavily sculptured Terebra. To cope with this situation and satisfy requirements of ICZN, we provided inclusive descriptions of genera and elaborated guidelines for their differentiation in Remarks. In cases similar to the one of Punctoterebra, we treated several morphological groups included in a phylogenetic genus separately, as each of them appeared morphologically more similar to one or several unrelated lineages, than to inferred congeners. One would argue that splitting such heterogeneous groups could have been a reasonable solution, leading at least to narrower and more robust generic diagnoses. Nevertheless, informative and seemingly more practical diagnoses would have been the only advantage of such a scheme, since they would not have made the task of delineating a morphological cluster from similar but unrelated lineages any easier. On the other hand, when sequenced species (~ 1/3 of currently accepted), and morphologically readily attributable species (~ 1/3 of species) are distributed among about 25-30 clearly defined genera, the remaining 20-30 percent of 'problematic' species, which do not match any proposed genus, end up in incertae sedis, which greatly compromises any proposed classification. However, the option of broadly defined genera, although not straightforward, allows for generally plausible placement of such divergent species. The proposed classification can be seen as a transition version that can be further elaborated with minimal rearrangement of the principal scheme established herein. The classification presented here although enclosing some compromises, fulfills its role to establish a framework consistent with the state-of-the-art phylogenetic relationships within Terebridae, and provides an account of morphological and molecular diversity in each proposed genus and subfamily, facilitating correct allocation of specimens and species.

Accounting for varying selection pressures and rates of evolution in species delimitation

Ideally, a set of morphological characteristics would be found for each delineated taxon that would allow its unmistakable recognition. However, there no reliable expectation that phylogenetically defined lineages will necessarily be readily distinguishable morphologically [START_REF] Bickford | Cryptic species as a window on diversity and conservation[END_REF][START_REF] Jörger | How to describe a cryptic species? Practical challenges of molecular taxonomy[END_REF]. The comprehension that molecular and morphological characters do not necessarily evolve with comparable rate underpins the concept of cryptic species that has profoundly changed our estimates of the global species diversity on the planet [START_REF] Knowlton | Sibling species in the sea[END_REF][START_REF] Bickford | Cryptic species as a window on diversity and conservation[END_REF]. It is to be expected that mechanisms which are accounted for morphological crypticism at the species level, be it morphological stasis or convergent evolution, can also act at a higher taxonomic level. On the contrary, descendants of a relatively recent radiation may exhibit wide range of morphologies under influence of different selection pressures on the diverging lineages as demonstrated in the terebrid genera Duplicaria, Myurella and Punctoterebra. Such cases of rapid divergence may lead to the erroneous phylogenetic hypotheses, if based solely on the morphology. Therefore, as shown with this revision of the Terebridae, the integrative approach is crucial to reconstruct phylogenetic relationships, when the molecular and morphological data considered separately lead to conflicting hypotheses of relatedness.

  clade F in relation to other studied Terebridae is sufficient, in our opinion, for recognizing it as a subfamily; the name Pervicaciinae is therefore restored and applied to it. Consequently, the two other main clades of Terebridae (clade A and the clade comprising the rest of the Terebridae), are also assigned the subfamily rank: Pellifroniinae n. subfam. for clade A, and Terebrinae for the Terebridae except clades A and F.

Finally

  , two species, Terebra walkeri E. A. Smith, 1899 and Euterebra mariato Pilsbry & Lowe, 1932 are excluded from Terebridae, and should be reassigned to Cerithiidae and Columbellidae respectively.

  Pellifronia Terryn & Holford, 2008 (Fig 2. A -D) Type species: Terebra jungi Lai, 2001; OD.

  2 ; P. jungi (Lai, 2001) 1 . Genus Bathyterebra n. gen. (Fig. 2 E -I) Type species: Terebra benthalis Dall, 1889; OD herein.Zoobank registration: urn:lsid:zoobank.org:act:EF1F9047-25AB-4400-A1D4-C705B757E114

Fig. 3

 3 Fig.3

Genus

  Partecosta Dance & Eames, 1966 (Fig. 3 F -O) Type species: Strioterebrum wilkinsi Dance & Eames, 1966 [=Terebra fuscobasis E. A. Smith, 1877]; OD.

Fig. 4

 4 Fig.4

Fig. 5

 5 Fig. 5

  Fig.9

  Type species: Terebra bathyrhaphe E. A.Smith, 1875; OD.

Fig. 10 Distribution:

 10 Fig. 10

  Pristiterebra Oyama, 1961 (Fig. 10 C) Type species: Terebra tsuboiana Yokoyama, 1922 †; OD. Genus Perirhoe Dall, 1908 (Fig. 10 E) Type species: Perirhoe circumcincta (Deshayes, 1857); OD.

