New species of the genus Otitoma Jousseaume, 1898 (Pseudomelatomidae, Conoidea) from the Western Pacific Ocean

Twelve new species are assigned to the genus Otitoma Jousseaume, 1898 in the family Pseudomelatomidae Morrison, 1966 and herein described: O. hadra sp. nov., O. neocaledonica sp. nov., O. rubiginostoma sp. nov and O. tropispira sp. nov. from New Caledonia; O. boucheti sp. nov., O. nereidum sp. nov. and O. sororcula sp. nov. from the Fiji Islands; O. xantholineata sp. nov. from the Solomon to the Fiji Islands; O. crassivaricosa sp. nov. from Fiji to Hiva Oa Island (Marquesas Archipelago); O. philpoppei sp. nov. from the Philippines but also reported from the Fiji Islands; O. elegans sp. nov. from the Fiji Islands and O. philippinensis sp. nov. from the Philippines. New data on O. carnicolor (Hervier, 1896) are provided. Otitoma mitra (Kilburn, 1986), from Southern Mozambique, is here considered a synonym of O. cyclophora (Deshayes, 1863). Drillia batjanensis Schepman, 1913, previously assigned to the genus Maoritomella Powell, 1942 in the family Borsoniidae Bellardi, 1875, is here assigned to the genus Otitoma. Photographs of the holotype of Drillia batjanensis are provided for the first time. In addition, color photographs of the type specimens of the following species are provided: Drillia kwandangensis Schepman, 1913, D. timorensis Schepman, 1913 and Mitrellatoma mitra Kilburn, 1986.


Introduction
For numerous years, the French Institut de Recherche pour le Développement (IRD) and the Musèum national d'Histoire naturelle (MNHN) of Paris have been conducting an intensive sampling program in the Indo-Pacific region, particularly in waters surrounding New Caledonia (Bouchet et al. 2008).As a result of such efforts an unprecedented amount of turriform gastropods has been accumulated.This material provided the basis for several studies which have been published over the last two decades, exploring the extreme morphological diversification among the Conoidean groups previously assigned to Turridae (sensu Powell 1966), but which has since been resolved in 13 distinct families (Bouchet et al. 2011).We had the opportunity to examine numerous lots of material assigned to Otitoma Jousseaume, 1898 following preliminary sorting.The generic name Otitoma Jousseaume, 1898 was rediscovered by Kilburn (2004), who treated Thelecytharella Shuto, 1969 as a synonym.Otitoma and Thelecytharella have recently been considered distinct genera by Wiedrick (2014) on the basis of the anal sinus structure.This feature, however, seems variable within certain genera belonging to the family Pseudomelatomidae Morrison, 1966 and we accordingly prefer to follow Kilburn in considering Thelecytharella a synonym of Otitoma.Kantor et al. (2008) demonstrated that shell features of turriform gastropods may be inadequate for supraspecific classification.The status of Thelecytharella, as a synonym of Otitoma, will therefore remain uncertain until anatomical and/or molecular features become known.Otitoma and Thelecytharella include respectively seven and 11 recognized Pleistocene to Holocene species (see WoRMS Editorial Board 2016).In the present paper, we describe 12 new species of Otitoma (= Thelecytharella), which represent the more distinctive species among the available material.

Material and methods
Inclusion of Otitoma in the family Pseudomelatomidae follows Bouchet et al. (2011) and Puillandre et al. (2011).The material examined is represented by empty shells.Descriptions and measurements were based on shells oriented spire up with the aperture facing the viewer (front view) and turned 90° counter clock-wise (side view).Photographs were made using a Motic SMZ-140-N2GG stereo microscope.SEM micrographs of selected samples mounted onto stubs and coated with gold were taken with a Hitachi S-2400 scanning electron microscope and a JEOL-5200 scanning probe microscope.

Diagnosis
Shell of small to moderate size, up to about 19 mm, with cylindric-fusiform or pupoid shape and constricted base.Sculpture predominantly spiral of weak to distinct cords sometimes plicate by axial ribs.
Outer lip edge sharp, preceded by a varix, anal sinus rounded, U-shaped.Stromboid notch distinct.Inner lip with a parietal pad which constricts entrance to anal sinus.Protoconch paucispiral and papilliform to multispiral bluntly or narrowly conical.
Remarks Shuto (1969: 208) proposed Thelecytharella as a subgenus of Euclathurella, selecting Agladrillia oyamai Shuto, 1965 from the Pleistocene of Kyushu (Japan) as the type species.In 1970, the same author erected the genus Lioglyphostomella to accommodate two Recent species from Indonesia, Drillia kwandangensis Schepman, 1913 andD. timorensis Schepman, 1913.Later, this author described Austropusilla (Metaclathurella) crokerensis Shuto, 1983 from the Arafura Sea (Shuto 1983).Subsequently, Kilburn (1995) moved the genus Thelecytharella to the Mangeliinae and recognized two South African species, T. oneili (Barnard, 1958) and T. metuloides Kilburn, 1995.In this paper we follow Kilburn (1995), who placed Metaclathurella and Lioglyphostomella in junior synonymy with Thelecytharella and demonstrated that Austropusilla is not related.A few years later, Sysoev (1997) described Thelecytharella kecil from Indonesia and demonstrated that it and T. vitrea (Reeve, 1845) are morphologically distinct and may actually represent a genus separate from Thelecytharella.Kilburn (2004) resurrected the genus Otitoma, previously considered a nomen dubium by Powell (1966: 126), and demonstrated that its type species, Otitoma ottitoma Jousseaume, 1898 from Yemen, is a synonym of the widely distributed Indo-Pacific Pleurotoma cyclophora Deshayes, 1863.This latter species had been assigned to the genus Daphnella Hinds, 1844, subgenus Hemidaphne Hedley, 1918 by Powell (1966).Kilburn (2004) described the similarities between Otitoma and Thelecytharella and considered the latter generic name a synonym of Otitoma.More recently, Wiedrick (2014) regarded both taxa as valid and described two new species, namely O. astrolabensis and "Otitoma" fergusoni, the latter assigned to genus Otitoma with doubt.Daphnella deluta Gould, 1860, described from the ambiguous locality "China Sea", was assigned to the genus Otitoma by Wiedrick (2014).We have not examined the type material, but judging from photographs of the lectotype (USNM 24225), the species appears to be raphitomine and its inclusion in Otitoma is uncertain.Conversely, Drillia batjanensis Schepman, 1913, described (Beets 1984;Shuto 1970;Sysoev 1997), but photographs of the holotype (ZMA 136858), clearly indicate that the species belongs to genus Otitoma.Otitoma batjanensis is similar to the holotype of O. crokerensis (Shuto, 1983) (see Australian Museum Collections).In this paper we follow Kilburn (2004) and tentatively treat Thelecytharella as a synonym of Otitoma, because in our opinion, the morphological features proposed by Wiedrick (2014) as diagnostic of the two groups are of doubtful value, at least at the supraspecific level.The systematic position of Otitoma (= Thelecytharella) has been a subject of debate.Shuto (1969) and Kilburn (1995) referred it to the subfamily Mangeliinae (now Mangeliidae), while Sysoev (1997)

