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A revision of the chilodontidae (Gastropoda: Vetigastropoda: Seguenzioidea) of Southern Africa and the south-western Indian ocean.

Vaceuchelus semilugubris (Deshayes, 1863) Figs 4L, 50, 51

Turbo semilugubris: Deshayes 1863: 72, pl. 9 [36], figs 9, 10; Martens 1880: 294; Herbert 1996: figs 70, 71 (syntype). Type loc.: Reunion.

Leptothyra semilugubris: Pilsbry 1888: 252, pl. 58, fig. 52a; Viader 1937: 55.

Vaceuchelus semilugubris: Poppe et al. 2006: 47.


Shell: Small, turbiniform, robust; length more or less equal to diameter (L/D=0.92-1.04); teleoconch of approx. 3.5 whorls, but apex usually badly eroded; sculpture cancellate; first whorl strongly shouldered at mid-whorl, almost horizontal above shoulder and vertical below it; shoulder develops into a strong spiral cord at end of whorl, another cord evident at level of abapical suture; axial pliculae do not develop until near end of whorl; this sculpture strengthening during second whorl with an additional cord arising between first cord and adapical suture; these two cords becoming the large peripheral and supra-peripheral cords of last adult whorl; 1-2 additional secondary cords arising above supra-peripheral cord during third whorl; axial pliculae stronger on second and subsequent whorls, prosocline and raised into granules where they cross spiral cords, granules somewhat angular on second whorl, but becoming progressively more rounded and bead-like on subsequent whorls; interstices quadrate; last adult whorl with 25-30 pliculae. Base with 5 spiral cords, the first level with suture, the fifth small and spiralling into umbilicus; umbilicus narrowly patent even in adults. Peristome oblique and nearly tangential, aperture subcircular; columella shallowly concave with a rounded tooth at its base; interior of outer lip of mature individuals subterminally thickened and set with approx. 10 ridge-like denticles, with an additional row of smaller granules just inside lip edge; apertural dentition absent in immature specimens, in these the outer lip is strongly notched at ends of spiral cords; interior nacreous, weakly angled beneath external cords, but labral denticles not extending into aperture as in-running ridges.

Microsculpture (Fig. 51A, B): Juvenile microsculpture comprising irregular vermiform spiral threads; subsequent microsculpture mostly granular, but with traces of scratchlike markings.

Protoconch (Fig. 51C): White, slightly exsert, eroded in most material; diameter ca 285 [micro]m; strongly sculptured with an irregular superficial reticulation showing traces of axial alignment, but no spiral element. Terminal lip more or less straight, but perhaps evincing signs of a broken beak-like projection; becoming concave toward abapical suture.

Colour: Shell white, variously patterned with black spots, blotches or axial bands, orientation of pattern prosocline in some specimens, opisthocline in others; markings generally finer on base; pattern fading to brown in old material. Occasional specimens with almost no black markings (Fig. 50F). Coloration of fresh specimens somewhat obscured by dirty buff intritacalx deposit, superficial calcareous encrustations and debris.

Dimensions: Largest specimen, length 3.55 mm, diameter 3.40 mm; smallest specimen with mature apertural dentition, length 2.6 mm, diameter 2.7 mm.

Operculum (Fig. 4L): Relatively tightly multispiral throughout.

Radula (Fig. 51D, E): Formula [infinity]+3+1+3+[infinity]; ca 50 transverse rows of teeth; transition from lateral to marginal series clear. Rachidian with broad, trigonal cusp, well-developed hood, and a distinct transverse basal ridge; cutting edge with a slender, acuminate central denticle and 2 or 3 progressively smaller, similarly shaped denticles on each side. Lateral teeth all of similar size, their cusps with coarse lateral denticles on both margins (3-5), those on the third lateral particularly strong. Marginals closely resembling those of Herpetopoma scabriusculum, the cusps of the inner ones with a strongly pectinate outer margin; outermost marginals shorter and with a slightly dilated, fringed cusp.

External anatomy: Only dried material available; evidently chilodontid but insufficient detail evident.

Type material: Two syntypes of Turbo semilugubris Deshayes, 1863, in MNHN, the one in better condition is here figured and designated lectotype (MNHN 24658) (Fig. 50A, B), length 3.10 mm, diameter, 3.05 mm.

Other material examined: REUNION: not further localised (M. Jay coll'n, MNHN; NMSA G4264, J269); off Cap La Houssaye (21.01797[degrees]S 55.23709[degrees]E), -12 m, hand-dredged sand, J. Drivas, i.1988 (NMSA K3035).

Distribution and habitat: Known only from the island of Reunion; Jay (2009) recorded it as living under stones at -10-15 m, but noted it to be rare.

Remarks: There appear to have been no published records of this species since its original description. Subsequent mentions of the name are either simple listings (Martens 1880; Viader 1937) or mere translation of the original description (Pilsbry 1888), and there is no evidence of new material having been examined. However, additional samples clearly referable to this species have been found on Reunion, confirming that the original provenance was correct. The occurrence of the species in Mauritius requires confirmation. Pilsbry (1888) and Viader (1937) mentioned this island, but it is not clear whether this was a generalisation or specifically intended to mean Mauritius rather than Reunion. Despite extensive searching for micro-molluscs in beach-drift samples and hand-dredged sand from near-shore reefs on Mauritius (by Kilburn and Herbert), no specimens of V. semilugubris have been collected. Nonetheless, it is clearly not common, even on Reunion. Given the proximity of Reunion and Mauritius, it would be surprising if the species did not occur on both islands.

Whether this species should be referred to Vaceuchelus or Herpetopoma requires further study. I have chosen to refer it to Vaceuchelus on account of the shape of the shell and its relatively coarse, cancellate sculpture. The presence of denticles inside the outer lip is a feature common to both Vaceuchelus and Herpetopoma. However, in the present species there is also a distinct denticle at the base of the columella that delineates a notch (albeit weak) at the junction of the basal and columella lips. This is a feature more typical of Herpetopoma.

The species is distinctive amongst small chilodontids of the south-western Indian Ocean on account of its bold, axial colour bands and relatively wide umbilicus. In fresh shells these bands are very dark purplish brown to almost black, but fade to a paler maroon-brown in old and sun-bleached specimens. They are clearly part of the shell, unlike the greyish axial bands seen in fresh material of other species (e.g. V. natalensis), which are part of the superficial intritacalx deposit. Rarely, specimens are almost devoid of black markings and may easily be confused with the sympatric V. jayorum (above). That species, however, has strong close-set axial pliculae on the first teleoconch whorl, has intermediary spiral cords on the last quarter of the final whorl, and does not develop labral dentition when mature. I suspect V. semilugubris may exhibit ecological variation in shell morphology, ranging from typical boldly marked specimens with coarse sculpture, to somewhat smaller specimens with little or no colour pattern and finer sculpture (Fig. 50F). However, since accurate information on habitat is not available, this remains a speculative observation.

Genus Perrinia H. & A. Adams, 1854

Perrinia: H. & A. Adams 1854 in 1853-54: 419. Type species: Monodonta angulifera A. Adams, 1853, by subsequent designation (Pilsbry 1890 in 1889-90: 15).

Shells of Perrinia species are small (length <20 mm) with relatively flat-sided whorls, resulting in a trochiform spire. The suture is frequently strongly indented and the periphery often angular and/or keeled. The apical whorls are sculptured only with axial pliculae and subsequent whorls with both spiral cords and axial pliculae that interact to produce a cancellate or foveolate sculpture (not obviously beaded). The columella has a single, relatively weak tooth or knob near its base, there is no inductural callus shield, the umbilicus is generally closed, and the interior of the outer lip bears spiral ridges that extend deep into the aperture. The juvenile microsculpture is granular and lacks vermiform spiral threads, and the adult microsculpture of close-set, scratch-like, axial marks is well developed.

Perrinia has long been regarded as a subgenus of Turcica H. & A. Adams, 1854 (type species Turcica monilifera A. Adams, 1854, by monotypy), but I consider Perrinia species to form a relatively well defined group which can be reasonably easily separated from the much larger Turcica s.s. species on account of their smaller size, stouter shells and strong spiral lirae inside the outer lip of the aperture. In the absence of data suggesting otherwise, I therefore accord Perrinia full generic status.

It should be noted that Kano et al. (2009) have shown that Turcica belongs within the Calliotropidae. This may also prove to be the case for Perrinia when sequence data become available, but I consider the overall facies of the shell of Perrinia to be closer to that of the Chilodontidae and thus maintain it here for the present.

Key to species of Perrinia in the south-western Indian Ocean

1 Shell with broad, low, opisthocline axial ribs angulifera

--Shell lacking such ribs, axial sculpture restricted to relatively narrow, prosocline pliculae

2 Shell small (length up to 6.0 mm); peripheral spiral cord at most weakly stellate konos

--Shell larger (length up to 11.0 mm); peripheral spiral cord strongly stellate stellata

Perrinia angulifera (A. Adams, 1853) Figs 4M, 5B, 6E, 52-54

Monodonta angulifera: A. Adams 1853: 176; Pilsbry 1890 in 1889-90: 416. Type loc.: Puerto Galero, Mindoro Is., Philippines, sandy mud, 6 fath. [-11 m] (Mus. Cuming).

Tectaria montrouzieri: Fischer 1878: 212-213; Souverbie & Montrouzier 1879: 31, pl. 3, fig. 6; Herbert 1996: 418, figs 18, 19 [= M. angulifera A. Adams, 1853]. Type loc.: Ile Art (New Caledonia).

Perrinia angulifera: Smith 1903: 618, pl. 35, fig. 27; Hylleberg & Kilburn 2003: 26; Heros et al. 2007: 209.

Calliostoma anguliferum: Hidalgo 1904-5: 255.

Canthariduspliciferus: Schepman 1908: 43, pl. 3, fig. 3. Type loc.: Siboga st'n 109, Pulu Tongkil, Sulu Archipelago, -13 m, Lithothamnion bottom. Syn. n.

Perriniaplicifera: Poppe et al. 2006: 45, pl. 15, fig. 7; Poppe & Tagaro 2008: 180, pl. 35, fig. 5.

Turcica montrouzieri: Hedley 1909: 354; Jansen 1996: 10, No 31.

Turcica (Perrinia) angulifera: Melvill 1928: 98.

Turcica concinna [non A. Adams, 1863]: Jay 2009.


Shell: Elevated trochiform (L/D=1.24-1.5); apical angle 50-60[degrees]; teleoconch of up to 7.5 whorls (apical ones frequently badly eroded or missing); apex truncated; spire whorls somewhat flat-sided, but still retaining a degree of curvature; periphery angular, but not keeled; suture inserted below peripheral angle, level with subperipheral cord creating a narrow channel. First teleoconch whorl sculptured initially only with axial pliculae ([+ or -] 30); 3-4 spiral cords develop during second whorl; cords crossed by crispate axial pliculae; third and subsequent whorls with 5-6 broad, rather uneven spiral cords above and including peripheral one; subsutural cord rendered coronate by well-developed triangular nodules arising from approximately alternate axial pliculae; nodules frequently apically bifid on later whorls; interaction of spiral and axial elements producing a foveolate sculpture with rectangular to D-shaped pits; shell also sculptured by broad, opisthocline (rarely orthocline) ribs, stronger in some specimens than others, 11-15 on last adult whorl; ribs for the most part arising from subsutural coronations (not invariably), strongest at periphery and rendering shell circumference undulant or even weakly stellate (infrequent). Base sculptured by 6-8 spiral cords; cords near umbilicus weaker than those toward periphery; axial pliculae evident in cord intervals, especially that between peripheral and subperipheral cords, weakest and almost obsolete near umbilicus; pliculae interact with cords causing some weak granulation of the latter; umbilicus closed in adults, but patent in very young individuals. Peristome oblique; aperture D-shaped, flattened at parietal and columella lips; columella (at maturity) with a relatively prominent tooth approximately one third of length from basal lip, occasionally also with a very low swelling apical to this; interior of outer lip with up to 16 spiral lirae running into aperture (only in mature shells), those nearest shell axis may terminate on base of columella and appear as small denticles below columella tooth; outer lip prosocline, its margin thin, but becoming thicker internally.