  4 comb. nov. Genus Microtrypetes Pilsbry & Lowe, 1932 (Fig. 10 F) Type species: Microtrypetes iola Pilsbry & Lowe, 1932; OD.Diagnosis: Shell small (<20mm), dull, with turreted outline due to subcylindric to clearly cylindric whorl outline. Sculpture of strong and straight to fine curving axial ribs and fine spiral striae. Subsutural band indistinct or weakly defined. Siphonal canal rather long, curved.Distribution: Eastern Pacific.Remarks:The genus can be differentiated from other New World genera of Terebridae by the stepped appearance of the spire due to cylindrical or sub-cylindrical whorl outline, by generally indistinct subsutural band, and by rather long for the family siphonal canal. Included species: Microtrypetes iola Pilsbry & Lowe, 1932 4 ; M. polypenus (Pilsbry & Lowe, 1932) 4 comb. nov. Duplicaria herberti Malcolm, Terryn & Fedosov new species (Fig. 11 A -C) Duplicaria mozambiquensis

  Type material: Holotype, MNHNIM-2013-52381, lv, 29.7 mm. Paratypes: Mozambique, Inhaca Island 26°03.1'S; 33°01.0'E, 50-53 m [INHACA 2011, Stn MD13]. Paratype 1, MNHN IM-2013-52373, 1 lv, 17.4 mm; Paratype 2, MNHN IM-2013-52379, 1 lv, 23.0 mm; Paratype 3, MNHN IM-2013-52383, 1 lv, 28.2 mm. Paratype 4, type locality, MNHN IM-2013-52385, 1 lv, 30.8 mm. Paratype 5, 26°05.0'S; 32°59.0'E, 0-35 m, 24.0 mm [INHACA 2011 Stn MA15], 1 lv, MNHN IM-2013-52405, 24.0 mm; Paratypes 6 -19, YT, Mozambique, Inhaca Island, 6-8 m, 14 lv, 22.4-32.5 mm; Paratype 20, SG, Mozambique, off Inhaca Island, 25°59'59.3"S; 32°54'43.2"E, 4 m, 1 lv, 24.0 mm; Paratypes 21 -25, JR, Mozambique, off Inhaca Island, 4-8 m, 4 lv, 30.0-39.5 mm; Paratype 26: GM, Mozambique, Inhambane, 1 lv, 26 mm; Paratype 27: YT, South Africa, Natal, off Durban Bluff, in sand dredging at bay head dump, 1 dd, 32.0 mm. Type Locality: Mozambique, 26°00.0'S -32°54.4'E, 4 m [INHACA 2011 Stn MR15] Zoobank registration: urn:lsid:zoobank.org:act:C26159E5-4A5A-4218-AC7F-A76103809749 Diagnostic nucleotide positions in COI alignment: see Table

Distribution:

  Known from South Mozambique to off Durban, RSA. Etymology: The species honours Dr David G. Herbert (formerly Chief Curator of Mollusca, KwaZulu-Natal Museum, RSA), who contributed with detailed observations on the D. mozambiquensis type series at the KwaZulu-Natal Museum and provided additional historical information.

  Malcolm, Terryn & Fedosov new species (Figs 3K, 11 E) Type material: Holotype, MNHN IM-2009-10163, 12.5 mm. Paratype 1, Type locality, MNHN IM-2009-10162, 1 lv, 11.8 mm. Type Locality: S Madagascar, Port d'Ehoala, 25°03'43.9''S; 46°57'42.9''E, 3-4 m [ATIMO VATAE Stn TP29].

  Malcolm, Terryn & Fedosov new species Fig. 12 (Fig. 12 A, B) Type material: Holotype, MNHN IM-2013-58123, 19 x 4.4 mm. Paratype 1, W Buka Is, N Bougainville, N/O Alis, 5°43'S; 154°03'E, 490-530 m [MADEEP Stn DW4278], MNHN IM-2013-45571, 1 lv, 29.5 x 6.3 mm; Paratype 2, same locality MNHN IM-2013-59946, 1 lv, 25.6 x 5.1 mm; Paratype 3, same locality, MNHN IM-2013-59944, 1 lv, 13.7 x 3.7 mm. Paratype 4: W Buka Is, N Bougainville, N/O Alis, 5°35'S; 153°58'E, 623-640 m, [MADEEP Stn DW4271], MNHN IM-2013-59945, 1 lv, 16.3 x 4.3 mm. Paratype 5: W Buka Is, N Bougainville, N/O Alis, 5°37'S; 153°59'E, 421m, [MADEEP Stn DW4270], MNHN IM-2013-45509, 1 lv, 27.1 x 5.8 mm. Paratype 6: SW Santa Isabel Is., N/O Alis, 8°24'S; 159°27'E, 362-432 m [SALOMON 2 Stn CP2193], MNHN IM-2013-30574, 1 lv, 33 x 6.1 mm (broken). Other material examined: Taiwan, 15°03' N; 116°31' E [NANHAI 2014 Stn DW4102], 1 lv, IM-2013-52276, 1 lv. Type locality: Papua New Guinea, off New Ireland, 2°21'S; 150°38'E, 496-609 m [KAVIENG 2014 Stn CP4422].