Remarks
This species, the type of the genus Otitoma, has a very broad distribution from the southern Red Sea and Mozambique to Japan, Fiji and Polynesia.Kilburn (2004) selected a neotype from Aden and redescribed the species on the basis of local material from Mozambique.The same author determined Daphnella crenulata Pease, 1868, described from Polynesia, to have been based on a juvenile specimen of O. cyclophora.Our material (Fig. 1A-C, G-L) matches that figured and discussed by Kilburn (2004) in morphological features and is, accordingly, considered as conspecific.Kilburn (1986) introduced Mitrellatoma mitra for a single shell from Southern Mozambique; the taxon was later cited by the same author in his list of species belonging to the genus Otitoma (Kilburn 2004), but without a formal European Journal of Taxonomy 304: 1-30 (2017) discussion of the reassignment.In our opinion, the holotype of Mitrellatoma mitra (Fig. 1E-F) is a juvenile specimen that perfectly matches the morphological features of the lectotype of Daphnella crenulata, and the former species is also considered a synonym of O. cyclophora here.
Otitoma cyclophora has previously been considered a littoral species (Kilburn 2004), but the present records indicate it can also live in deep water (down to 600 m).The available specimen from deep water (Stn DW626) measures 13.7 × 4.0 mm, aperture height 6.4 mm and is the maximum size reported for the species.As already noted by Kilburn (2004), O. cyclophora has a well-developed secondary spiral sculpture of numerous closely-spaced, fine spiral threads (Fig. 1I).

Description
Shell claviform-fusiform (b/l 0.36-0.37;a/l 0.44-0.50),somewhat pupoid.Teleoconch of 4½-4¾ weakly and evenly convex whorls with periphery just below mid-whorl height on earlier two whorls, at mid-whorl on later ones.Suture shallow, bordered by a sub-sutural fold, demarcated by a shallowly concave sutural ramp.Axial ribs opisthocline and rounded, equal in width to or wider than intercostal space, extend from whorl base to sub-sutural fold but becoming rather weak on sutural ramp.12-13 axial ribs on penultimate whorl, 12-16 on last whorl.Spiral sculpture (Fig. 1P) of narrow, widely-spaced spiral cords, not forming distinct nodules where crossing axial ribs.First two teleoconch whorls with a sub-sutural cord, 1-2 weak cords (abapical one stronger) on sutural ramp and two main cords below ramp, 1-2 additional main cords develop on subsequent two whorls.Last whorl with 8-9 spiral cords on base and rostrum.Interspaces between spiral cords sculptured by dense, closely-spaced fine spiral threads irregularly plicate by axial incremental lines.Aperture rather narrow, slightly constricted at entrance to siphonal canal, the latter short, slightly expanded terminally, somewhat obliquely truncated.

Remarks
This species was originally described from Lifou Island and cited by Bouge & Dautzenberg (1914).It has subsequently been ignored until Kilburn (2004)

Description
Shell claviform-fusiform (b/l 0.36-0.44;a/l 0.43-0.50),somewhat pupoid.Teleoconch of 3½-4¾ convex whorls with median periphery.Suture distinct, bordered by a prominent spiral cord, demarcated by a distinctly concave sutural ramp.Axial ribs rather straight, opisthocline and distinctly rounded, broader than intervals between them, developed mainly in peripheral region, becoming much weaker to obsolete and slightly arcuate on sutural ramp.8-11 axial ribs on penultimate whorl, 11-14 on last whorl.Spiral sculpture of prominent, narrow, widely-spaced spiral cords, becoming conspicuously nodulose where crossing axial ribs (Fig. 2F).Earlier 1-2 whorls with a subsutural cord and two cords on abapical part of whorl, increasing to 3 and 4 on penultimate and last whorl, respectively.Sutural ramp with 1-2 weak spiral cords on abapical side.Last whorl with 9-12 spiral cords on base and rostrum.Interspaces between spiral cords lacking secondary sculpture (Fig. 2G).Aperture rather narrow and almost linear, but strongly constricted at entrance to siphonal canal.Siphonal canal short, relatively wide, expanding slightly distally.Inner lip with moderately thick callus with distinct parietal pad.Outer lip edge sharp, preceded by a wide, low, rounded varix.Stromboid notch distinct.Anal sinus rather deep, apically rounded, asymmetrically U-shaped, with strongly constricted opening.Protoconch papilliform, of ¼-1½ MORASSI M. et al., New species of the genus Otitoma whorls (Fig. 2H-I), diameter 0.62-0.89mm.Color white or yellowish-white with an amber-orange band bordering the adapical suture.