Microsculpture (Fig. 53B, C): Juvenile shell with finely granular microsculpture, vermiform spiral threads not evident; adult shell with well-developed scratch-like microsculpture, which is filled with intritacalx deposit in fresh shells.

Protoconch (Fig. 53A, C): Translucent white; diameter ca 260 um; not projecting above first teleoconch whorl, shell apex thus appearing truncated and somewhat tilted; missing or badly eroded in most specimens; surface sculptured with a fine, irregular granulation; terminal lip strongly angled above mid-whorl.

Colour: Ground colour generally greyish white to pale buff; basal spiral cords commonly marked with brown flecks; some specimens with a reddish or brown spiral band below suture and another at periphery; entire surface covered with off-white, chalky intritacalx deposit. One specimen almost entirely brownish grey, with a dark, ash-grey intritacalx. Most shells encrusted to some degree with other organisms, frequently coralline algae and bryozoans.

Dimensions: Largest NMSA specimen (E4265), length 16.0 mm, diameter 11.6 mm.

Operculum (Fig. 4M): Initially tightly multispiral, but whorls broadening with growth and becoming more openly multispiral.

Radula (Fig. 53D-F): Formula [infinity]+3+1+3+[infinity], with ca 60 transverse rows of teeth; lateral flanges of rachidian well developed creating a distinct hood, cusp with a strong transverse basal ridge, the apex broadly trigonal with stout denticles, a larger, lanceolate median one and 2 or 3 smaller ones on each side. Laterals overlapping extensively, their cusps trigonal and noticeably asymmetrical; central denticle largest, lateral denticles well developed on outer margin, progressively decreasing in size toward tooth shaft; inner margin with fewer denticles. Marginals numerous and slender, cusps of inner ones recurved and coarsely pectinate, outer ones spathulate distally with a feathered edge.

External anatomy (Figs 5B, 6E): Body white with pale brownish maculations on sides of foot and underside of epipodium, snout heavily marked (transversely) with dark brown to black. Snout expanded laterally, but not strongly so; free margin of cephalic lappets relatively coarsely divided (2-3 digits); right post-ocular peduncle well developed, arising beside base of eyestalk, somewhat flattened and with a distinct dorsal groove; right subocular tentacle not evident; left neck lobe with numerous tentacles of varying size, right lobe with finely fimbriate margin and approx. 3 tentacles in anterior half; 6 or 7 large epipodial tentacles on each side, with frequent smaller intermediary tentacles of varying size; an indistinct epipodial sense organ present at base of larger epipodial tentacles, none evident under neck lobes.

Type material: Three syntypes of M. angulifera A. Adams, 1853, in NHMUK (1968215), the largest is here figured and designated lectotype (Fig. 52A), length 12.9 mm, diameter 8.9 mm. Holotype of C. pliciferus Schepman, 1908 (Fig. 52C), in ZMAN (3.08.028). Holotype of T. montrouzieri Fischer, 1878 (Fig. 52B), in MHNB (Herbert 1996) and there is an 'ex auctore' specimen in MNHN, but this has no type status.

Regional material examined (all NMSA unless indicated otherwise): KENYA: Shimoni (4.6482[degrees]S 39.3814[degrees]E), dredge 1+2, J.D. Taylor (NHMUK). MAURITIUS: off Tombeau Bay (20.1017[degrees]S 57.5025[degrees]E), -25-25 m, CSIR Water Research (L2724); REUNION: off Baie de St-Paul, Marion-Dufresne 32, st'n DC85 (21.00[degrees]S 55.25[degrees]E), -58-70 m, dredged, 1982 (MNHN); off St-Gilles-les-Bains, Marion-Dufresne 32, st'n DC56 (21.083[degrees]S 55.200[degrees]E), -170-225 m, dredged, 1982 (MNHN); off Souris Chaude (21.383[degrees]S 55.667[degrees]E), -65 m, hand-dredged sand, J. Drivas, 1993 (L548); Reunion, not further localized (M. Jay coll'n, MNHN). RODRIGUES: 160 miles south (21.350[degrees]S 65.867[degrees]E), Anton Bruun 2, st'n 124F (USNM 716607). MOZAMBIQUE: Nacala area, Fernao Veloso Bay (14.4312[degrees]S 40.7003[degrees]E), -8-10 m, x.1998, C. Fernandes (J. Rosado coll'n); 50 miles SE of Beira (20.5000[degrees]S 35.7167[degrees]E), -62 m, Anton Bruun 8, St'n 400C, IIOE (USNM 718524); off Lacerda lighthouse, 50 km north of Maputo (25.56167[degrees]S 32.84472[degrees]E), -50-56 m, dredged J. Rosado, v.2010 (J Rosado coll'n); off Ponta Techobanine (26.68132[degrees]S 32.95093[degrees]e), -68-75 m, dredged J. Rosado, xii.2005 (J. Rosado coll'n). SOUTH AFRICA: KwaZulu-Natal: off Kosi Bay (26.8916[degrees]S 32.9266[degrees]E), -51 m, sand, stones, large algae, dredged NMDP, RV Sardinops, st'n ZA48, (S8959); SE of Kosi River mouth (26.9100[degrees]S 32.9217[degrees]E), living, -50 m, medium sand, algae, dredged NMDP, RV Meiring Naude, st'n ZA9, (D6180, D6182); ditto (26.9167[degrees]S 32.9300[degrees]E), -65 m, sponge, gorgonians, medium sand, dredged NMDP, RV Meiring Naude, st'n ZA12, (D8183); ditto (26.9217[degrees]S 32.9233[degrees]E), living, -50 m, medium sand, rubble, dredged NMDP, RV Meiring Naude, st'n ZA11, (D8960); off Boteler Point (27.00[degrees]S 32.92[degrees]E), -70 m, coral rubble, dredged NMDP, RV Meiring Naude, st'n ZB5, (D6382, E1762); ditto (27.013[degrees]S 32.905[degrees]E), living, -50 m, dead coral rubble, lithothamnion, dredged NMDP, RV Meiring Naude, st'n ZB2, (D9209); ditto (27.013[degrees]S 32.918[degrees]E), -70 m, some coarse sand, some shell rubble, dredged NMDP, RV Meiring Naude, st'n ZB4, (D7412); ditto (27.0183[degrees]S 32.9200[degrees]E), -78 m, coarse sand, dredged NMDP, RV Meiring Naude, st'n ZB6, (D7476); NE of Dog Point (27.0800[degrees]S 32.8867[degrees]E), -56-57 m, sand, lithothamnion pebbles, dredged NMDP, RV Sardinops, st'n ZC8, (S5072); ditto (27.08000[degrees]S 32.89167[degrees]E), living, -65 m, sand, lithothamnion pebbles, dredged NMDP, RV Sardinops, st'n ZC9, (S7597); SE of Rocktail Bay (27.2017[degrees]S 32.8300[degrees]E), living, -60 m, coarse sand, dredged NMDP, RV Sardinops, st'n ZD9, (S5250); NE of Liefeldt's Rocks (27.7167[degrees]S 32.6650[degrees]E), -50 m, lithothamnion, medium sand, dead coral rubble, dredged NMDP, RV Meiring Naude, st'n ZJ6, (E4265); ditto (27.72000[degrees]S 32.66167[degrees]E), living, -50 m, lithothamnion, stones, some coarse sand, dredged NMDP, RV Meiring Naude, st'n ZJ1, (E4344); ditto (27.7230[degrees]S 32.6633[degrees]E), -50 m, medium sand with some stones, dredged NMDP, RV Meiring Naude, st'n ZJ5, (E3417).

Other material examined: ARABIAN SEA: Gulf of Oman, Townsend (NMGW). ANDAMAN ISLANDS: Port Blair, Winckworth coll'n (NHMUK). NEW GUINEA: off west side of Lolorua Is., SW of Port Moresby, -13-18 m, Ponder & Colman (AMS); New Britain, J. Brazier (AMS C11849). AUSTRALIA: Flinders Passage, 7(?) fath. [-13 m] Challenger (NHMUK); Gulf of Carpentaria, 10 mls SW of Mapoon, -128 m (AMS); off Murray Is., Torres Strait, -9-15 m, C. Hedley (AMS); east of Banks Is., Torres Strait, -18 m, BMR st'n 522 (AMS); Cape York Peninsula, Albany Passage, 4-14 fath. [-7-26 m], C. Hedley (AMS); V2 ml west of North Direction Is., -36.5 m, Great Barrier Reef Exped'n, dredged, st'n 16, 1929 (AMS); 2 mls NE of west side of Gillet Cay, Swains Reef, southern Great Barrier Reef, 30-40 fath. [-55-73 m] (AMS). NEW CALEDONIA (all ORSTOM, MNHN): Secteur de Poum, st'n 1027 (20[degrees]03'S 163[degrees]51'E), -29 m; secteur de Poum, st'n 1017 (20[degrees]08'S 163[degrees]51'E), -21 m; secteur de Noumea, st'n 56 (22[degrees]10'S 166[degrees]15'E), -11 m; secteur de Noumea, st'n 285 (22[degrees]24'S 166[degrees]26'E), living, -19 m; secteur de Noumea, st'n 271 (22[degrees]15'S 166[degrees]21'E), -22 m; secteur de Koumac, st'n 942 (20[degrees]37'S 164[degrees]13'E), living, -15 m.

Distribution and habitat (Fig. 54): Indo-West Pacific; from SE Asia and northern Australia, to the Andaman Islands, Maldive archipelago (Smith 1903), Mascarene Islands and the continental margin of the western Indian Ocean, from Muscat (Melvill 1928) south to northern South Africa. Off Zululand, this species was dredged, in relative abundance, amongst old coral rubble and pebbles encrusted with Lithothamnion growths lying on a substratum of coarse bioclastic sand, at depths of -50-80 m (living specimens -50-65 m). Cantharidus pliciferus was also found on a substratum described as a 'Lithothamnion bottom' (Schepman 1908). In fully tropical areas the bathymetric range evidently extends into shallower water (to -10 m). Deep-water material from Reunion (-170-225 m) comprised only long-dead specimens that probably originated in shallower habitats on the island's steeply shelving coast.

Remarks: South-western Indian Ocean examples of this species generally have fewer (11-15), broader axial ribs than is typical (18 on last adult whorl in NHMUK types) and the axial pliculae between the spirals cords are less close-set. However, in other respects they are indistinguishable and I have little hesitation in referring them to Adams' species. Such small differences are not unexpected in material from such widely separated localities. Canthariduspliciferus Schepman, 1908 from the Sulu Archipelago is simply a subadult specimen of the present species and Tectaria montrouzieri Fischer, 1878 from New Caledonia, is a very typical adult (Herbert 1996), albeit rather small.

The broad, opisthocline ribs of this species set it apart from all others of the genus, even those of similar size, e.g. P. chinensis (Sowerby, 1888) from Hong Kong (Fig. 71A, B), P. maculata (Brazier, 1877) from northern Australia (Fig. 71C, D) and P. elisa (Gould, 1849) from Singapore. Amongst local species, P. konos is very much smaller and P. stellata has a carinate and strongly stellate periphery.