  Malcolm, Terryn & Fedosov new species (Fig. 12 E, F) Type material: Holotype, MNHN IM-2013-61875, 22.4 x 4.3 mm. Paratype 1, type locality, MNHN IM-2013-61877, 1 lv, 22.2 x 4.2 mm; Paratype 2, type locality, MNHN IM-2013-61995, 1 lv, 27.2 x 5.1mm. Paratype 3, New Ireland, 2°26'S; 149°55'E, 240-242 m [KAVIENG 2014 Stn DW4485], MNHN IM-2013-58887, 1 lv, 19.2 x 3.9 mm. Paratype 4, South China Sea. YT, 200 m, 1 dd, 23.1 mm. Paratype 5 -7, Philippines, off Aliguay Island, YT, dredged at 150-200 m, 3 lv, 20.6-29.5 mm. Type locality: N Macclesfield bank; South China Sea 16°07'N; 114°23'E, 161-m [ZhongSha 2015 Stn DW4144]. Zoobank registration: urn:lsid:zoobank.org:act:F1081F38-DB65-4EDF-B73C-8743D92009A9 Diagnostic nucleotide positions in COI alignment: see Table

  Malcolm, Terryn & Fedosov new species (Fig. 13 A, B) Type material: Holotype, MNHN IM2013-61448: 17.2 x 3.5 mm. Paratypes: Guadeloupe, off Fajou, 16°22'N; 61°34', 80 m [KARUBENTHOS 2012 Stn GD02]. Paratype 1, MNHNIM-2013-20542, 1 lv, 6.5 x 2 mm; Paratype 2, MNHNIM-2013-20541, 1 lv, 5.7 x 2 mm; Paratype 3, MNHN IM-2013-20539, 1 lv, 10.4 mm x 2.6 mm. Paratype 4, Guadeloupe, Port Louis, 16° 22' 46'N; 61°34'W, 66 m [KARUBENTHOS 2012 Stn GD35], MNHN IM-2013-9110, 1 lv, 7 x 2.1 mm. Type Locality: S Marie-Galante, Guadeloupe. 15°50'N, 61°19'W, 305-312 m [N/O Antea, KARUBENTHOS 2015 Stn DW4638]. Zoobank registration: urn:lsid:zoobank.org:act:57E995EE-24F0-47A0-9DDC-E9D54948B056 Diagnostic nucleotide positions in COI alignment: see Table 11

  .

Fig. 13

 13 Fig.13

  and classification of the Conoidea (= Toxoglossa)(Gastropoda). Bulletin of the Natural History Museum of London (Zoology), 59: 125-170. TERRYN, Y. 2007. A Collectors Guide to Recent Terebridae: (Mollusca: Neogastropoda). Conchbooks, Hackenheim. TERRYN, Y. & HOLFORD, M. 2008. The Terebridae of Vanuatu with a revision of the genus Granuliterebra Oyama, 1961. Visaya, Suppl. 3: 1-118. VERDES, A., ANAND, P., GORSON, J., JANNETTI, S., KELLY, P., LEFFLER, A., SIMPSON, D., RAMRATTAN, G. & HOLFORD, M. 2016. From mollusks to medicine: a venomics approach for the discovery and characterization of therapeutics from Terebridae peptide toxins. Toxins 8: 117.

  1 comb. nov.; B. zhongshaensis Malcolm, Terryn & Fedosov 1 sp. nov.

	Subfamily Pervicaciinae Rudman, 1969

Terebra Bruguière, 1789

  Dimidacus Iredale, 1929 (nomen novum pro Terebrina Bartsch, 1923, non Rafinesque, 1815); type species: Terebra cingulifera Lamarck, 1822; by typification of replaced name; Myurellina Bartsch, 1923; type species: Terebra ornata Gray, 1834; OD; Subula Schumacher, 1817 (unnecessary replacement name for Terebra); type species: Buccinum subulatum; typification of replacement names; Terebrina Bartsch, 1923 (invalid: junior homonym of Terebrina Rafinesque, 1815); type species: Terebra cingulifera Lamarck, 1822; OD; Terebrum Montfort, 1810 (unnecessary amendation of Terebra Bruguière, 1789); type species: Buccinum subulatum Linnaeus, 1767; by typification of replaced name; Paraterebra Woodring, 1928; type species: Terebra texana Dall, 1898; OD; Panaterebra Olsson, 1967; type species: Terebra robusta Hinds, 1844; OD]

	(Figs 4, 5)
	Type species: Buccinum subulatum Linnaeus, 1767; Type by subsequent monotypy.