Remarks
Among its described congeners, Otitoma sororcula sp.nov.most closely resembles Otitoma kwandangensis (Schepman, 1913) (Fig. 2J-K).The two species are nearly indistinguishable from each other in teleoconch morphology, but differ in protoconch structure, which in O. sororcula sp.nov. is papilliform, of 1¼-1½ whorls while in Otitoma kwandangensis it "is broken at the tip and the preserved part consists of 2.7 volutions" (Shuto 1970: 166, plate X, fig. 19).Otitoma sororcula sp.nov. is remarkably smaller than the type specimen of Drillia kwandangensis (4.3-6.6 mm vs 8.2 mm, mean 5.15 mm in length) and differs in color (white or yellowish-white with an amber-orange band bordering the adapical suture and tinged with the same color on the adapical part of the base vs a uniform red-brown).The new species is comparable to Otitoma timorensis (Schepman, 1913), in protoconch features and shape, but judging from the photographs of the holotype ZMA 138096 (Fig. 2L-N), Otitoma sororcula sp.nov.differs in lacking a secondary spiral sculpture, having a different number of spiral cords (3 instead of 2 on penultimate whorls), which are more distinctly nodulous where crossing the axial ribs, and in having broader and more bulging axial ribs.

Etymology
The specific epithet comes from the Greek 'xanthos' and Latin 'lineata', alluding to the brownish orange spiral cords.

Description
Shell cylindric-fusiform (b/l 0.37-0.41;a/l 0.48-0.54).Teleoconch of 4-5 weakly and evenly convex whorls with periphery at mid-whorl height.Suture shallow and bordered by a sub-sutural fold with a prominent spiral cord.Sutural ramp concave on earlier whorls, more shallowly concave on later ones.Axial sculpture absent (holotype) or consisting of ribs restricted to earlier two teleoconch whorls.Spiral sculpture consisting of prominent, angular and well-spaced cords.First two teleoconch whorls with a sub-sutural cord and two cords between sutural ramp and whorl base, increasing to 3 on antepenultimate whorl, 3-4 and 4-5 on penultimate and last whorl, respectively.Sutural ramp sculptured with none (usually first whorl) to 1 weak cord on earlier whorls; later two whorls with two spiral cords, abapical one strongest.Last whorl with 12-14 spiral cords on base and rostrum.Interspaces between spiral cords lacking secondary sculpture (Fig. 3E).Surface covered only by relatively coarse axial incremental lines.Aperture pyriform, constricted at entrance to siphonal canal.Siphonal canal relatively wide, expanding slightly distally, oblique, not notched.Inner lip covered with thin callus, most of its parietal region occupied by a thin pad.Outer lip edge sharp, preceded by a wide, low and rounded varix.Stromboid notch distinct.Anal sinus U-shaped, its entrance only slightly constricted by parietal pad.Protoconch domed, of 1½-1¾ smooth whorls with distinct protoconch/teleoconch transition, white, diameter 0.77-0.90mm (Fig. 3F-G).Teleoconch yellowish-white to pale orange-yellow, with light brownish orange spiral cords and aperture edged with brownish orange.

Remarks
Otitoma xantholineata sp.nov. is similar in shape and number of spiral cords to O. kagoshimaensis from the Upper Pleistocene of Moeshima Island, Japan (3 on first teleoconch whorl increasing to 4-5 on later whorls), based on a single specimen of the latter, which is lacking the spire apex.However, O. kagoshimaensis is sculptured by axial ribs on all teleconconch whorls (Shuto 1965: 156)  European Journal of Taxonomy 304: 1-30 (2017) a more shallowly concave sutural ramp compared to O. kagoshimaensis.Otitoma xantholineata sp.nov. is superficially similar to two Recent species: Drillia batjanensis Schepman, 1913 from Indonesia (Fig. 3H-I) and O. crokerensis (Shuto, 1983) from the Arafura Sea (Fig. 3J-L).Otitoma xantholineata sp.nov. is readily distinguished from D. batjanensis in protoconch structure (domed, of 1½-1 ¾ whorls rather than bluntly conical, of 3 whorls), and by spiral sculpture features.The two species have a comparable number of spiral cords on spire whorls, but these are of nearly uniform strength in Otitoma xantholineata sp.nov., while in D. batjensis they are more "keel-like", with those on abapical side of the whorl more prominent than the others.Furthermore, O. xantholineata sp.nov.has vestigial axial sculpture on earlier whorls and distinct axial incremental lines on the entire surface, while there is no indication of axial sculpture in D. batjensis.Otitoma xantholineata sp.nov.may superficially resemble O. crokerensis (Shuto, 1983), from which it is distinctly different in protoconch features (O.crokerensis has the same protoconch structure as D. batjensis), in having fewer and narrower spiral cords, in the profile of spiral whorl(evenly convex instead of slightly shouldered) and in the more defined siphonal canal.

Etymology
The specific epithet comes from a combination of the Greek 'tropis' (a keel) and the Latin 'spira' (spire), alluding to the strong spiral sculpture of the keel-like cords.

Description
Shell fusiform (b/l 0.34-0.36;a/l 0.42-0.50).Teleoconch of 6-7 convex whorls with periphery at middle on earlier whorls, posterior to mid-whorl on later ones.Suture moderately deep, bordered by a prominent sub-sutural fold bearing a spiral cord, demarcated by a distinctly concave sutural ramp.Sculpture consisting of widely-spaced, narrow, keel-like spiral cords.Four earliest teleoconch whorls with a sub-sutural cord and two cords on abapical half of whorl, increasing to 3 on fifth and 4 on later two whorls.Last whorl with 11-15 spiral cords on base and rostrum.Sutural ramp with a narrow weak cord on its abapical part, becoming stronger on later two whorls (in paratype 5 (MNHN IM-2000-32595) comparable in strength to main spiral cords).Interspaces between spiral cords sculptured by numerous, very fine and faint spiral threads varying in number (up to about 13) (Fig. 3P).Axial sculpture represented only by incremental lines, more evident on sutural ramp, where they are arcuate in conformity with curvature of anal sinus.Aperture rather narrow, lanceolate, strongly constricted at entrance to siphonal canal.Siphonal canal long for genus, not notched terminally.Inner lip with moderately thin callus, European Journal of Taxonomy 304: 1-30 (2017) parietal region with a pad formed at intersection with outer lip.Outer lip edge sharp, preceded by a wide, strong, rounded varix.Stromboid notch distinct, deep.Anal sinus deep, U-shaped, with strongly constricted opening.Protoconch conical, of 2½-2¾ smooth whorls.Protoconch diameter 0.81-0.92mm (Fig. 3Q-R).Color white or cream, with yellowish-orange spiral cords.