Perrinia konos (Barnard, 1964) comb. n. Figs 4N, 55-57

Turcica konos: Barnard 1964: 20, fig. 2f; Kensley 1973: 44, fig. 111. Type loc.: off Umkomaas, KZN south coast, South Africa, 40 fath. [-73 m].


Shell: Small, elevated-trochiform to turriculate (L/D=1.26-1.50); apical angle 55-60[degrees]; teleoconch of 6-7 whorls; whorls flat-sided and peripherally angled, peripheral angle marked by a strong keel, base flattened; sutures channelled and inserted below peripheral angulation, spire thus rendered pagodaform; insertion of suture level with subperipheral spiral cord. First teleoconch whorl more or less evenly rounded and sculptured only with strong, evenly spaced, axial pliculae (14-20); peripheral cord (angulation) begins to develop during second whorl, axial pliculae become nodular at angulation; whorls more flat-sided from third whorl onward and with 2 or 3 further spiral cords, one immediately below suture and 1 or 2 between this and peripheral keel; subsutural cord with adapically pointed triangular nodules which project into sutural channel; nodules with a thin lamellate ridge running into suture; middle spiral cord(s) thinner than the other two and at most obsoletely granular where crossed by axial sculpture; peripheral cord becomes keel-like with growth and is rendered stellate in apical view by radiating triangular nodules, 13-20 on body; axial sculpture of prosocline pliculae persists throughout, but pliculae more numerous than either the subsutural or peripheral nodules; whorl surface rendered somewhat cancellate/foveolate by interaction of spiral and axial sculpture. Base with 4 (rarely 5) evenly spaced spiral cords, outer one weakly undulant, others progressively smoother toward columella; interval between peripheral and subperipheral cords deeply concave and forms suture channel in spire whorls; innermost basal spiral very close to columella; umbilicus patent in juveniles, but closed by thickened columella in adults. Aperture subcircular, somewhat flattened in parietal and columellar regions; columella (at maturity) with a bulge-like swelling at its base, rarely forming a distinct tooth; interior of outer lip with up to 11 spiral lirae running into aperture (evident only in fully mature shells), one where basal portion of lip joins columella not enlarged; outer lip prosocline, not obviously thickened; interior nacreous.

Microsculpture (Fig. 56B, C): Initial whorls lacking vermiform spiral microsculpture; later whorls with distinct scratch-like microsculpture.

Protoconch (Fig. 56A, C): White to translucent white, strongly exsert; diameter 260280 um; apical beak scarcely evident; surface evidently smooth; terminal lip roundly angled between mid-whorl and adapical suture.

Colour: Ground colour yellowish white to pale orange, commonly buff; frequently with spots and blotches of a deeper shade beneath suture and at periphery; basal cords often spotted with dark brown; occasional specimens almost entirely pink with deeper pink markings; intritacalx deposit generally cream-yellow, only visible in the freshest specimens; shell frequently encrusted with bryozoans, sponges, tubicolous worms and encrusting Foraminifera.

Dimensions: Attaining length 6.0 mm, diameter 4.2 mm in South Africa, but reaching length 6.2 mm, diameter 4.9 mm in northern Madagascar (MNHN).

Operculum (Fig. 4N): Initially tightly multispiral, but whorls broadening with growth and becoming more openly multispiral.

Radula: Formula [infinity]+3+1+3+[infinity], with ca 70 transverse rows of teeth; similar to that of P. angulifera in that the cusp of rachidian is strongly dentate, with a sharply pointed central denticle and several similar but smaller lateral denticles on each side; laterals with a sharply pointed central cusp; marginals very numerous and fine.

External anatomy: Little preserved material available, but evidently similar to that of P. angulifera.

Type material (four syntypes of Turcica konos Barnard, 1964, in SAMC): SOUTH AFRICA: KwaZulu-Natal: off Umkomaas, 40 fath. [-73 m] (A9257, 2 adult, 1 immature and 1 broken).

Material examined (all NMSA unless indicated otherwise): MADAGASCAR: West of Cap d'Ambre (12.133[degrees]S 48.933[degrees]E), -238-249 m, Campagne Miriky, dredged, st'n DW3196, (MNHN); west of Nosy Be (13.417[degrees]S 47.950[degrees]E), -71-158 m, Campagne Miriky, dredged, st'n DW3230, 3.vii.2009 (MNHN); in front of Baie Narendry (14.483[degrees]S 47.450[degrees]E), living, -48-139 m, Campagne Miriky, dredged, st'n DW3238, 6.vii.2009 (MNHN); Secteur de Manantenina (24.38333[degrees]S 47.53333[degrees]E), -154-168 m, Exped'n Atimo Vatae, dredged Nosy Be 11, st'n DW3522, 1.v.2010 (MNHN); ditto (24.38333[degrees]S 47.51667[degrees]E), -200-220 m, Exped'n Atimo Vatae, dredged Nosy Be 11, st'n DW3523, 1.v.2010 (MNHN); north of Sainte Luce (24.59833[degrees]S 47.53500[degrees]E), -80-86 m, Exped'n Atimo Vatae, dredged Nosy Be 11, st'n DW3530, 2.v.2010 (MNHN); ditto (24.6567[degrees]S 47.5283[degrees]E), -86-87 m, Exped'n Atimo Vatae, dredged Nosy Be 11, st'n DW3532, 2.v.2010 (MNHN); between Lokaro & Ste Luce (24.8450[degrees]S 47.4783[degrees]E), -99-101 m, Exped'n Atimo Vatae, dredged Nosy Be 11, st'n DW3518, 30.iv.2010 (MNHN); ditto (24.865[degrees]S 47.467[degrees]E), -80-83 m, Exped'n Atimo Vatae, dredged Nosy Be 11, st'n DW3519, 30.iv.2010 (MNHN); south east of Faux-Cap (25.78333[degrees]S 46.0333[degrees]E), -133-178 m, Exped'n Atimo Vatae, dredged Nosy Be 11, st'n CP3620, 15.v.2010 (MNHN). MOZAMBIQUE: NE of Maputo, -65 m, dredged J. Rosado, (D. Slater coll'n); Maputo transect, Campagne Mainbaza, RV Vizconde de Eza, dredged, st'n CP3130 (23.883[degrees]S 33.117[degrees]E), living, -112-127 m, 9.iv.2009 (MNHN); off Maputo, -55-100 m, dredged J. Rosado, 2008 (D. Slater coll'n); off Inhaca Is (26.020[degrees]S 33.066[degrees]E) -75-125 m, dredged J. Rosado, 2006 (D. Slater coll'n); off Ponta Techobanine (26.68132[degrees]S 32.95093[degrees]E), -60-115 m, dredged J. Rosado (D. Slater coll'n). SOUTH AFRICA: KwaZulu-Natal: off Kosi River Mouth (26.9100[degrees]S 32.9433[degrees]E), -75 m, coral rubble, sandstone, marine growths, dredged NMDP, RV Meiring Naude, st'n ZA13, (D8999); off Black Rock (27.165[degrees]S 32.865[degrees]E), -150 m, sand, dredged NMDP, RV Sardinops, st'n ZCC2, (S5096); off Rocktail Bay (27.1850[degrees]S 32.8483[degrees]E), -100 m, sand, dredged NMDP, RV Sardinops, st'n ZD4, (S5172); off Rocktail Bay (27.19[degrees]S 32.85[degrees]E), -100 m, sandstone rubble, dredged NMDP, RV Meiring Naude, st'n ZD1, (E1419); off Lala Neck (27.2267[degrees]S 32.8217[degrees]E), -75 m, coarse sand, sandstone, coral, dredged NMDP, RV Sardinops, st'n ZDD4, (S3771); off Gobey's Point (27.430[degrees]S 32.742[degrees]E), living, -55-100 m, sand, shell rubble, dredged NMDP, RV Meiring Naude, st'n ZG2, (E1440); Sodwana Bay (27.533[degrees]S 32.683[degrees]E), -100 m, dredged (A5803); off Jesser Point (27.553[degrees]S 32.713[degrees]E), living, -85 m, sponge, coral rubble, dredged NMDP, RV Meiring Naude, st'n ZH4, (E2936); NE of Gipsy Hill (27.753[degrees]S 32.663[degrees]E), -110 m, sponge, stones, dredged NMDP, RV Meiring Naude, st'n ZK4, (E7473); ditto (27.773[degrees]S 32.657[degrees]E), -6370 m, sand, sandstone, growths, dredged NMDP, RV Sardinops, st'n ZK25, (S3476); ditto (27.777[degrees]S 32.653[degrees]E), -74-87 m, sandstone rocks, slightly muddy sand, dredged NMDP, RV Sardinops, st'n ZK23, (E7462, S6370); Leadsman Shoal (27.800[degrees]S; 32.616[degrees]E), living, -100 m, dredged A.D. Connell, iv.1980 (B4059); off Gipsy Hill (27.8117[degrees]S 32.6570[degrees]E), -100-125 m, broken shell, dredged NMDP, RV Meiring Naude, st'n ZK9, (E3245); ditto (27.828[degrees]S 32.637[degrees]E), -47-50 m, fine sand, dredged NMDP, RV Meiring Naude, st'n ZK1, (S1136); NE of Leven Point (27.917[degrees]S 32.647[degrees]E), -250 m, coarse sand, dredged NMDP, RV Meiring Naude, st'n ZL5, (S1141); SE of Cape Vidal (28.320[degrees]S 32.607[degrees]E), -110 m, sponge rubble, dredged NMDP, RV Meiring Naude, ZM5, (E4907); off St Lucia Lighthouse (28.500[degrees]S 32.917[degrees]E), -100 m, mud & pebbles, dredged (A5716); Port Durnford-Richard's Bay (29.007[degrees]S 32.200[degrees]E), -152 m, mud and stones, dredged NMDP, RV Meiring Naude, st'n ZQ7, (V3922); off Glenton Reef (29.245[degrees]S 32.037[degrees]E), -200-210 m, sandy mud, dredged NMDP, RV Meiring Naude, st'n ZRR9, (S459); SE of Sheffield Beach (29.5050[degrees]S 31.7617[degrees]E), -100-105 m, glutinous grey mud, dredged NMDP, RV Meiring Naude, st'n XX139, (E5033); off Umlaas Canal (30.018[degrees]S 31.053[degrees]E), -150 m, coarse sand, numerous spatangoids, pebbles, dredged NMDP, RV Meiring Naude, st'n XX70, 9.viii.1985 (E7598); SE of Green Point (30.250[degrees]S 30.905[degrees]E), living, -100 m, fine sand & rubble, dredged NMDP, RV Meiring Naude, st'n XX92, 8.vii.1986 (D5989); off Umzinto (30.36[degrees]S 30.85[degrees]E), -84 m, dredged NMDP, RV Meiring Naude, st'n X2, 15.vii.1982 (D5426); off Park Rynie (30.375[degrees]S 30.855[degrees]E), -136 m, sponge rubble, dredged NMDP, RV Meiring Naude, 5.iii.1981 (B3889); ditto, living, -110 m, sponge rubble, dredged R. Kilburn, 2.iii.1981 (W7457); ditto, 100 m, sand and sponge rubble, dredged NMDP, RV Meiring Naude, 4.iii.1981 (B3719); off Port Shepstone (30.775[degrees]S 30.538[degrees]E), -70 m, eroded shell & rubble, dredged NMDP, RV Meiring Naude, 4.iii.1981 (B3655); off Margate (30.91[degrees]S 30.47[degrees]E), -100-110 m, sponge, dredged NMDP, RV Meiring Naude, st'n X6, 22,vii.1982 (B8786); off Trafalgar (31.020[degrees]S 30.382[degrees]E), -120 m, sand, sponge, dredged NMDP, RV Meiring Naude, st'n X4, 22.vii.1982 (B8866); off Mpahlana River (31.118[degrees]S 30.280[degrees]E), living, -100 m, sponge rubble, dredged NMDP, RV Meiring Naude, st'n XX3, 15. vi.1983 (C5248); between Mpahlana & Umyameni rivers (31.14[degrees]S 30.27[degrees]E), -100 m, sponge rubble, dredged NMDP, RV Meiring Naude, st'n XX8, (C5378). Eastern Cape: off Mtamvuna River (31.1483[degrees]S; 30.2617[degrees]E), -111 m, sponge, dredged NMDP, RV Meiring Naude, st'n A14, 18.viii.1981 (E278, E870); ditto (31.15[degrees]S 30.27[degrees]E), -137 m, rocks, sponge, dredged NMDP, RV Meiring Naude, st'n A16, 18.viii.1981 (E6994); off Kwanyana River (31.183[degrees]S 30.223[degrees]E), -100 m, sponge rubble, dredged NMDP, RV Meiring Naude, st'n XX10, (C5331,C5484); off Port Grosvenor (31.4083[degrees]S 29.9500[degrees]E), -80 m, lithothamnion sheets, dredged NMDP, RV Meiring Naude, st'n D20, 16.viii.1981 (E217, E6978); ditto (31.41[degrees]S 29.95[degrees]E), living, -80 m, worn coral nodules, dredged NMDP, RV Meiring Naude, st'n D17, 16. viii.1981 (E177, E346); ditto (31.4360[degrees]S 29.9516[degrees]E), -100-115 m, sand, some mud, solitary coral, shells, dredged NMDP, RV Meiring Naude, st'n D3, viii.1981 (C1333); off Mbotyi (31.550[degrees]S 29.863[degrees]E), -200 m, sandstone slabs, coarse sand, dredged NMDP, RV Meiring Naude, st'n F13, 4.vii.1986 (C9755); off N'tafufu River (31.578[degrees]S 29.662[degrees]E), -50 m, mud, sand, dredged NMDP, RV Meiring Naude, st'n H6, 14.viii.1981 (E241); off Mgazi River (31.738[degrees]S 29.537[degrees]E), -250 m, muddy sand, dredged NMDP, RV Meiring Naude, st'n J11, 4.vii.1985 (E7423); off Qora River (32.557[degrees]S 28.800[degrees]E), -100 m, coarse sand, some sponge rubble, dredged NMDP, RV Meiring Naude, st'n U6, (C5189); off Kei River (32.822[degrees]S 28.520[degrees]E), -138 m, coarse sand, dredged NMDP, RV Meiring Naude, st'n Z4, (C5116).