[Synonyms:

Triplostephanus Dall, 1908

; type species: Terebra triseriata Gray, 1834; OD; Cinguloterebra

[START_REF] Oyama | On some new facts of the taxonomy of Terebridae[END_REF] type species: Terebra hedleyana Pilsbry, 1905 (=Terebra adamsii E. A. Smith, 1873)

; OD;

Genus Hastula H. Adams & A. Adams, 1858 733

  are found in C5 as well. The combination of a very high spire and flattened whorl outline makes this morphotype very distinctive 607 among Terebridae. 608 While the three described morphotypes represent extremes of the conchological diversity of Terebra, 609 there is a plethora of intermediate forms. For example, Terebra taurina (Lightfoot, 1786) and Terebra 610 fujitai Kuroda & Habe, 1952 of the clade C3 have bifurcate ribbed subsutural band, similar to fenestrata 611 morphotype, although notably lower. Similarly, a bifurcate nodulose subsutural band is present in 612 Terebra corrugata Lamarck, 1822, a species showing close affinity to Terebra subulata in the clade C1. 613The apparent lack of phylogenetic signal in the distribution of these morphotypes across the clade C 614 precludes assignment of any taxonomic rank to them. Moreover, the general lack of homogeneity in shell 615 characteristics within subclades of Terebra makes their establishment as supraspecific taxonomic units 616 pointless as well. 617Taking into consideration extreme conchological variation within Terebra, it is difficult to suggest 618 general diagnostic criteria of this genus. However, it should be noted, that prevailing spiral sculpture 619 below subsutural band allows differentiation of Terebra from most other terebrid genera. Similarly, 620 bifurcate subsutural band has not been recorded in any species outside Terebra. These characters in 621 combination with the presence of a complete complex of foregut organs and radular teeth of hypodermic 622 type allow for reliable differentiation of Terebra spp. 623

	Fig.	734	(Fig. 6 A -E)
		735	Type species: Buccinum strigilatum 1 Linnaeus, 1758; SD Cossmann, (1896: 53).
		736	[Synonyms: Impages E. A. Smith, 1873; type species: Terebra caerulescens Lamarck, 1822 (= Buccinum
		737	hecticum Linnaeus, 1758); SD, Cossmann (1896); Acuminia Dall, 1908; type species: Buccinum
		738	lanceatum Linnaeus, 1767; OD; Hastulina Oyama, 1961; type species: Terebra casta Hinds, 1844; OD;
		739	Egentelaria Rehder, 1980; type species: Terebra stylata Hinds, 1844; OD.]
		740	Definition: This genus includes species closely related to Hastula strigilata (Linnaeus, 1758), based on
		741	the molecular phylogenetic analysis (Gorson et al. submitted) and those that show a combination of
		742	conchological, anatomical and distribution characteristics closely comparable to Hastula strigilata or any
		743	genetically proven member of the genus.
		744

  Abretia H. & A. Adams, 1853 (invalid: junior homonym of Abretia Rafinesque, 1814), type species Terebra cerithina Lamarck, 1822; SD, Dall, 1908, The Nautilus, 21(11): 125; Abretiella Bartsch, 1923;(replacement name for Abretia H. & A. Adams, 1853); type species: Terebra cerithina Lamarck, 1822; by typification of replaced name; Nototerebra Cotton, 1947; type species: Terebra albida Gray, 1834; OD]