Remarks
Otitoma tropispira sp.nov. is distinguished from its described congeners by its rather large size (attains about 19 mm in maximum length) and the sculpture of its keel-like spiral cords, somewhat reminiscent of members of the pseudomelatomid genus Turridrupa Hedley, 1922.However, members of that genus have a weak to absent parietal pad, which does not constrict entrance to the anal sinus, and the interior part of the aperture is sculptured by spiral plicae.Furthermore, the protoconch of O. tropispira sp.nov. is smooth, while that of Turridrupa bears axial riblets on the last 1½-2 whorls (Kilburn 1988;Powell 1966).

Description
Shell fusiform (b/l 0.33; a/l 0.52).Teleoconch of 6 whorls, moderately convex, with a slightly angular shoulder posterior to mid-whorl.Suture shallow, bordered by a narrow and weak sub-sutural cord and a shallowly concave sutural ramp.Axial sculpture absent.Spiral sculpture of widely-spaced and rather narrow cords.Three earliest teleoconch whorls with sub-sutural fold and two cords on abapical half of whorl; fourth whorl with two additional cords, one secondary cord at abapical margin of sutural ramp and one main cord margining whorl base.Penultimate whorl with three main and four weaker cords, increasing in strength in such a way that there are 7 cords of nearly uniform strength on last whorl (Fig. 4C).Last whorl with 21 spiral cords on base and siphonal canal.Interspaces between cords sculptured by 2-5 spiral threads (Fig. 4D).Aperture rather narrow and ovate, constricted at entrance to siphonal canal.Siphonal canal long for genus, not notched terminally.Inner lip with moderately thin callus, parietal region with a pad formed at junction of outer lip.Outer lip edge sharp, preceded by a wide, low, rounded varix.Stromboid notch moderately deep.Anal sinus deep, U-shaped, with constricted opening.Protoconch narrowly conical, of 2¾ smooth whorls, diameter 0.76 mm (Fig. 4E-F).Color yellowish-white.

Remarks
In its large size (up to about 15 mm in length) and yellowish-white shell, Otitoma boucheti sp.nov.may superficially resemble O. tropispira sp.nov., but is readily distinguished from the latter in having more numerous, but much narrower and less prominent spiral cords, and a longer siphonal canal.

Etymology
The specific epithet alludes to the origin of the type material (New Caledonia).

MORASSI M. et al., New species of the genus Otitoma
rather straight, opisthocline and rounded, equal to or wider than intercostal space, developing mainly in peripheral region, not reaching suture.12-15 axial ribs on penultimate whorls; on last whorl 11-17 ribs, but reduced to 8-10 in specimens with ribs fading on last portion of whorl.Sculptured by widely-spaced spiral cords, crossing low axial ribs, not forming distinct nodules.Earliest two teleoconch whorls with 2-3 spiral cords between sutural ramp and whorl base, increasing to 4-6 on later two whorls.Sutural ramp with 2 main cords on abapical side and 1-2 finer cords posterior to these.Last whorl with 12-17 main spiral cords, interstices with none to 1-2 secondary cords and with finer interstitial threads on base and rostrum.Under SEM (Fig. 4J-K) intervals between main cords seen to be sculptured by 2-4 rather inconspicuous, microscopic spiral threads and numerous granules.Entire surface covered by coarse axial growth lines.Aperture rather narrow and almost linear, but constricted at entrance to siphonal canal.Siphonal canal relatively wide, expanding slightly distally, oblique, not notched terminally.Inner lip with moderately thin callus, but distinct parietal pad.Outer lip edge sharp, preceded by a wide, low, rounded varix.Anal sinus moderately deep, apically rounded, asymmetrically U-shaped, with strongly constricted opening.Stromboid notch distinct.Protoconch domed to papilliform, of 1¼-1¾ whorls; first whorl smooth, last portion with numerous irregular axial riblets, diameter 0.67-0.88mm (Fig. 4L-M).Protoconch and earlier teleoconch whorls violaceous in some specimens (Fig. 4O-N).Two color forms: a) white or yellowish-white tinged with light brownish-orange on sub-sutural ramp, base and tip of rostrum, and aperture rimmed with darker brownish orange; b) a uniform light brown (paratype 6).

Remarks
Otitoma neocaledonica sp.nov. is morphologically similar to O. oneili (Barnard, 1958), but differs from the latter in numerous characters.The new species is smaller in length (up to 8.2 mm vs up to 12.3 mm in maximum length), with different proportions (a/l 0.45-0.52vs 0.32-0.41),fewer secondary cords on the sub-sutural ramp and the presence of weaker, inconspicuous microscopic cords in the interspaces between the main cords.Otitoma neocaledonica sp.nov.has a different protoconch structure (domed-papilliform and of 1¼-1¾ whorls instead of bluntly conical, of about 3 whorls).Furthermore, the protoconch and the earlier teleoconch whorls are violaceous in most specimens of Otitoma neocaledonica sp.nov., a feature not reported in O. oneili (Kilburn 1995 and pers. obs).

Etymology
The specific epithet refers to the Greek 'hadros' (thick), alluding to the small but very thick shell.