Distribution and habitat (Fig. 57): South-western Indian Ocean; from northern Madagascar to Mozambique, extending south to the Great Kei River, Eastern Cape, South Africa; -47-250 m, living specimens -55-110 m, on varied substrata, but usually associated with sponge and coral rubble.

Remarks: Distinctive amongst local species on account of its small size and turriculate profile. Specimens from Madagascar often rather broad and with a more coarsely stellate peripheral cord. Perrinia angulifera (A. Adams, 1853) has a similar elevated-trochiform spire, but is less pagodaform and much larger (length up to 16 mm). The most similar sympatric species is P. stellata (A. Adams, 1864), but that also generally attains a larger size (length up to 9.6 mm in south-eastern Africa) and has a much more strongly stellate periphery and stronger basal cords.

Schepman (1908) described a number of new species of this genus (placed as a subgenus of Calliostoma) from Indonesia, two of which, P. squamocarinata and P. nigromaculata, resemble the present taxon. P. squamocarinata (syntype examined, ZMAN 3.08.084, Fig. 71E, F) (additional figures provided by Poppe et al. (2006) and Poppe & Tagaro (2008)) differs in having stronger granules on the spiral cords above the peripheral keel on the last adult whorl, and the base retains an umbilicus (albeit small) and has five granular spiral cords (usually only four in P. konos) with strong axial pliculae in their intervals. P. nigromaculata (holotype examined, ZMAN 3.08.087, figured by Poppe et al. 2006) is less strongly keeled than P. konos and is larger (length ca 10 mm), has finer, more numerous spiral cords and a much more prominent tooth at the base of the columella (additional figures provided by Tsuchida & Ikebe (1990), Tsuchida & Kurozumi (1992), Poppe et al. (2006) and Poppe & Tagaro (2008)). Also from the Philippines, the recently described Perrinia cecileae Poppe, Tagaro & Dekker, 2006 has a similar elevated conical profile and is of a similar size, but it has more numerous spiral cords above the periphery, a weaker peripheral keel on the spire whorls, and much stronger columella teeth. Turcica (Perrinia) morrisoni Ladd, 1966 from the Miocene and Recent of the Marshall Islands is more depressed, has a more granular sculpture and fewer, stronger peripheral stellations.

Perrinia stellata (A. Adams, 1864) Figs 4O, 58-60

Turcica stellata: A. Adams 1864a: 508; Pilsbry 1890 in 1889-90: 418, pl. 67, fig. 77; Yen 1942: 177, pl. 11, fig. 11 (holotype); Mastaller 1979: 31; Smythe 1979: 64; idem 1882: 39; Glayzer et al. 1984: 318. Type loc.: China Seas (Cuming) [erroneous, here emended to the Gulf of Suez].

Tectaria armata: Issel 1869: 192, 289, pl. 2, fig. 7; Moazzo 1939: 183. Type loc.: Gulf of Suez, Red Sea.

Euchelus (Perrinia) stellata: MacAndrew 1870: 443.

Turcica (Perrinia) stellata: Melvill & Standen 1901: 351; Tomlin 1927: 298 [=Tectaria armata Issel, 1869]; Melvill 1928: 98; Lamy 1938: 82, fig. 9; Biggs 1973: 350; Kilburn 1977: 176.

Calliostoma stellatum: Hidalgo 1904-5: 255.

Tectarius armatus: Sturany 1903: 263, 278.

Turcica (Turcica) stellata: Bisacchi 1931: 182.

Perrinia stellata: Kendall & Skipwith 1969: 855; Bosch et al. 1995: 33, fig. 28; Hoenselaar & Dekker 1998: 199; Rusmore-Villaume 2008: 20; Zuschin et al. 2009: 99, pl. 9, figs 5-7; Bandel 2010: 470, fig. 15d-f.


Shell: Elevated-trochiform to turriculate (L/D=1.10-1.65); apical angle 42-63[degrees]; teleoconch of up to 8 whorls; whorls flat-sided and periphery angular, marked by strong, stellate, keel-like spiral cord, below which is a distinct supra-sutural sulcus. First teleoconch whorl rounded and sculptured only with close-set, curved, axial pliculae ([+ or -] 35); weak spiral cords develop toward end of second whorl; supra-sutural cord rapidly strengthening during third and subsequent whorls, and developing characteristic squamose, stellate projections; projections becoming fewer and larger with growth ([+ or -] 15 on last adult whorl); uppermost cord also strengthens, but to a lesser extent and becomes coronated by apically orientated, scale-like granules; these 1.5-2 times as numerous as peripheral projections; interval between peripheral cord/keel and abapical suture progressively deeper and more channelled with growth; development of remaining spirals between adapical suture and periphery variable, sometimes distinct, sometimes obsolete; axial sculpture becoming less obvious with growth; apical whorls (not first) somewhat cancellate, becoming more foveolate with elongate D-shaped pits on later whorls; pliculae in supra-sutural sulcus usually stronger and less close-set than those above peripheral keel. Base with 4 welldefined spiral cords and a fifth adjacent to columella; outermost cord strongest and frequently set with scale-like granules, in large specimens the other basal cords may also be somewhat granular; umbilicus closed. Peristome oblique; aperture D-shaped; columella more or less straight, a single weak tooth or rounded bulge present at its base in mature specimens; outer lip strongly notched at ends of peripheral and basal cords, particularly in subadults; interior of outer lip weakly thickened with in-running ridges at maturity, but most local material somewhat subadult in this respect; interior of aperture nacreous, angled beneath peripheral cord.

Microsculpture (Fig. 59B, C): Initial whorls somewhat worn in all available material, but evidently lacking vermiform spiral microsculpture; later whorls with distinct scratchlike microsculpture.

Protoconch (Fig. 59A, C): Apex missing or heavily encrusted in most specimens; protoconch remaining only in some juveniles and rather worn even in these; translucent white; diameter ca 260 um; moderately exsert; terminal lip roundly angled between mid-whorl and apical suture; superficial sculpture eroded in all available material.

Colour: Shell milky-white, fresh specimens with a cream to dirty buff intritacalx deposit, often with broad, dark greyish axial bands, particularly in juveniles. Many specimens with heavy, whitish or pinkish coralline encrustation.

Dimensions: Largest specimen (holotype of Tectaria armata), length 15.5 mm. Southeast African specimens develop mature apertural features at a smaller size than those from the north-western Indian Ocean and Red Sea, and never attain such a large size (largest southern African specimen, length 9.6 mm).

Operculum (Fig. 4O): Initially tightly multispiral, but whorls broadening with growth and becoming more openly multispiral, although somewhat less so than in P. angulifera and P. konos.

Radula: Unknown.

External anatomy: Like that of P. angulifera; only small specimens available, but no clear differences are apparent.

Type material: Holotype of Turcica stellata A. Adams, 1864 (Fig. 58A), in NHMUK (1968214). Holotype of Tectaria armata Issel, 1869 (Fig. 58F), in MCSNG.