		807	H. puella (Thiele, 1925) 1 ;	814	H. stylata (Hinds, 1844) 1 ;
	Table 4	808	H. raphanula (Lamarck, 1822) 1 ;	815	H. tenuicolorata Bozzetti, 2008 1 ;
		809	H. rufopunctata (E. A. Smith, 1877) 2 ;	816	H. tiedemani Burch, 1965 2 ;
		810	H. salleana (Deshayes, 1859) 1 ;	817	H. venus Aubry, 2008 2 ;
		811	H. sandrogorii Ryall, Terryn & Rosado, 2017 2 ;	818	H. verreauxi (Deshayes, 1857) 1 ;
		812	H. solida (Deshayes, 1857) 1 ;	819	H. westralica (Aubry, 1999) 2 ;
		813	H. strigilata (Linnaeus, 1758) 1 ;	820	H. daniae (Aubry, 2008) 3 .
			Genus Oxymeris Dall, 1903
			(Fig. 6 F -H)
			Type species: Buccinum maculatum Linnaeus, 1758; OD.
			[Synonym:
				788	H. exacuminata Sacco, 1891 2 ;
				789	H. filmerae (G. B. Sowerby III, 1906) 2 ;
		771	2 ;	790	H. hamamotoi Tsuchida & Tanaka, 1999 2 ;
		772	H. albula (Menke, 1843) 1 ;	791	H. hastata (Gmelin, 1791) 1 ;
		773	H. androyensis Bozzetti, 2008 2 ;	792	H. hectica (Linnaeus, 1758) 1 ;
		774	H. anomala (Gray, 1834) 2 ;	793	H. imitatrix (Auffenberg & Lee, 1988) 2 ;
		775	H. anosyana (Bozzetti, 2016) 2 ;	794	H. inconstans (Hinds, 1844) 2 ;
		776	H. apicitincta (G. B. Sowerby III, 1900) 2 ;	795	H. knockeri (E. A. Smith, 1872) 2 ;
		777	H. bacillus (Deshayes, 1859) 2 ;	796	H. lanceata (Linnaeus, 1767) 1 ;
		778	H. casta (Hinds, 1844) 1 ;	797	H. leloeuffi Bouchet, 1983 2 ;
		779	H. celidonota (Melvill & Sykes, 1898) 2 ;	798	H. lepida (Hinds, 1844) 2 ;
		780	H. cernohorskyi Burch, 1965 2 ;	799	H. marqueti (Aubry, 1994) 2 ;
		781	H. cinerea (Born, 1778) 1 ;	800	H. maryleeae R. D. Burch, 1965 2 ;
		782	H. continua Deshayes, 1859 2 ;	801	H. matheroniana (Deshayes, 1859) 1 ;
		783	H. crossii (Deshayes, 1859) 1 ;	802	H. nana (Deshayes, 1859) 2 ;
		784	H. cuspidata (Hinds, 1844) 2 ;	803	H. natalensis (E. A. Smith, 1903) 1 ;
		785	H. denizi Rolán & Gubbioli, 2000 2 ;	804	H. parva (Baird, 1873) 1 ;
		786	H. engi Malcolm & Terryn, 2017 2 ;	805	H. penicillata (Hinds, 1844) 1 ;
		787	H. escondida (Terryn, 2006) 2 ;	806	H. philippiana (Deshayes, 1859) 2 ;
				20

Table 5

 5 and most other terebrid genera. Finally, widely set deep spiral grooves differentiate Perirhoe spp. from Oxymeris.

		Included species:
	861	Oxymeris albida (Gray, 1834) 2 ;	872	O. felina (Dillwyn, 1817) 1 ;
	862	O. areolata (Link, 1807) 1 ;	873	O. gouldi (Deshayes, 1857) 2 ;
	863	O. barbieri (Aubry, 2008) 2 ;	874	O. lineopunctata (Bozzetti, 2008) 2 ;
	864	O. caledonica (G. B. Sowerby III, 1909) 1 ;	875	O. maculata (Linnaeus, 1758) 1 ;
	865	O. cerithina (Lamarck, 1822) 1 ;	876	O. ngai Thach, 2016 2 ;
	866	O. chlorata (Lamarck, 1822) 1 ;	877	O. senegalensis (Lamarck, 1822) 2 ;
	867	O. consors (Hinds, 1844) 1 ;	878	O. strigata (G. B. Sowerby I, 1825) 1 ;
	868	O. crenulata (Linnaeus, 1758) 1 ;	879	O. suffusa (Pease, 1869) 2 ;
	869	O. dillwynii (Deshayes, 1859) 2 ;	880	O. swinneni Terryn & Ryall, 2014 2 ;
	870	O. dimidiata (Linnaeus, 1758) 1 ;	881	O. trochlea (Deshayes, 1857) 2 ;
	871	O. fatua (Hinds, 1844) 2 ;	882	O. troendlei (Bratcher, 1981 1 .
		Genus Myurella Hinds, 1845
			(Fig. 7 A -D)
		Type species: Terebra affinis Gray, 1834; SD, Cossmann, 1896
		[Synonyms: Clathroterebra Oyama, 1961; type species: Terebra fortunei Deshayes, 1857; OD;
		Decorihastula Oyama, 1961; type species: Terebra affinis Gray, 1834; OD; Terenolla Iredale, 1929; type
		species: Terebra pygmaea Hinds, 1844; OD]

  Type species: Terebra pseudopertusaBratcher & Cernohorsky, 1985; OD herein.