Description
Shell somewhat biconic (b/l 0.41-0.48;a/l 0.56-0.57).Teleoconch of 3-3¼ weakly and evenly convex whorls, with a slight shoulder at mid-whorl height.Suture shallow, bordered by a weakly prominent sub-sutural fold followed by a shallowly concave sutural ramp.Axial sculpture of opisthocline ribs, subequal to or narrower than intercostal spaces, abruptly ending at level of abapical side of sutural ramp (Fig. 4S).14-16 axial ribs on penultimate and 18-19 ribs on last whorl.First teleoconch whorl with 3 spiral cords between the sutural ramp, sutures of following whorl increasing to 4-5 on penultimate and last whorl.Spiral cords closely-spaced, not plicate where crossing axial ribs.Sutural ramp with 1 spiral cord increasing to 2 cords on later two whorls.Last whorl with 19-22 main spiral cords and 1 occasional secondary cord in interspaces on base and rostrum.Interspaces between cords lacking secondary sculpture.Aperture rather narrow, ovate, constricted at entrance to siphonal canal.Siphonal canal short, relatively wide.Inner lip covered with a thick callus.Outer lip edge sharp, preceded by a strong, wide varix.Stromboid notch weak.Anal sinus relatively broad and moderately deep, a C-shaped notch, its entrance weakly constricted.Protoconch domed, of up to 1½ smooth whorls, diameter 1.04-1.09mm.Protoconch white, teleoconch yellowish-white flecked with orange-yellow on sutural ramp and along some axial ribs on last whorl.

Remarks
Otitoma hadra sp.nov. is readily distinguished from its described congeners by its rather small, thick shell (about 5 mm in maximum length), with a relatively large protoconch and well developed sculpture (both spiral and axial).

Etymology
The specific epithet is a ombination of the Latin words 'rubiginosus' (rusty) and 'stoma' (aperture), alluding to the rust-like color of the outer lip edge and parietal pad nodule.

Description
Shell cylindric-pupoid (b/l 0.38-0.45;a/l 0.44-0.55).Teleoconch of 2½-3½ whorls which are weakly and evently convex to roundly shouldered above mid-whorl height.Suture shallow, bordered by a moderately prominent sub-sutural fold with two closely spaced cords, followed by a shallow, concave sutural ramp.Axial sculpture of opisthocline ribs, subequal to or narrower than intercostal space, extending on spire whorls from suture to suture, becoming much narrower and strongly arcuate on sutural ramp.13-16 axial ribs on penultimate whorl and 16-18 on last whorl.First teleoconch whorl MORASSI M. et al., New species of the genus Otitoma with 2-3 spiral cords between whorl base and sutural ramp, increasing to 4-5 on penultimate whorl and 5-6 on last whorl.Spiral cords somewhat plicate where crossing axial ribs.Interspaces between main cords with none to 1-2 weaker cords (Fig. 5D-E).Sutural ramp with 1 weak spiral cord on first whorl, increasing to two on following whorl(s) (abapical one slightly more prominent); additional fine, inconspicuous cords on abapical side of sutural ramp of last whorl (in holotype).Base sculptured by 13-15 spiral cords, with none to 1-2 interstitial cords.Surface covered by dense axial incremental lines, particularly evident on sutural ramp, where they are sinuous in conformity with curvature of anal sinus.Aperture narrow and almost linear, but constricted at entrance to siphonal canal.Siphonal canal wide, expanding slightly distally, not notched.Inner lip covered with a thin callus and parietal pad.Outer lip edge sharp, preceded by a wide, low and rounded varix.Stromboid notch deep.Anal sinus deep, rounded, asymmetrically U-shaped, its entrance slightly constricted.Protoconch domed and flat, of 2¼-2½ whorls; first whorl smooth, subsequent whorl with numerous inconspicuous, widely-spaced axial riblets and sparse microgranules (Fig. 5H).Protoconch white, diameter 0.71-0.89mm.Teleoconch white, with outer lip edge and parietal pad tinged with brown-orange.

Remarks
Otitoma rubiginostoma sp.nov. is a morphologically distinctive species.The profile of the whorls and type of axial ribbing (i.e., axial ribs becoming strongly arcuate on sutural ramp) may cause it to superficially resemble O. cyclophora, but the two species are otherwise very different in numerous characters.Otitoma rubiginostoma sp.nov. is much smaller in length (up to 4.7 mm vs 13.7 mm), with fewer teleoconch whorls (2½-3½ instead of 4½-5) and it has a different shape (cylindric-pupoid instead of cylindric-fusiform).O. rubiginostoma sp.nov.has strong axial ribs while in most specimens of O. cyclophora these are weak to obsolete, particularly on the last two teleoconch whorls.The protoconch of O. rubiginostoma sp.nov. is much larger than that of O. cyclophora (0.71-0.89 mm vs 0.52-0.59mm) with different a shape and number of whorls (domed and flat of 2¼-2½ vs papilliform of about 1¾ whorls).The two species further differ in color: O. rubiginostoma sp.nov.has a white shell with the outer lip edge and parietal pad tinged with brown-orange while O. cyclophora has a uniform pale buff shell.The new species is similar to O. vitrea and O. kecil, both of which possess large, domed protoconchs, but their teleoconch characters are otherwise too different for meaningful comparison.

Etymology
The specific epithet refers to the Latin 'elegans', meaning 'characterized by refined grace', alluding to the aesthetically pleasant appearance of the shell.