Regional material examined (all NMSA unless indicated otherwise): KENYA: Shimoni (4.6482[degrees]S 39.3814[degrees]E), dredge 1+2, J.D. Taylor (NHMUK); ditto, dredge 6, J.D. Taylor (NHMUK); ditto (4.6536[degrees]S 39.3799[degrees]E), dredged in channel between mainland and Wasini Is., <12 fath. [-22 m], coral sand, J.D. Taylor (NHMUK); Kichangani, Shimoni, from broken fish traps, J.D. Taylor (NHMUK). TANZANIA: Entrance to Dar-es-Salaam harbour, -7.3 m, W. Rudman (AMS). MADAGASCAR: Banc du Leven, NW of Madagascar (12.533[degrees]S 47.667[degrees]E), -35-150 m, Benthedi Exped'n, st'n 5, dredged, 1977 (MNHN). MOZAMBIQUE: Mozambique Is. (15.03[degrees]S 40.73[degrees]E), R. Kilburn, ix.1974 (G7930); Bazaruto Archipelago, 0.5 miles west of Santa Carolina Is. (21.619[degrees]S 35.332[degrees]E), 3 fath. [-5.5 m], sand and shell debris, dredged, P. & E. Roscoe, 8.i.1976 (G7096); off Ponta da Barra lighthouse, 'Office Reef' (23.7869[degrees]S 35.5410[degrees]E), -24 m, dived, xi.2003 (D. Slater coll'n); off Lacerda Lighthouse (25.56167[degrees]S 32.84472[degrees]E), -52-55 m, dredged, vi.2010 (J. Rosado coll'n); NE of Maputo, -65 m, dredged J. Rosado, ii.2010 (D. Slater coll'n); off Ponta Techobanine (26.68132[degrees]S 32.95093[degrees]E), -60 m, dredged J. Rosado, xii.2005 (D. Slater coll'n). SOUTH AFRICA: KwaZulu-Natal: off Kosi Bay (26.905[degrees]S 32.925[degrees]E), living, -45 m, sand, stones and large algae, dredged NMDP, RV Sardinops, st'n ZA49, (S8937); Kosi Bay, main reef, 1-4 km south of estuary mouth (26.9210[degrees]S 32.8861[degrees]E), -18 m, underwater pump, D. Herbert & K. Bloem, 6.v.1990 (S2260); ditto, living, -20-22 m, underwater pump, D. Herbert & K. Bloem, 5.v.1990 (S1985); SE of Kosi River Mouth (26.9217[degrees]S 32.9183[degrees]E), living, -50 m, algae, shells, dredged NMDP, RV Meiring Naude, st'n ZA20, (E1364); off Boteler Point (27.0083[degrees]S 32.9117[degrees]E), -50 m, dead coral rubble and lithothamnion, dredged NMDP, RV Meiring Naude, st'n ZB7, (D7437); off Hully Point (27.3367[degrees]S 32.7700[degrees]E), -30-40 m, fine muddy sand, dredged NMDP, RV Meiring Naude, st'n ZF7, (E2904); off Sodwana Bay (27.553[degrees]S 32.686[degrees]E), -50 m, shell and coral sand with foraminiferans, dredged CSIR Water Research (A5849); ditto, -46 m, sediment at base of drop-off in canyon, dredged UND Marine Geoscience Unit, 7.xi.1992 (V2629); NE of Liefeldt's Rocks (27.7167[degrees]S 32.6650[degrees]E), -50 m, lithothamnion, medium sand, dead coral rubble, dredged NMDP, RV Meiring Naude, st'n ZJ6, (E4264); SE of Mission Rocks (28.2917[degrees]S 32.5433[degrees]E), -50 m, old coral rubble, lithothamnion, dredged NMDP, RV Meiring Naude, st'n ZN1, (E7350); Aliwal Shoal, off Scottburgh (30.2833[degrees]S 30.8333[degrees]E), ca -14 m, underwater pump, D. Herbert, (S8689); Lander's Reef, off Park Rynie (30.3333[degrees]S 30.8166[degrees]E), -34 m, sand, D. Herbert, (S6010).

Other material examined: PERSIAN GULF: Khor-al-Bazm, Abu Dhabi, H.E.J. Biggs (NMSA G6968 and NHMUK); Dabai [Dubai] R. Winckworth (NHMUK). GULF OF OMAN: Jask, Iran, R. Winckworth (NHMUK). PAKISTAN: Mekran Coast, V.W. MacAndrew (NHMUK). RED SEA: Suez, J.J. Walker and R. MacAndrew (NHMUK); Suez Canal, living, Cambridge University Exped'n (NHMUK 1928.3.30.236); Great Bitter Lake, Suez, living, dredged at -9 m, muddy sediment with coarse sand covered thickly by Sargassum, C. Beets (H. Dekker coll'n, large specimens); Nuweiba, Sinai, beach, J. Wise (AMS); Eilat, north beach, H. Kurutz & J. Wise (AMS); Gulf of Aqaba, Dahab, south side of Ras el Kura, sand beach, coral reef, reef flat (H. Dekker coll'n); Hurghada, Egypt, R. Kilburn (NMSA L376); 16 km S of Hurghada Egypt, sandy beach stones, dead coral, coral reef (H. Dekker coll'n). GULF OF ADEN: Aden, H. Burnup coll'n (NMSA G4296).

Distribution and habitat (Fig. 60): Persian Gulf, Gulf of Oman, Arabian Sea, Red Sea, Suez Canal and East Africa south to the KwaZulu-Natal south coast (Scottburgh area); living material collected from -18-50 m in South Africa, but probably also occurring in shallower water in truly tropical areas; available data on habitat preferences is inconclusive. A record from Zambales in the Philippines (Hidalgo 1904-05) requires confirmation.

Remarks: A characteristic species though somewhat variable in size and shell proportions. The only local species with which it might be confused is P. konos, but that species is smaller, never has such strongly developed peripheral projections, has weaker basal cords and more widely spaced axial pliculae on the apical whorls. Turcica (Perrinia) morrisoni Ladd, 1966 from the Marshall Islands, is smaller, less elevated and has much more obvious dentition inside the outer lip.

Tomlin (1927) followed by Lamy (1938) synonymised Tectaria armata Issel, 1869 from the Red Sea, with the present species. The figure of the holotype here provided (Fig. 58F) indicates that this was fully justified. Tomlin (1927) also questioned the validity of the original Cumingian locality data, China Seas. This is a notoriously vague locality which should be rejected in view of the known inaccuracy of the provenance attached to much Cuming material. With the exception of one unconfirmed record from the Philippines (Hidalgo 1904-05), the species has been recorded subsequently only from the western Indian Ocean. I here emend the type locality to be the Gulf of Suez.

Genus Pholidotrope gen. n.

Etymology: From Greek pholidotos (scaly) and trope (a turn); in reference to the scalelike sculpture on the spiral cords. Gender feminine.

Type species: Pholidotrope gloriosa sp. n.

Diagnosis: Shell small, profile conical with narrowly indented suture; suture level with subperipheral cord; sculpture coarsely cancellate with scale-like projections on spiral cords; columella with a single basal tooth; interior of outer lip thickened and set with ridge-like denticles, that nearest columella largest and separated from basal columella tooth by a U-shaped notch; denticles do not extend into aperture as in-running ridges; umbilical and parietal region covered by glossy inductural callus shield; aperture obliquely tangential to base of last adult whorl; outer lip with low subterminal external varix; protoconch exsert.

Remarks: Pholidotrope resembles Clypeostoma in possessing a well developed basal callus shield, but differs in being smaller, having a more conical profile, coarser sculpture, only one tooth on the columella and a subterminal external varix behind the outer lip. It also resembles Mirachelus Woodring, 1928 from the western Atlantic and eastern Pacific, but species of that genus lack an expanded basal callus and a subterminal external labral varix. Perrinia likewise lacks both these features, and in addition its aperture is spirally corded within and the dentition at the base of the columella is weaker.

Pholidotrope gloriosa sp. n. Figs 61, 62

Etymology: From Latin gloriosa (famous, glorious); in reference to Iles Glorieuses, near which the type material was collected.


Shell: Trochiform, small, spire conical, base somewhat flattened (L/D=1.18); teleoconch of 5.5 whorls; initial whorls rounded, becoming more flat-sided with growth; suture narrowly indented; sculpture of spiral cords and lamellate axial pliculae; first whorl sculptured only by relatively widely spaced axial pliculae (13), second whorl with 16 axial pliculae and 2 developing spiral cords; third whorl with 17 axial pliculae and a third spiral cord developing at shoulder; fourth and fifth whorls with 3 strong spiral cords and 18 and 19 axial pliculae respectively; cords and pliculae subequal in strength, their intersections produced into scale-like nodules, those on shoulder cord somewhat more rounded; suture level with subperipheral cord; cord intervals wider than cords themselves, that between peripheral and subperipheral cord deep. Base with 4 spiral cords, progressively weaker toward centre; umbilicus narrowly patent in juvenile, but umbilical and parietal regions covered by a thin, smooth inductural callus spreading from aperture at maturity. Peristome markedly oblique, more or less in one tangential plane; aperture D-shaped; columella more or less straight, but with a distinct, roundly trigonal tooth near its base; outer lip internally thickened, its edge flaring outward; inner thickening with 10 ridge-like denticles, that nearest columella largest, creating a U-shaped notch between it and columella tooth; interior of aperture nacreous, labral denticles not extending internally as in-running spiral ridges; exterior of outer lip subterminally thickened by a low collabral varix.

Microsculpture (Fig. 62A, B): Early whorls with close-set, microscopic granules, vermiform spiral threads not evident; fine, prosocline, scratch-like marks on later whorls.

Protoconch (Fig. 62C): White, globose and distinctly exsert, diameter ca 280 um; surface worn, but with evidence of a fine, flocculent microsculpture; terminal lip convex.

Colour: White.

Dimensions: Holotype, length 4.7 mm, diameter 3.9 mm.

Operculum, radula and external anatomy: Unknown.

Holotype: MADAGASCAR: NW of Madagascar, west of Banc du Leven (12.53[degrees]S 47.67[degrees]E), -35-150 m, Benthedi Exped'n, st'n 5, dredged, 1977 (MNHN 24656).

Paratype: MADAGASCAR: NW of Madagascar, west of Iles Glorieuses (11.487[degrees]S 47.303[degrees]E), -250 m, rock dredge, Benthedi Exped'n, st'n 8, dredged, 19.iii.1977 (MNHN 24657).

Distribution and habitat: Known only from off north-western Madagascar, -35-250 m, presumably on rocky substrata.

Remarks: The small size and conical profile of Pholidotrope gloriosa renders it distinctive amongst south-western Indian Ocean chilodontids. The only similar species, Perrinia konos, is more elevated, less coarsely sculptured and has less well-developed dentition at the base of the columella.

Genus Granata Cotton, 1957

Granata: Cotton 1957: 126. Type species: Stomatella imbricata Lamarck, 1816, by original designation.

Nomenclatural remarks: Cotton (1957), believing Stomatella auricula Lamarck, 1816 to be the type species of Stomatella Lamarck, 1816 (3), proposed a new generic taxon, Granata, for Stomatella imbricata Lamarck, 1816. However, there have been differing opinions concerning the type species designation for Stomatella. Early workers such as Thiele (1924, 1929) and Wenz (1938), following Gray (1847) and Pilsbry (1890 in 1890-91), believed it to be S. imbricata and, recognising the true affinities of that species, placed Stomatella near Euchelus and Danilia. In contrast, most recent authors (e.g. Keen 1960; Hickman & McLean 1990) have followed Cotton (1957) who observed that Anton had designated Stomatella auricula as the type species of Stomatella, in his Verzeichniss der Conchylien (Anton 1838, but cited as 1839), and that this pre-dated Gray's (1847) more widely known designation of S. imbricata. Macpherson and Gabriel (1962) claimed that no such designation existed in the Verzeichniss der Conchylien and believed Granata to be an 'absolute [objective] synonym' of Stomatella. Iredale and McMichael (1962) listed S. imbricata as the 'logotype' (type species by subsequent designation) of Stomatella, citing Dujardin in Dictionnaire Universel d' Histoire Naturelle (d'Orbigny 1839-1849) and giving a date of 'ante 1845'. The most recent analysis of the dates of publication for the various volumes and livraisons of d'Orbigny's Dictionnaire (Evenhuis 1990), however, gives the date for volume 12, livraison 133, pages 1-64 (entries Stellion to Strombides in the dictionary, Stomatella on p. 47) as 9 September 1848.

Examination of Anton's Verzeichniss shows that a type designation is in fact present, though somewhat concealed. In the introductory pages to this work, Anton stated "... den Gattungen (deren typusart mit Versalbuchstaben gedruckt ist)" [".. .the genera (whose type species are printed in capital letters)"] (Anton 1838: vi) and entry 1222 under Stomatella is printed "LUTEA = St. auricula Lam. = Patella lutea L.". However, since Stomatella lutea was not one of the nominal species originally included by Lamarck (1916) in Stomatella, it is not eligible for subsequent designation as the type species (ICZN 1999: Art. 69.1). This not withstanding, since Anton clearly indicated that he considered Patella lutea to be a synonym of Stomatella auricula, which was one of the species originally included in Stomatella, this is to be considered a valid fixation of the latter species as the type species of the genus (ICZN 1999: Art. 69.2.2). Dated 1838 (Cernohorsky 1978a), it is evidently the earliest designation available. Like Anton, Lamarck himself considered Patella lutea and Stomatella auricula to be synonymous (Lamarck 1822: 210), though Pilsbry (1890 in 1890-91) believed P. lutea to be an unidentifiable entity.