			1203	N. intumescyra (Lima, Tenorio & Barros,
			1204	2007) 2 comb. nov.;
			1205	N. jacquelinae (Bratcher & Burch, 1970) 2 comb.
			1206	nov.;
			1207	N. juanica (Dall & Simpson, 1901) 2 comb.
			1208	nov.;
			1209	N. lamyi (Terryn, 2011) 2 comb. nov.;
			1210	N. larvaeformis (Hinds, 1844) 1 comb. nov.;
			1211	N. leptapsis (Simone, 1999) 2 comb. nov.;
			1212	N. limatula (Dall, 1889) 1 comb. nov.;
			1213	N. lucana (Dall, 1908) 2 comb. nov.;
			1214	N. mugridgeae (García, 1999) 2 comb. nov.;
			1215	N. nassula (Dall, 1889) 2 comb. nov.;
			1216	N. pacei (Petuch, 1987) 2 comb. nov.;
	Table		1217 1218	N. panamensis (Dall, 1908) 2 comb. nov.; N. pedroana (Dall, 1908) 2 comb. nov.;
			1219	N. petiveriana (Deshayes, 1857) 2 comb. nov.;
			1220	N. plicata (Gray, 1834) 2 comb. nov.;
			1221	N. protexta (Conrad, 1846) 2 comb. nov.;
			1222	N. puncturosa (Berry, 1959) 1 comb. nov.;
			1223	N. rancheria (Bratcher, 1988) 2 comb. nov.;
			1224	N. riosi (Bratcher & Cernohorsky, 1985) 2 comb.
			1225	nov.;
			1226	N. roperi (Pilsbry & Lowe, 1932) 3 comb. nov.;
			1227	N. rushii (Dall, 1889) 2 comb. nov.;
			1228	N. sanjuanense (Pilsbry & Lowe, 1932) 2 comb.
			1229	nov.;
			1230	N. shyana (Bratcher & Burch, 1970) 2 comb.
			1231	nov.;
			1232	N. simonei (Lima, Tenorio & Barros, 2007) 2
			1233	comb. nov.;
			1234	N. specillata (Hinds, 1844) 1 comb. nov.;
			1235	N. spirosulcata (Simone, 1999) 2 comb. nov.;
			1236	N. sterigma (Simone, 1999) 2 comb. nov.;
			1237	N. sterigmoides (Simone & Gracia, 2006) 1
			1238	comb. nov.;
			1239	N. stohleri (Bratcher & Burch, 1970) 2 comb.
	1200	N. hondurasiensis (Gargiulo, 2016) 2 comb.	1240	nov.;
	1201	nov.;	1241	N. tiarella (Deshayes, 1857) 3 comb. nov.;
	1202	N. intertincta (Hinds, 1844) 3 comb. nov.;	1242	N. variegata (Gray, 1834) 1 comb. nov.;
			1243	N. vinosa (Dall, 1889) 2 comb. nov.
	1244
	1160 1245	2 comb. nov.; Genus Maculauger n. gen. N. alagoensis (Lima, Tenorio & Barros, 2007) 2 1161
	1246		comb. nov.; (Fig. 7 E, F) 1162

ACKNOWLEDGMENTS

The material in this paper originates from several shore-based expeditions and deep sea cruises, conducted respectively by MNHN and Pro-Natura International (PNI) as part of the Our Planet Reviewed programme (SANTO 2006, INHACA 2011, ATIMO VATAE and KAVIENG 2014; PI Philippe Bouchet) and/or by MNHN, Institut de Recherche pour le Développement (IRD) and other partners as part of the Tropical Deep-Sea Benthos programme (SALOMON 2, EXBODI, TERRASSES, MAINBAZA, MIRIKY, KARUBENTHOS 2, MADEEP, NANHAI 2014, ZHONGSHA 2015; PIs Philippe Bouchet, Sarah Samadi, Wei-Jen Chen, Tin-Yam Chan). Incountry scientific partners included the University of Papua New Guinea (UPNG); National Fisheries College (Kavieng, Papua New Guinea); Universidade Eduardo Mondlane (Maputo); Parc National de la Guadeloupe, and Université des Antilles; University of Taipei and National Taiwan Ocean University (Keelung, Taiwan). Funders and sponsors included the Total Foundation, Stavros Niarchos Foundation, European Regional Development Fund (ERDF), the French Ministry of Foreign Affairs and Fonds Pacifique. Materials from expedition PANGLAO 2005 (a joint project of MNHN, University of San Carlos, Cebu City, and the Philippines Bureau of Fisheries and Aquatic Research); sampling in Congo arranged by Bernard Thomassin. Access to ship time on the French Oceanographic Fleet is gratefully acknowledged; the Taiwan and South China Sea cruises were supported by bilateral cooperation research funding from the Taiwan Ministry of Science and Technology (MOST 102-2923-B-002-001-MY3, PI Wei-Jen Chen) and the French National Research Agency (ANR 12-ISV7-0005-01, PI Sarah Samadi). Additional materials were used from fieldwork expeditions organized in collaboration with the Smithsonian Tropical Research Institute in Panama in 2006-2008 with funding to MH (NSF-CHE 0610202). All expeditions operated under the regulations then in force in the countries in question and satisfy the conditions set by the Nagoya Protocol for access to genetic resources. The authors would like to thank Philippe Maestrati, Virginie Héros, Barbara Buge and Julien Brisset for their help in curating the vouchers. The authors are grateful to Peter Dance for his remarks on taxonomy, and to Yuri Kantor and John Taylor for generously sharing their data on anatomy. The authors are grateful to Andreia Salvador for images of the NHMUK type specimens (photos copyrighted by NHMUK). This work was supported in part by the CONOTAX project funded by the French National Research Agency (grant number ANR-13-JSV7-0013-01), by funding from the Camille and Henry Dreyfus Teacher-Scholar Award and NSF awards CHE-1247550 and CHE-1228921 to MH and by the Russian Science Foundation grant RSF 16-14-10118 (PI Yuri Kantor) to AF.