Description
Shell fusiform (b/l 0.38; a/l 0.55).Teleoconch of slightly less than 7 convex whorls, with periphery at middle of the 3 earliest whorls, posterior to mid-whorl on the later whorls.Whorls separated by a distinct suture bordered by a spiral cord.Sutural ramp moderately concave.Axial sculpture consisting of opisthocline ribs, subequal to broader than intercostal spaces, extending on spire whorls from suture to abapical side of sutural ramp, prominent on earlier whorls, but becoming remarkably weaker on last whorl.21 axial ribs on penultimate whorl and 17 on last whorl.Spiral sculpture consists of widelyspaced, narrow, weakly prominent spiral cords, somewhat plicate where crossing axial ribs.Earliest three whorls with two cords between sutural ramp and abapical suture, increasing to three cords on fourth and antepenultimate whorls and four on penultimate and last whorls.Last whorl with 13 spiral cords on base and rostrum.Insterpaces between cords sculptured by numerous fine spiral threads (up to 14) (Fig. 5K-L).Aperture rather narrow, lanceolate, constricted at entrance to siphonal canal.Siphonal canal moderately long, not notched terminally.Inner lip with moderately thin callus, parietal region with a pad formed by termination of outer lip.Outer lip edge sharp, preceded by a wide, low, rounded varix.Anal sinus moderately deep, apically rounded, asymmetrically U-shaped, not strongly constricted at opening.Stromboid notch moderately deep.Protoconch of 1½ smooth, domed whorls with few sparse microscopic granules when observed under SEM (Fig. 5N), diameter 0.86 mm.Color yellowish-white with brown-orange spiral cords.

Remarks
Although only a single specimen of Otitoma elegans sp.nov. is available, this species differs distinctly from its described congeners in having only a few, widely-spaced primary spiral cords with well developed secondary spiral sculpture, and a peculiar color pattern (yellowish-white with brown-orange spiral cords).O. elegans sp.nov.further differs from its congeners of similar size in possessing a paucispiral and domed protoconch instead of multispiral and conical.

Etymology
The specific name is derived from the country of the origin of the type.

Description
Shell fusiform (b/l 0.33-0.34;a/l 0.46-0.47).Teleoconch of 6 convex whorls with periphery below mid-whorl height on earliest four whorls, at mid-whorl on later two whorls.Whorls separated by a distinct suture bordered by a weakly prominent spiral cord.Sutural ramp rather wide and shallowly concave.Axial sculpture consisting of opisthocline ribs, subequal to broader than intercoastal spaces, extending on spire whorls from suture to abapical side of sub-sutural ramp, prominent on earlier whorls but becoming much weaker on last whorl.15 axial ribs on penultimate whorl, becoming obsolete on last whorl.Spiral sculpture consisting of widely-spaced, narrow, moderately prominent spiral cords, plicate where they cross axial ribs.First three teleoconch whorls with two cords between sub-sutural ramp and abapical suture, increasing to three cords on following three whorls.Last whorl with 13-15 spiral cords on base and rostrum.Intercostal spaces sculptured by numerous very fine spiral threads (up to about 13) (Fig. 5R-S).Aperture rather narrow, lanceolate, constricted at entrance to siphonal canal.Siphonal canal long for genus, not notched terminally.Inner lip with moderately thin callus, parietal region with a pad formed by termination of outer lip.Outer lip preceded by a wide, prominent, rounded varix; edge of lip sharp.Anal sinus moderately deep, asymmetrically U-shaped, with constricted opening.Stromboid notch moderately deep.Protoconch narrowly conical, of three smooth whorls (Fig. 5T-U), diameter 0.80 mm.Color yellowish-white tinged, with light brown on sub-sutural ramp, along peripheral cord and at level of abapical suture.

Remarks
This species is characterized by its large size, wide sutural ramp and well developed sculpture on the abapical side of each whorl.It is very similar to an undescribed species occurring in the Philippines and represented in the available material by a single large (17.9 × 6.3 mm), but damaged specimen from the Solomon Islands [SALOMON 1, Stn DW1768], which will be described by other authors elsewhere.The main difference is that in O. philippinensis sp.nov.there are two peripheral spiral cords on the earliest three whorls, increasing to 3 on later whorls; one very weak spiral cord is also present at the level of the abapical suture of the later whorls.In the undescribed species there are two main peripheral spiral cords bordered by a third, weaker cord on the abapical part of the sutural ramp and one (earlier whorls) increasing to 2 cords (later two whorls) at the level of the abapical suture.A further distinguishing feature is represented by its color pattern.Otitoma philippinensis sp.nov. is yellowish-white, tinged light brown on the sutural ramp, along the peripheral cord and at the level of the abapical suture, while the undescribed species is more vividly patterned with two well defined and intense brown bands.

Description
Shell fusiform-cylindrical (b/l 0.37-0.40;a/l 0.50-0.52).Teleoconch of up to 5¾ weakly convex whorls.Suture distinct, bordered by a moderately prominent sutural fold, followed by a moderately concave sutural ramp.Axial sculpture consisting of opisthocline axial ribs, rather straight and rounded, equal to or wider than intervals, developing mainly in peripheral region, gradually disappearing on abapical portion of sutural ramp.13-14 axial ribs on penultimate whorl, 11-12 on last whorl.Spiral sculpture consisting of widely-spaced cords, not forming distinct nodules at their intersection with axial ribs.First two teleoconch whorls with 3 spiral cords between sutural ramp and whorl base, increasing to 5-6 on later two whorls.Last whorl with 12 spiral cords on base and rostrum.Interspaces between spiral cords covered by microscopic spiral threads, 4-7 between spiral cords on later two teleoconch whorls (Fig. 6C-D).Aperture narrowly elliptical, constricted at entrance to siphonal canal.Siphonal canal short, slightly expanded at its end, and somewhat obliquely truncate.Outer lip edge sharp, preceded by a wide, prominent rounded varix.Stromboid notch strong.Anal sinus rather deep, apically rounded, its entrance strongly constricted by parietal pad.Protoconch narrowly conical, of 3¼ whorls, smooth except for presence of sparse microgranules, diameter 0.82-0.84mm (Fig. 6E-H).Color yellowish-white with yellowish-orange spiral cords.

Remarks
This species differs from its congeners by the combination of a narrowly conical protoconch, strong axial sculpture and well-developed secondary spiral sculpture.It may superficially resemble some specimens of O. cyclophora, from which it differs mainly in having broader axial ribs, a less constricted entrance to the anal sinus, and different protoconch features (narrowly conical, of 3¼ whorls rather than bluntly papillose, of about 1¾ whorls; see Kilburn 2004: 268).