Remarks: Hickman (1998) believed Granata to be a monotypic genus endemic to the southern half of Australia. However, I follow other recent authors (e.g. Sasaki 2000; Poppe et al. 2006) in referring additional auriform chilodontids to this genus. While these may not be as depressed and haliotiform as the type species, they almost certainly belong to the same lineage. With these taxa included in Granata, it is evident that the genus is widely distributed in the Indo-West Pacific. In addition to being conchologically similar, G. sulcifera, like G. imbricata, has an enlarged right hypobranchial gland and secretes a noxious white mucus when irritated, and has a similar radula morphology.

The relationship of Granata and Stomatolina Iredale, 1937 (type species Stomatella rufescens Gray, 1847, by original designation) needs to be further investigated. Both stomatelline and chilodontid taxa have been referred to Stomatolina. Although the matter could be easily resolved by examination of the radula and external anatomy, the difficulty lies in obtaining reliably identified specimens of S. rufescens. On the evidence available (Iredale 1937), it seems more probable that Stomatolina is stomatelline, perhaps close to Pseudostomatella Thiele, 1924.

Though perhaps related to Granata, species of Hybochelus Pilsbry, 1890 (type species Stomatella cancellata Krauss, 1848, by original designation) differ in having a less expanded last adult whorl, an open umbilicus (typically) which is bordered by somewhat stronger cords, and an operculum which almost completely closes the aperture (pers. observ. Hybochelus mysticus (Pilsbry, 1890)). The genus occupies conchological morphospace somewhat intermediate between that of typical Euchelus and Granata. As indicated earlier (see Excluded taxa, p. 398), Krauss's Stomatella cancellata is an extralimital species and is not included in this revision.

Key to species of Granata in the south-western Indian Ocean

1 Aperture very broad, ratio of maximum:minimum aperture diameter 1.25-1.40; cream patterned with reddish to reddish brown spots that remain distinct on last adult whorl; axial sculpture of rather coarse, regular, crispate pliculae cumingii

--Aperture not as broad, ratio of maximum:minimum aperture diameter 1.04-1.16; initially with purplish to greyish brown spots, becoming more densely pigmented on last adult whorl; axial sculpture of rather uneven growth-lines sulcifera

Granata cumingii (A. Adams, 1854) comb. n. Figs 63-65

Stomatella cumingii: A. Adams 1854a: 834, pl. clxxv, fig. 38; idem 1854b: 74; Sowerby 1874: pl. v, fig. 32; Pilsbry 1890 in 1890-91: 13, pl. 52, fig. 67. Type loc.: none originally given; here designated to be tropical East Africa.

Stomatia cumingii: Chenu 1959 in 1959-62: 364, fig. 2709.

This species is similar to Granata sulcifera (below) and does not warrant a full redescription. It differs most obviously in shape and colour, having a distinctly more elongate aperture (ratio of maximum:minimum aperture diameter 1.25-1.40 in G. cumingii compared with 1.04-1.16 in G. sulcifera) and a more consistent colour pattern of reddish or reddish brown spots on a cream ground that remain distinct even on the last adult whorl. In addition, the sculpture of G. cumingii is somewhat coarser, the axial pliculae more crispate and regular, and the interstices more obviously iridescent. Although smaller, the three specimens listed below, are almost identical with the holotype and are undoubtedly conspecific therewith.

Besides mention in late nineteenth century iconographies (Sowerby 1874; Pilsbry 1890 in 1890-91), this species seems not to have been discussed in the literature subsequent to its original description. No locality data accompanied the holotype and, until now, the provenance of the species has remained unknown. Granata lyrata (Pilsbry, 1890), from Japan, which is also reportedly more strongly sculptured than G. sulcifera (Sasaki 2000), as a less elongate aperture and has greyish rather than reddish maculations.

Microsculpture: Similar to that of G. sulcifera.

Protoconch: Missing in all available specimens.

Operculum: Oligospiral; like that of G. sulcifera.

Radula: Unknown.

External anatomy: Only one, badly contracted specimen available, but evidently similar to that of G. sulcifera.

Type material: Holotype of Stomatella cumingii A. Adams, 1854, in NHMUK (1968201), maximum diameter 31.1 mm, height 19.8 mm (Fig. 63); provenance unknown.

Other material examined: KENYA: Kikambala (3.828[degrees]S 39.829[degrees]E), B. Hooper, don. A. Jenner, iii.1972 (NMSA F8740). MOZAMBIQUE: Conducia Bay (14.9128[degrees]S 40.7178[degrees]E), living, on rock on muddy bottom above LST, K. Grosch, ix.1976 (NMSA L1440). MADAGASCAR: Secteur de Lavanono (25.48333[degrees]S 44.93167[degrees]E), -14-18 m, tombant calcaire avec surplombs, Exped'n Atimo Vatae, st'n BS04, 30.v.2010 (MNHN).

Distribution and habitat (Fig. 65): Known only from the tropical western Indian Ocean, from southern Kenya and northern Mozambique to southern Madagascar; has been found living intertidally, on a rock on a muddy substratum, near low water.

Granata sulcifera (Lamarck, 1822) Figs 4C, 6D, 65-67

Stomatella sulcifera: Lamarck 1822: 210, No 3; Delessert 1841: pl. 33, fig. 3a, b; Krauss 1848: 93; A. Adams 1850: 30; 1854a: 833, pl. clxxiv, fig. 3; H. Adams & A. Adams 1854 in 1853-54: 435, 1858: pl. 49, fig. 8b, c (operculum); Brazier 1877: 46; Pilsbry 1890 in 1890-91: 11, pl. 52, fig. 59; Sowerby 1892: 46; Melvill & Standen 1895: 126; 1899: 178; 1901: 345; Hidalgo 1904-5: 259; Couturier 1907: 171; Hedley 1907: 478; 1909: 353; Schwartz 1910: 115; Dautzenberg 1929: 337 [543]; 1932: 79; Dautzenberg & Bouge 1933: 410; Mermod & Binder 1963: 136, fig. 206; Davies 1972: 253; Smythe 1982: 40, pl. 1e; Glayzer et al. 1984: 318; Kilburn & Rippey 1982: 42, pl. 9, fig. 5. Type loc.: 'les mers de la Nouvelle-Hollande' (Australia).

Stomatella articulata: A. Adams 1850: 30; 1854a: 834, pl. clxxiv, fig. 2; Pilsbry 1990 in 1890-91: 13, pl. 52, fig. 43; Sowerby 1874: pl. iv, fig. 22; Sowerby 1892: 46; Schwartz 1910: 115; Dautzenberg 1932: 79; Barnard 1951: 117, pl. xiv, fig. 17; 1963: 244, fig. 12b (radula); Paes da Franca 1960: 55, pl. 1, fig. 7; Macnae & Kalk 1969: 37, 118, 127; Kensley 1973: 44, fig. 108. Type loc.: 'In insulis Pacificis', but also given as 'Australia; Lord Hood's Is. [Tuamotu], South Seas, on the pearl oyster' [Cuming].

Stomatella elegans [non Gray, 1847]: Biggs & Grantier 1960: 387; Bosch & Bosch 1982: 37.

Stomatella (Stomatella) sulcifera: Kilburn 1972: 394 [= S. articulata].

?Hybochelus (Granata) sulcifera: Habe 1964: 11, pl. 4, fig. 16.

Granata sulcifera: Bosch et al. 1995: 32, fig. 27; Jansen 1996: 8, No 25; Sasaki 2000: 55, No 9; Poppe et al. 2006: 33, pl. 12, fig. 4; Poppe & Tagaro 2008: 172, pl. 31, fig. 54, 6.

not Stomatella sulcifera: A. Adams 1854a: 833, pl. clxxiv, fig. 3; Chenu 1859 in 1859-62: 363, fig. 2703; Sowerby 1874: pl. ii, fig. 11. The figures in these works depict a shell with bold axial bands (cf. Kilburn 1972).


Shell: Depressed turbiniform to auriform (L/D=0.74-0.90), last adult whorl expanding rapidly; teleoconch of up to 4 whorls; suture indented but not channelled, level with peripheral cord on spire whorls but descending below this near aperture; first teleoconch whorl initially more or less smooth, 3-4 spiral cords develop toward end of whorl; second whorl with 4-5 cords and further cords arising by intercalation with growth on subsequent whorls; end of penultimate whorl with 4-6 first-order cords; intervals between cords wider than cords themselves and usually with a weaker intermediary lira (frequently more than one near end of last adult whorl); axial pliculae develop during second and third whorls; pliculae initially rather regular and producing somewhat cancellate sculpture (second whorl), becoming finer, more close-set and irregular with growth; last adult whorl also with strong, irregular growth-lines; upper cords granular, those nearer periphery smoother. Base similarly sculptured, but cords lower; umbilicus lacking; aperture large, ovate; maximum:minimum aperture diameter 1.04-1.16; columella nacreous and lacking denticles, strongly concave, its junction with basal lip scarcely delimited; interior of outer lip not obviously thickened and lacking denticles or ridges; interior of aperture highly nacreous when fresh, with weak anglulations underlying external cords.

Microsculpture: Vermiform spiral threads not evident on juvenile shell, but surface of early whorls generally worn; later whorls with fine scratch-like axial microsculpture (Fig. 67A).

Protoconch (Fig. 67B): White, diameter 250-270 um; usually missing, damaged or badly eroded; protrudes slightly above first teleoconch whorl; sculptured with a coarsely flocculent sculpture with some traces of spiral threads (perhaps resembling that of Clypeostoma salpinx when fresh); apex weakly beaked; terminal lip with a welldeveloped projection just above mid-whorl, angular in some specimens rounded in others.

Colour: Initially whitish, with dark purplish or greyish brown spots appearing on cords during third whorl; subsequent whorls spotted, blotched or washed with similar shades, last whorl sometimes heavily so; in living specimens coloration frequently obscured by a dirty brownish periostracal layer.

Dimensions: Largest specimen, maximum diameter 21.6 mm, height 19.2 mm.

Operculum (Fig. 4C): Oligospiral; somewhat thicker than in other chilodontid genera; maximum diameter approx. half maximum diameter of aperture; frequently damaged. Radula (Fig. 67C, D): Formula [infinity]+(3-4)+1+(3-4)+[infinity], with 90-100 transverse rows of teeth; rachidian relatively weakly hooded, cusp acutely trigonal with margins serrated by lateral denticles. Lateral teeth overlapping extensively, their cusps similar to that of rachidian, but slightly asymmetrical and a little larger; whether the fourth tooth should be considered a lateral or a marginal is debateable (Barnard (1963) likewise observed a gradual transition from laterals to inner marginals). Remaining marginals numerous, longer and more slender, with recurved, pectinate cusps. In terms of its general form this radula is similar to that of the type species, G. imbricata, but in that species the tooth cusps are finer and more elongate (Hickman & McLean 1990; Hickman 1998).