Included species: 625

Terebra achates Weaver, 1960 2 ; T. adamsii (E. A. Smith, 1873) 2 ; T. albocancellata Bratcher, 1988 T. binii (Aubry, 2014) 2 ; T. boucheti (Bratcher, 1981) 2 ; T. bratcherae Cernohorsky, 1987 3 ; T. caddeyi [START_REF] Bratcher | [END_REF] T. circinata Deshayes, 1857 2 ; T. cognata E. A. Smith, 1877 3 ; T. commaculata (Gmelin, 1791) 2 ; T. connelli (Bratcher & Cernohorsky, 1985) 2 ; T. consobrina Deshayes, 1857 1 ; T. corrugata Lamarck, 1822 1 ; T. cossignanii Aubry, 2008 2 ; 651 T. cumingii (Deshayes, 1857) 1 ; 652 T. deshayesii Reeve, 1860 2 ; 653 T. donpisori Terryn, 2017 3 ; 654 T. elliscrossi (Bratcher, 1979) 1 ; 655 T. erythraeensis Terryn & Dekker, 2017 3 ; 656 T. eximia (Deshayes, 1859) 2 ; 657 T. fenestrata (Hinds, 1844) 1 ; 658 T. fernandae Aubry, 1995 3 ; 659 T. fernandesi Bouchet, 1983 3 ; 660 T. fijiensis (E. A. Smith, 1873) 3; 661 T. floridana (Dall, 1889) 3 ; 662 T. formosa Deshayes, 1857 1 ; 663 T. fujitai (Kuroda & Habe, 1952) 1 ; 664 T. funiculata Hinds, 1844 1 ; 665 T. gabriellae Aubry, 2008 3 ; 666 T. gaiae Aubry, 2008 3 ; 667 T. giorgioi Aubry, 1999 3 ; 668 T. grayi E. A. Smith, 1877 3 ; 669 T. guineensis Bouchet, 1983 3 ; 670 T. guttata (Röding, 1798) 1 ; 671 T. helichrysum Melvill & Standen, 1903 3 ; 672 T. histrio Deshayes, 1857 3 ; 673 T. hoaraui (Drivas & Jay, 1988) 1 ; 674 T. insalli (Bratcher & Burch, 1976) 2 ; 675 T. irregularis Thiele, 1925 3 ; 676 T. jacksoniana (Garrard, 1976) 3 ; 677 T. jenningsi (Burch, 1965) Diagnosis: Shell small to medium-sized (20-65 mm). Whorls outline flattened to convex with predominant pustulose, or seldom cancellate sculpture. Whorl periphery shifted abapically, leading to truncated appearance of shell, with shell base shortly constricted to short siphonal canal. Aperture wide, especially in its anterior portion.

Distribution: Western Pacific

Remarks: A characteristic truncated outline with a short siphonal canal and broad aperture allows for easy recognition of the Pristiterebra spp. among other terebrid genera. Species in Pristiterebra are nearest to the nodulose species of Neoterebra, i.e. N. tuberculosa, N. glauca and N. petiveriana, which can be recognised by the presence of a distinct subsutural groove. Some species of Gradaterebra and Partecosta have similar shell outline, but both can be differentiated from Pristiterebra by dominating axial sculpture or, rarely, complete lack of sculpture, and (Partecosta) by generally smaller adult size.

Included species:

Pristiterebra bifrons (Hinds, 1844) Diagnosis: Small (<25 mm) often dull coloured shells with flattened to bulbous whorls, axial structure of prominent ribs, spiral structure absent, microscopical or represented by fine cords; last whorl inflated.

Distribution: Indian Ocean and SW Pacific

Remarks. The genus mainly comprises small species from Australia and South Africa that can be differentiated based on adult size <25mm. 11.