Description
Shell cylindric-pupoid (b/l 0.39-0.40;a/l 0.55-0.59).Teleoconch consisting of 2¾-3½ whorls which are weakly and evenly convex.Last whorl with a shallowly excavated base.Whorls separated by a shallow suture, bordered by 2 closely spaced spiral cords, but lacking a distinct sub-sutural fold.Subsutural ramp shallow and concave.First teleoconch whorl with 2 closely spaced cords at suture, 1 weak cord on sutural ramp and 3 spiral cords on abapical part of whorl.On following whorl additional cords develop on sutural ramp, including 2-3 spirals more prominent than others, with 8-9 spiral cords below sutural ramp, increasing to 14-15 on last whorl.Spiral cords closely spaced, somewhat irregular in strength, rendered plicate by dense, moderately coarse incremental growth lines covering entire surface (Fig. 6L).Base sculptured by 14-15 spiral cords with 1 fine interstitial thread between.Axial sculpture restricted to penultimate whorl, consisting of irregularly spaced, opisthocline axial ribs, narrower than intercostal spaces, extending from suture to suture, but becoming narrower and arcuate on sutural ramp (about 13 on penultimate whorl in holotype).Aperture narrow and almost linear, but constricted at entrance to siphonal canal.Siphonal canal wide, expanding slightly distally, not notched.Inner lip covered with a thin callus and weak parietal pad.Outer lip preceded by a wide, low and rounded varix, its edge sharp, with a relatively shallow (for genus) stromboid notch.Anal sinus moderately deep, rounded, asymmetrically U-shaped, its entrance only slightly constricted by parietal pad.Protoconch domed, of 2½-2¾ whorls, smooth (Fig. 6M-N), diameter 0.89-0.90mm.Color white or white with three light brownish orange bands (on sutural ramp, on base of last and penultimate whorl and on base) and aperture edged with brownish orange.

Remarks
Otitoma nereidum sp.nov. is readily distinguished from its congeners on the basis of a combination of features including protoconch structure (paucispiral and domed), sculpture of the numerous axial ribs incised by strong spiral cords, and the relatively long siphonal canal.

Discussion
According to the WoRMS register of marine species the genus Otitoma Jousseaume, 1898 is represented by seven Recent species while Thelecytharella Shuto, 1969, retained as a full genus, includes 11 species consisting of nine Recent and two Pleistocene species.According to Wiedrick (2014), the genus Thelecytharella differs from Otitoma in having a broadly domed instead of bulbous protoconch, with a weak parietal callus pad and "open" anal sinus rather than restricted as in Otitoma.Although described by Kilburn (1995) as "bluntly conical, of about 3 whorls", species belonging to Thelecytharella show a remarkable variability in protoconch shape, varying from multispiral and bluntly conical (as in Otitoma oneili Barnard, 1958) to paucispiral and bulbous (Otitoma timorensis Schepman, 1913) and the stated difference is therefore very doubful.Similarly, the taxonomic value of anal sinus structure in supraspecific classification is uncertain.In the Pseudomelatomidae genus Inquisitor Hedley, 1918, for example, the anal sinus varies from U-shaped with constricted entrance to openly U-shaped with diverging sides (Kilburn 1988: 258-259).We, therefore, prefer to follow Kilburn (2004) in considering Thelecytharella as a synonym of Otitoma.However, Kantor et al. (2008) recently pointed out that shell features of members of Turridae are useful for species-level identification but are inadequate for supraspecific classification.For this reason, the status of Thelecytharella as a synonym of Otitoma will remain uncertain until anatomical and/or molecular features of species become known.Otitoma (=Thelecytharella) is a rather variable assortment of species including several different morphological subgroups indicative of remarkable diversification, but the dividing line between groups is too nebulous to permit recognition of separate genera or even subgenera.Agladrillia oyamai Shuto, 1965, type species of Thelecytharella, is characterized by a multispiral, bluntly conical protoconch of three whorls, and a teleoconch with predominant axial sculpture consisting of numerous weaker spiral cords.Similar features occur in O. oneili (Barnard, 1958) and O. metuloides (Kilburn, 1995), while O. neocaledonica sp.nov.differs in protoconch structure but otherwise conforms in sculpture to this group.Shuto (1970) proposed the genus Lioglyphostomella for the two Recent species Otitoma kwandangensis and O. timorensis, both described by Schepman (1913), which differ from the "typical" group in having stronger, more prominent axial ribs crossed by few spiral cords forming low nodules where crossing the axials (Shuto 1970: 165).Otitoma sororcula sp.nov., the third member of this small species-group, is very similar in teleoconch features to D. kwandangensis from which it differs in protoconch structure.The generic name Lioglyphostomella was considered a synonym of Thelecytharella by Kilburn (1995), while Bouchet et al. (2011: 293) retained it as a valid genus, but without a discussion.
European Journal of Taxonomy 304: 1-30 (2017) A further morphological subgroup includes species which resemble Austropusilla (Metaclathurella) crokerensis Shuto, 1983, namely the Recent Otitoma batjanensis (Schepman, 1913) and O. xantholineata sp.nov.These species have a predominantly spiral sculpture differing from Agladrillia oyamai Shuto, 1965 in lacking axial sculpture or in retaining vestigial ribs on earlier teleoconch whorls (O.xantholineata sp.nov).Austropusilla (M.) crokerensis Shuto, 1983 andO. batjanensis (Schepman, 1913) have a large conical protoconch of three whorls similar to that occurring in Agladrillia oyamai, while in O. xantholineata sp.nov. the protoconch is somewhat bulbous, of 1½-1¾ whorls.Some large size Otitoma species (up to 19 mm in length), namely O. tropispira sp.nov., O. boucheti sp.nov.and O. kagoshimaensis (Shuto, 1965), a Pleistocene species from Japan originally assigned to Turridrupa, have prominent keel-like spiral cords, somewhat reminiscent of those in the pseudomelatomid genus Turridrupa Hedley, 1918.However, members of the latter genus bear spiral plicae in the interior part of the aperture and have the last half to two protoconch whorls sculptured by axial riblets (Kilburn 1988: 235;Powell 1966: 54).Sysoev (1997: 344) noted that O. vitrea (Reeve, 1845) and O. kecil (Sysoev, 1997) (both as Thelecytharella) "form a distinct group characterized by a very small (5.5 mm high) pupoid shell without axial sculpture" and suggested the possibility that these species may represent a separate genus.A further feature of these species is that the teleoconch consists of only a few whorls, usually three, while in the "typical" A. oyamai and O. oneili there are up to 6-7 teleoconch whorls.Otitoma philpoppei sp.nov. is morphologically similar in shape and protoconch features to O. vitrea, but has distinct axial ribs on the first teleoconch whorl, obscuring the limit of this subgroup.
Species assigned to the genus Otitoma show a remarkable variability in spiral sculpture.For example, in the morphologically related O. kecil, O. vitrea and O. philpoppei sp.nov., the spiral sculpture is rather weak and there is no trace of secondary sculpture.Other species have a developed primary spiral sculpture but lack secondary threads between cords (examples are exhibited by Otitoma metuloides, O. sororcula sp.nov., O. kwandangensis and O. xantholineata sp.nov.).On the contrary, O. cyclophora and the majority of species have a distinct secondary sculpture (examples are O. boucheti sp.nov., O. carnicolor, O. crassivaricosa sp. nov., O. elegans sp. nov., O. philippinensis sp. nov., O. timorensis and O. tropispira sp. nov.).In this latter "group", the spiral threads may appear almost linear (O.philippinensis sp.nov., O. tropispira sp.nov.) or somewhat plicate where they cross axial incremental lines (Figs 3P,5S).Otitoma neocaledonica sp.nov.has relatively weak spiral interstitial threads, but a well-developed sculpture of microscopic granules (Fig. 4K).Secondary sculptural features seem poorly correlated with subgroups recognized on the basis of other characters.There are numerous cases of morphologically similar species differing in absence/presence of secondary spiral threads (O.xantholineata sp.nov.vs O. kagoshimaensis, O. kwandangensis vs O. timorensis and O. metuloides vs O. oneili); therefore, this feature should only be considered of specific value.(1971) proposed the genus Maesiella with Maesiella maesae McLean & Poorman, 1971 from Pacific Mexico selected as the type species.While discussing the taxon, McLean noted that "Thelecytharella has the appearance of a crassispirine genus related to Maesiella but having a broadly open rather than constricted sinus" (McLean 1971: 123).The four species currently accepted as belonging to Maesiella (see WoRMS Register), have a reported distribution from Mexico and Panama to Venezuela, are morphologically very similar to Otitoma (= Thelecytharella) as presently construed, and the stated difference in anal sinus structure is, as previously remarked, of uncertain value.We have not examined material of Maesiella and therefore cannot provide a more objective evaluation; however, in our opinion further studies will probably demonstrate that the two taxa are synonymous.