External anatomy (Fig. 6D): General body colour yellowish white, epipodium paler with a few scattered black blotches, also on sides of the foot; cephalic tentacles, forehead and snout usually with grey-brown to black pigmentation. Cephalic lappets, neck lobes and epipodial fold well developed, forming an almost continuous sensory skirt around aperture margin, as in other auriform vetigastropods (e.g. Haliotis spp.). Cephalic lappets moderately broad with close-set, stubby processes on free margin; snout laterally expanded; post-ocular peduncle present on right (in both sexes), arising from posterior base of right eyestalk; peduncle with a longitudinal dorsal groove evident in some specimens; a smaller subocular tentacle emerging from ventral base of right eyestalk (illustrated also in G. lyrata by Kano 2008: fig. 4). Neck lobes originate beneath eyestalks, overlapping snout flanges, and extend posteriorly for approximately half length of animal; free margin of both lobes microscopically fimbriate with micropapillate tentacles of 2 or 3 sizes emerging from beneath margin; more numerous on left lobe than right; neck lobes narrowing posteriorly and merge seamlessly with epipodial fold. Edge of epipodial fold set almost throughout with epipodial tentacles of varying size, their number depending upon animal size (approx. 10 major epipodial tentacles on each side in large specimens, with numerous smaller intermediaries). An epipodial sense organ is present at base of most of the larger epipodial tentacles, but usually small and indistinct; none evident below neck lobes.

Type material: Holotype of Stomatella sulcifera Lamarck, 1822, in MHNG, "Nouv. Hollande", figured by Mermod & Binder (1963: fig. 206) and Poppe et al. (2006: fig. 26). Three syntypes of Stomatella articulata A. Adams, 1850, in NHMUK (1968113), one here illustrated and designated lectotype, maximum diameter 19.0 mm, height, 15.5 mm (Fig. 66F-H).

Regional material examined (selected, all NMSA unless indicated otherwise): TANZANIA: Dar-es-Salaam (6.7805[degrees]S 39.3104[degrees]E), I.F. Lambert (F8571). MOZAMBIQUE: Bazaruto Is., north reef (21.5197[degrees]S 35.4915[degrees]E), E. Roscoe, 1.xii.1974 (9647); Santa Carolina Is., Bay north of Battleship Rock (21.6187[degrees]S 35.3371[degrees]E), found under dead coral by itself, instead of in a colony, E. Roscoe (G2100); Santa Carolina Is., west bay (21.6187[degrees]S 35.3371[degrees]E), E. Roscoe, 1969-1973 (G99, J9645, J9646, J9648); ditto, living, R. Kilburn, 21.viii.1974 (K3084; L2695); Bazaruto Is., Ponta Gengareme (21.6638[degrees]S 35.4313[degrees]E), E. Roscoe, 1972-1974 (G2168, J9644); Inhaca Is., living, vii.1969 (L1437); ditto, bay west of lighthouse at Cabo Inhaca (25.9741[degrees]S 32.9778[degrees]E), living, under stones at low water, R. Kilburn & D. Herbert, x.1993 (L1442); ditto, sheltered west coast (26.0093[degrees]S 32.9075[degrees]E), living, under rock and dead coral blocks at low water, R. Kilburn & D. Herbert, x.1993 (L1446). MADAGSACAR: Pointe Barrow (25.20333[degrees]S 44.32167[degrees]E), -4 m, Exped'n Atimo Vatae, st'n TA31, 28.v.2010 (MNHN); secteur du Cap Malaimpioka (25.3650[degrees]S 44.8367[degrees]E), living, -10-17 m, sable et algues, Exped'n Atimo Vatae, st'n BP36, (MNHN); secteur de Lavanono (25.440[degrees]S 44.935[degrees]E), living, -14-18 m, tombant calcaire avec surplombs, Exped'n Atimo Vatae, st'n BS03, 29.v.2010 (MNHN); NW Rocher de l'Albatros (25.4700[degrees]S 44.9400[degrees]E), living, -12-14 m, fond rocheux et cailloux, Exped'n Atimo Vatae, st'n BB01, 25.v.2010 (MNHN); Ambatomainty (25.4383[degrees]S 44.9417[degrees]E), living, intertidal platier a galets basaltiques, Exped'n Atimo Vatae, st'n BM03, 25.v.2010 (MNHN); Ambatobe, pres Soamanitse (25.4567[degrees]S 44.9567[degrees]E), living, rochers et sable grossier, Exped'n Atimo Vatae, st'n BM02, v-vi.2010 (MNHN); Ambatobe, Bavarama (25.465[degrees]S 44.960[degrees]E), living, intertidal platier a galets basaltiques, Exped'n Atimo Vatae, st'n BM06, 28-29.v.2010 (MNHN); Andramara (25.48000[degrees]S 44.97167[degrees]E), living, intertidal roche basaltique, dalles sableuses, Exped'n Atimo Vatae, st'n BM10, (MNHN); Cap Sainte Marie (25.58167[degrees]S 45.12667[degrees]E), living, -15 m, dalles sableuses et blocs, Exped'n Atimo Vatae, st'n BV20, (MNHN); Cap Sainte Marie (25.6050[degrees]S 45.1617[degrees]E), intertidal platier calcaire, Exped'n Atimo Vatae, st'n BM16, (MNHN); Faux-Cap (25.5700[degrees]S 45.53167[degrees]E), -1-10 m, piscine de sable fin, protegee, Exped'n Atimo Vatae, st'n BV09, (MNHN); entree Est Baie des Galions (25.155[degrees]S 46.755[degrees]E), living, -10 m, brossage sur rares cailloux, Exped'n Atimo Vatae, st'n TB10, 11.v.2010 (MNHN); Cap Ranavalona (25.0717[degrees]S 46.9617[degrees]E), living, intertidal platier greseux et algues, Exped'n Atimo Vatae, st'n TM02, 27.iv.2010 (MNHN); Plage Libanona (25.0417[degrees]S 46.9950[degrees]E), -4-5 m, fond rocheux corallien, Exped'n Atimo Vatae, st'n TB07, 9.v.2010 (MNHN); Cap d'Antsirabe (25.0433[degrees]S 46.9967[degrees]E), living, intertidal platier rocheux, Exped'n Atimo Vatae, st'n TM 14, 6.v.2010 (MNHN); Plage Monseigneur (25.0350[degrees]S 46.9983[degrees]E), living, intertidal platier rocheux avec algues, Exped'n Atimo Vatae, st'n TM01, iv-v.2010 (MNHN); Pointe Flacourt (25.025[degrees]S 47.000[degrees]E), living, -2-4 m, rochers dans sable, Exped'n Atimo Vatae, st'n TB13, 15.v.2010 (MNHN); Port de Fort Dauphin (25.0267[degrees]S 47.0000[degrees]E), living, -2-4 m, cailloux sur sable, Exped'n Atimo Vatae, st'n TV21, 15.v.2010 (MNHN); Pointe Evatra, crique (24.9683[degrees]S 47.1017[degrees]E), living, -3-8 m, fond rocheux et gazon d'algues, Exped'n Atimo Vatae, st'n TR05, 30.iv.2010 (MNHN); sud de la Baie de Lokaro (24.9500[degrees]S 47.1067[degrees]E), -5-6 m, limon et sable sur roche, Exped'n Atimo Vatae, st'n TV11, 6.v.2010 (MNHN); Ilot de Lokaro (24.9417[degrees]S 47.1183[degrees]E), intertidal sable, mode battu, Exped'n Atimo Vatae, st'n TM05, 30.iv.2010 (MNHN); Sainte Luce, sud Ilot Souillac (24.76333[degrees]S 47.20667[degrees]E), living, -4-7 m, Exped'n Atimo Vatae, st'n TA37, (MNHN). REUNION: Not further localised (M. Jay coll'n, MNHN). SOUTH AFRICA: KwaZulu-Natal: Between Bhanga Neck and Kosi Bay, reef off marker 13 north (26.93[degrees]S 32.90[degrees]E), living, -9-14 m, D. Herbert & F. Wiercx et al. dived, 7&12.v.1990 (S1674); Leadsman Shoal, Raggie Reef, 1-2km North of Leven Point (27.90[degrees]S 32.62[degrees]E), -9-14 m, mixed algal and coral reef, D. Herbert & NPB, dived, 13.v.1998 (E2520); Umhlali (29.50[degrees]S 31.23[degrees]E), H. Burnup coll'n (8655); Umhlali, Thompson's Bay, Charles Pool (29.5229[degrees]S 31.2278[degrees]E), R. & E. Kilburn, J. Marais 1972-1979 (9332, 9517, B6720, S9043); Tongaat (29.583[degrees]S 31.133[degrees]E), H. Burnup coll'n (6369); Umdloti (29.6760[degrees]S 31.1158[degrees]E), W. Falcon coll'n (6366, 8656); Umdloti (29.6829[degrees]S 31.1127[degrees]E), rocky intertidal zone, low shore pools D. Herbert, L. Davis & T. Nangammbi, 27.ii.2005 (W2790); Durban (29.85[degrees]S 31.02[degrees]E), H. Burnup coll'n (6370); ditto, W. Falcon coll'n (A4558); ditto. S. Fenwick (A2009); ditto, R. Kilburn (5708, B5676); Durban Bay (29.8742[degrees]S 31.0559[degrees]E), dredgings B.J. Young, don. xi.1976-1979 (A5187, B2306, B2307); Durban, Vetch's pier (29.866975[degrees]S 31.052026[degrees]E), living, R. & E. Kilburn, 6.iv.1970 (V3925); Durban area, Reunion Rocks (29.9860[degrees]S 30.9648[degrees]E), living, rocky intertidal zone, D. Herbert, 19.iii.2003 (W546); Isipingo tidal pool and adjacent rocks (29.9996[degrees]S 30.9476[degrees]E), R. Kilburn, D. Herbert & R. Fregona 25.iii.1985 (9023, D609); Widenham, intertidal rocks (30.216829[degrees]S 30. 798644[degrees]E), low shore spring tide, living under large rocks with spaces below, together with arcid and carditid bivalves, leg. D. Herbert & L. Davis, 23.ix.2010 (W7463); Aliwal Shoal, off Scottburgh (30.2833[degrees]S 30.8333[degrees]E), -14-20 m, D. Herbert, dived, 25.x.1992 (S7972); ditto, -25-27 m, hand-dredged sand and reef debris, D. Herbert, 4.iv.1992 (S7160); ditto, -10-20 m, hand-dredged sand, D. Herbert, (S8016); Park Rynie, Rocky Bay (30.3364[degrees]S 30.7353[degrees]E), J. Marais (S9056); Mtwalume (30.4833[degrees]S 30.6333[degrees]E), living, intertidal rock pools, R. Kilburn & D. Herbert, 12.viii.1984 (B8666, V3927); Port Shepstone (30.75[degrees]S 30.45[degrees]E), H. Burnup coll'n (A4556, A4557); Port Shepstone area, Shelly Beach (30.817[degrees]S 31.658[degrees]E), W.G. Rump, ii.1930 (6367); Port Edward area, Leisure Bay (31.0214[degrees]S 30.2485[degrees]E), J.P. Marais (S9042). Eastern Cape: Pondoland Coast, A. Filmer, ex Transvaal Mus. 1978, H. Becker coll'n (B6845); Mzamba (31.08[degrees]S 30.20[degrees]E), beach drift, J. P. Marais, iv.1992 (S8342); ditto, R. Kilburn & D. Herbert, 12-30.v. 1986 (D2953); between Mzamba and Mtentu Rivers, don. J. Stannard, vii.1988 (E5989); Msikaba Is., north side (31.3248[degrees]S 29.9682[degrees]E), R. Kilburn et al,, viii.1983 (C5503); Port Grosvenor (31.38[degrees]S 29.90[degrees]E), R. Kilburn, 30.iv.1976 (A4852); Mbotyi (31.465[degrees]S 29.736[degrees]E), living, R. Kilburn & D. Herbert, v-vi.1985 (C8270); Mbotyi east (31.4588[degrees]S 29.7484[degrees]E), sheltered bay, loose rocks and crevices, fine silt, large pool behind reef, R. Kilburn & J. McKay, 26-27.iv.1976 (A5263); Lwandile/Mdumbi (31.883[degrees]S 29.266[degrees]E), R. Kilburn & R. Fregona, vii.1981 (C31); Coffee Bay (31.98[degrees]S 29.15[degrees]E), W. Tyson, ex Albany Mus. 1980 (B5565).