DESCRIPTIONS OF NEW SPECIES

Abbreviations

Description: Shell small with orthocline spire and rather wide apical angle. Protoconch wide paucispiral, comprising 2.5 whorls with small nucleus. Teleoconch of 9 whorls Early teleoconch whorls convex in outline with strong sharply convex ribs; subsequent whorl's outline with narrow constriction formed by indentation with straight ribs below the suture. Suture distinct and ragged. Mature whorls with less convex outline, weaker and more numerous ribs, 30 on the penultimate whorl. Spiral sculpture of very weak threads. Last whorl bulbous at its base but elongate with wide aperture. Columella straight with weak fold. Overall colour white with extensive patches of light tan axial flammules on mature whorls.

Habitat: Found at depths between 470-494 m.

Distribution: Only known from type locality.

Etymology: Named for the Zhongsha sand bars, which form a subsurface atoll that sits on top of the Macclesfield Bank.

Remarks: Bathyterebra zhongshaensis n. sp. is similar to the deep-water species Bathyterebra coriolisi (Aubry, 1999) and Bathyterebra benthalis (Dall, 1889). Bathyterebra coriolisi (Fig. 2E) is similar in colour, in the sculpture of the early whorls and in the shape of its aperture. However, Bathyterebra coriolisi differs in having a much narrower shape with a strong subsutural groove and fewer ribs on the early whorls creating a band with nodes. Its subsutural groove becomes weaker and weaker with just a slight indentation on mature whorls. Bathyterebra benthalis (Fig. 2I) is similar in colour and in overall sculpture of the shell, but differs from the Bathyterebra zhongshaensis n. sp. in the outline of its mature whorls being flat and the ribs on the early whorls are extremely convex creating nodes which dominate the profile of the early whorls.

Table

Description: Shell small; protoconch multispiral, transparent with 3 whorls tinged yellow. Protoconch / teleoconch transition clear-cut with visible sinusigera. Teleoconch of holotype with 15 whorls; overall whorls outline straight to convex. Weak axial sculpture of 15 narrow axial ribs with wider interspaces increasing with maturity to 25 on penultimate whorl. Deep subsutural groove crossed obliquely by ribs. Subsutural band with strong elongate nodes; subsequently, two strongly projecting spiral cords at top and middle of whorl dominating sculpture forming a reticulate pattern of deep indentations with thin ribs. The upper cord projects in outline beyond the subsutural band. On the mature whorls interspaces sometimes with one or two weak spiral striae. Reticulate sculpture continuing on shell base; aperture long with weak callus; columella with 2 weak folds. Shell lacks any pattern, coloured in shades of white to yellow.

Habitat: Deep water 60-310 m.

Distribution: Only known from off Guadeloupe.

Etymology: Named after the type locality, the French Caribbean island of Guadeloupe.

Remarks: A number of Terebridae species have been described from the Western Atlantic and Caribbean seas which have a similar strong reticulate sculpture. These include Neoterebra limatula (Dall, 1889) which has more than 2 strong spiral cords in addition to the subsutural band (Fig. 13C) and has a paucispiral protoconch. Neoterebra colombiensis (Simone & Gracia, 2006) is very similar to Neoterebra guadeloupensis n. sp. but the spiral sculpture in the former species is less dominant giving a more uniform reticulate sculpture with nodes at intersections. Furthermore, Neoterebra colombiensis has a paucispiral protoconch in comparison to the multispiral protoconch of the N. guadeloupensis n. sp. Neoterebra simonei (De Lima, Tenorio & De Barros, 2007) is similar in colour and sculpture, however, it has 3 spiral cords in addition to the subsutural band and has a distinctive paucispiral protoconch of 1.5 whorls. Other similar species from the region e.g. Neoterebra intumescyra (De Lima, Tenorio & De Barros, 2007), Neoterebra alagoensis (De Lima, Tenorio & De Barros, 2007) (Fig. 9F) all have paucispiral protoconchs and differing sculpture patterns. 11.

Maculauger sudchinensis

Description: Shell sized to 21 mm, with rather narrow apical angle. Protoconch paucispiral of 1.5 translucent whorls. Teleoconch of holotype with 14 whorls; outline of teleoconch whorls slightly convex. Subsutural band formed by deep punctures. Axial ribs narrow with wide interspaces; ribs circa 18-20 on penultimate whorl, straight on subsutural band, becoming oblique at subsutural groove and then curved on abapical part of whorls. Axial ribs intersected by 2 to 3 evenly spaced spiral grooves. Aperture long with straight columella. Ground colour chalky white, sometimes with vague brown marks on subsutural band.

Distribution: Only known from the N Macclesfield Bank.

Habitat: Found at depths of 160-220 m.

CAPTIONS