MORASSI M. et al., New species of the genus Otitoma
From the data available in the literature, the majority of species belonging to the genus Otitoma (= Thelecytharella) are from the Indo-Pacific region, particularly the central-western (Indonesia and Philippines) and south-western (New Caledonia and Lifou) to the central Pacific (Howland Island, Cook Islands).Of the twelve species here described, four are presently reported from New Caledonia (O. hadra sp. nov., O. neocaledonica sp. nov., O. tropispira sp. nov. and O. rubiginostoma sp. nov.), while O. carnicolor (Hervier, 1896) occurs in the vicinity of Lifou Island.Six species, including five here described, are from Fiji Islands (O. astrolabensis, O. boucheti sp. nov., O. elegans sp. nov., O. nereidum sp. nov., O. sororcula sp. nov. and O. xantholineata sp. nov.) (Kilburn, 1986).However, O. oneili (Barnard, 1958) actually has a much broader distribution, occurring also from the Philippines to the Solomon Islands (pers.obs.).Perplexingly, O. cyclophora Deshayes, 1863, type species of Otitoma, is the sole Otitoma species presently reported from the Red Sea (Dekker & Orlin 2000) and the northern Indian Ocean; it seems probable that more intense sampling in these still poorly explored areas will yield recognition of additional undescribed species.
Abbreviations: a = aperture length a/l = ratio of aperture length to total shell length b = shell width b/l = ratio of shell width to total length l = total shell length MNHN = Muséum national d'Histoire naturelle, Paris, France MZB = Museo di Zoologia dell'Università di Bologna, Bologna, Italy o.d.= original designation pers.obs.= personal observation spm(s) = specimen(s) Stn = station USNM = National Museum of Natural History, Washington DC, USA ZMA = Naturalis Biodiversity Center, Leiden, the Netherlands
, while in O. xantholineata sp.nov.axial sculpture is absent or restricted to earlier teleoconch whorls.In O. kagoshimaensis the interspaces between spiral cords are "covered by numerous minute spiral lines", while O. xantholineata sp.nov.has no interstitial sculpture.Furthermore, O. xantholineata sp.nov.has . Otitoma philpoppei sp.nov.occurs from the Philippines to Fiji, while O. philippinensis sp.nov., O. batjanensis, O. kecil, O. kwandangensis, O. timorensis and O. vitrea are reported from the Philippines.Otitoma crokerensis is only known from the Arafura Sea.Otitoma crassivaricosa sp.nov.has a wide Pacific distribution from Fiji to Hiva Oa, Marquesas Archipelago.At present only three taxa are reported from Southern Africa and Mozambique, namely O. oneili, O. metuloides and O. cyclophora (= O. mitra)