Other material examined: PERSIAN GULF: not further localised, Lebour (NMSA H6196). PAKISTAN: Karachi, R. Winckworth (NHMUK). INDIA: Mannar, R. Winckworth (NHMUK); Gulf of Mannar, Tuticorin, Koswari Is. (8.8704[degrees]N: 78.2255[degrees]E), beach-drift, R. Kilburn, 12.x.2000 (NMSA L5243). SEYCHELLES: Cerf Is., R.C. Wood (NMSA J7964). AUSTRALIA: Lizard Is., Queensland, J.D. Taylor (NHMUK).

Literature records: MADAGASCAR: Diego-Suarez [Antsiranana] (12.267[degrees]S 49.283[degrees]E), Decary (Dautzenberg 1932); fle Ste-Marie, entre l'lle aux Nattes et Ilampy (17.071[degrees]S 49.836[degrees]E) (Dautzenberg 1929); Lambetabe [Lambelabe] (24.783[degrees]S 43.933[degrees]E) (Dautzenberg 1929); Faux-Cap (25.567[degrees]S 45.517[degrees]E), Decary (Dautzenberg 1932); Cap Ste-Marie (25.599[degrees]S 45.137[degrees]E), Decary (Dautzenberg 1932, as Stomatella articulata).

Distribution and habitat (Fig. 65): Indo-West Pacific; from Japan (Sasaki 2000) and the Tuamotu Archipelago (Couturier 1907) in the east, to the eastern seaboard of Africa, extending south to the northern Eastern Cape (Coffee Bay, 32.00[degrees]S); common in the low intertidal and shallow subtidal down to -18 m (empty shells to -45 m), living specimens most often found under stones and dead coral blocks, sometimes in small groups, in both sheltered and somewhat exposed habitats; often where the rock rests on muddy sand and where conditions are somewhat anoxic (Kilburn 1972; Kilburn & Rippey 1982, and pers. observ.). Shells of living specimens frequently encrusted with tubes of spirorbid polychaetes, sometimes heavily so.

Remarks: Granata sulcifera is the only chilodontid species commonly found living intertidally in southern Africa and is easily recognised by its low spire and rapidly expanding last adult whorl. Vaceuchelus gemmula and V. natalensis may also be found intertidally in South Africa, but are less frequently encountered, and are probably often overlooked on account of their small size.

Granata elegans (Gray, 1847) from north-eastern Australia, G. lyrata (Pilsbry, 1890) (not Stomatia lirata A. Adams, 1850 - a species of Pseudostomatella) from Japan, and G. maculata (Quoy & Gaimard, 1834), described from Vanikoro Is., need to be examined for comparison as potential synonyms, but G. sulcifera predates all. Much museum material identified as G. elegans is in fact referable to G. sulcifera, and so probably are some literature references (e.g. Bosch & Bosch 1982); unfortunately the whereabouts of the type material of Gray's species is unknown. G. lyrata reportedly differs from G. sulcifera in having a less rapidly expanding last adult whorl and in being more strongly sculptured (Sasaki 2000).

This species has also been recorded from Pleistocene shorelines in the southern and eastern Cape, South Africa (Schwarz 1910; Barnard 1962; Davies 1972). Although such a range extension would have been quite possible during warmer interglacial periods, these records require confirmation since there may well have been confusion with the somewhat similar Pseudostomatella orbiculata (A. Adams, 1850), which is not uncommon in raised beach deposits in the Algoa Bay-Mossel Bay area (Kilburn & Tankard 1975). The same may also apply to Sowerby's (1892) record of Stomatella articulata from the Bairstow collection (i.e. Port Elizabeth).


List of ZooBank LSIDs for new genus- and species-group names.

Ascetostoma gen. n.:

Clypeostoma gen. n.:

Pholidotrope gen. n.:

Clypeostoma reticulatum sp. n.:

Danilia boucheti sp. n.:

Danilia textilis sp. n.:

Herpetopoma serratocinctum sp. n.:

Herpetopoma stictum sp. n.:

Pholidotrope gloriosa sp. n.:

Vaceuchelus cretaceus sp. n.:

Vaceuchelus jayorum sp. n.:


Material from South African waters was collected on board the RV Meiring Naude and RV Sardinops and was made possible through the allocation of ship's time by the CSIR and South African Division of Sea Fisheries respectively. The dredging work undertaken formed part of the Natal Museum Dredging Programme led by Dick Kilburn.

Deep-water material was collected off Mozambique on board RV Vizconde de Eza during the Mainbaza cruise, a joint effort of MNHN and Instituto Espanol de Oceanografia (IOE), and off Madagascar in 2009-2010 on board FV Miriky and FV Nosy Be 11 owned by the Societe des Pecheries de Nosy Be (Groupe Unima); further material was collected intertidally and by diving in south Madagascar during Expedition Atimo Vatae . These Mozambique-Madagascar expeditions (2009-2010) were funded by the Total Foundation, Prince Albert II of Monaco Foundation, and Stavros Niarchos Foundation, and were conducted by the MNHN and Pro-Natura International (PNI) as part of their "Our Planet Reviewed' programme (PI Philippe Bouchet). In addition, two earlier cruises yielded further chilodontid lots in the MNHN: Benthedi cruise, conducted in 1977 on board RV Noroit in the northern part of the Mozambique Channel (PI Bernard Thomassin); and MD32, conducted in 1982 on board RV Marion-Dufresne around Reunion. I am most grateful to Philippe Bouchet for allowing me to study this MNHN material, as well as that from Reunion in the collection of the late Maurice Jay (per Mireille Guillaume). Additional material from their private collections was loaned by Henk Dekker, Johan Marais, Jose Rosado and Dave Slater.

For their assistance in allowing me access to the collections under their care and for answering queries, I thank Ian Loch and Winston Ponder (AMS); Nathalie Memoire (MHNB); Virginie Heros and Philippe Maestrati (MNHN); Henry McGhie (MMUM); Tom Kemp (OXUM); Jon Ablett, Amelia MacLellan, Joan Pickering and Kathie Way (NHMUK); Abel Prieur (UCBL); Paul Greenhall (USNM); Shirley Slack-Smith and Fred Wells (WAM); Robert Moolenbeek (ZMAN). For photographs of type specimens I thank: Maria Tavano (MCSNG); Philippe Maestrati (MNHN); Bob Hershler and Yolanda Villacampa (USNM), and Dick Kilburn.

Philippe Bouchet and Gary Rosenberg provided valuable input regarding nomenclatural issues, and Anders Waren and Bruce Marshall provided much appreciated comments during the review process. Linda Davis catalogued type material and prepared line drawings from my anatomical sketches. SEM facilities were made available by the Centre for Electron Microscopy, University of KwaZulu-Natal.

This study forms part of a programme of research on southern African molluscs supported by a grant from the National Research Foundation (NRF) of South Africa (GUN 61261).


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(1) Chilodontinae Wenz, 1938 (Mollusca) is homonymous with Chilodontidae Macalister, 1876 (Ciliophora) and Chilodontinae Eigenmann, 1910 (Pisces). An application has been submitted to the International Commission on Zoological Nomenclature (Case 3555) to emend the spelling of Chilodontidae Wenz, 1938 to Chilodontaidae (Herbert & Bouchet 2011). While such a case is under consideration by the Commission, prevailing usage is to be retained (ICZN 1999: Art. 82.1).

(2) Beu and Climo (1974) cited the type species of Heliciella as H. costellata O.G. Costa, 1861, by monotypy. In fact, O.G. Costa (1861) described two species within his new genus Heliciella, H. costellata and H. mutabilis. The type species of Heliciella was not fixed until Dall (1927: 134) formally designated it to be H. costellata [although Monterosato (1884: 109) might also be deemed to have done the same]. A subsequent designation ofH. mutabilis as the type species of Heliciella by Bouchet and Waren (1988: 86) is invalid.

(3) Although Lamarck (1816) is traditionally cited as the author of Stomatella, the genus has recently been credited to Bowdich (1822) (Bouchet & Rocroi 2005). This stems from the fact that Lamarck's plate of Stomatia and Stomatella in his Tableau Encyclopedique (Lamarck 1816: pl. 450) has no associated legend and thus no species are listed under these names and it is not evident which figures he considered to belong to the respective genera. The legend is taken to have been eventually published in 1827 (Evenhuis 2003). Prior to this, however, Lamarck clarified his concept of these genera in his Histoire naturelle des animaux sans vertebres (Lamarck 1822; April), where he described a number of species, but this was predated by a publication in which Bowdich (1822; February) discussed Stomatella, citing and illustrating a single species, Stomatella imbricata, thus suggesting that Bowdich was the first to validly propose the genus. However, in reallity a legend was published in 1816 for the Stomatia and Stomatella plate in the Tableau Encyclopedique, in a 16-page "Liste des objets representes dans les planches de cette livraison" that is unfortunately lacking in many sets of the work (Evenhuis & Petit 2003; Petit 2011). The author of Stomatella is thus indeed Lamarck (1816). This superficially trivial nomenclatural fact is of considerable significance, for had Bowdich (1822) been the first to validly propose the Stomatella, then its type species would (by monotypy) be Stomatella imbricata and not Stomatella auricula. This in turn would mean that the trochid subfamily currently known as the Stomatellinae ought no longer be known by that name, and the Stomatellinae/idea would in fact be an earlier name for the Chilodontidae. Fortunately such is not the case, but I document the matter to provide clarification.

D. G. Herbert

KwaZulu-Natal Museum, P. Bag 9070, Pietermaritzburg, 3200 South Africa and School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, 3206 South Africa;

Chilodontidae of the south-western Indian Ocean:
geographical distribution, bathymetric range and
principle habitat preferences. Where possible,
bathymetric data are given only for live collected
samples. Distribution: IWP--Indo-West Pacific;
MI--Mascarene Islands; RSA--South Africa;
SWIO--south-western Indian Ocean; WIO--western
Indian Ocean.

     Species                   Distribution

1    Clypeostoma reticulatum   SWIO
2    Clypeostoma salpinx       SWIO
3    Clypeostoma meteorae      WIO
4    Clypeostoma nortoni       IWP
5    Danilia boucheti          SWIO
6    Danilia textilis          SWIO
7    Ascetostoma               SWIO
8    Euchelus asper            IWP
9    Euchelus atratus          IWP
10   Herpetopoma (s.s.)        IWP
11   Herpetopoma (s.s.)        WIO
12   Herpetopoma (s.s.)        WIO
13   Herpetopoma (s.s.)        MI
14   Herpetopoma (s.l.)        SWIO
15   Herpetopoma (s.l.)        IWP
16   Herpetopoma (s.l.)        IWP
17   Perrinia angulifera       IWP
18   Perrinia konos            SWIO
19   Perrinia stellata         WIO
20   Pholidotrope gloriosa     SWIO
21   Vaceuchelus cretaceus     SWIO
22   Vaceuchelus jayorum       SWIO
23   Vaceuchelus gemmula       RSA
24   Vaceuchelus natalensis    SWIO
25   Vaceuchelus               MI
26   Granata cumingii          WIO
27   Granata sulcifera         IWP

     Species                   Depth range in SWIO

1    Clypeostoma reticulatum   228-230 m, dead shells
2    Clypeostoma salpinx       60-277 m, living
3    Clypeostoma meteorae      80-257 m, dead shells
4    Clypeostoma nortoni       90-333 m, dead shells
5    Danilia boucheti          238-249 m, dead shells
6    Danilia textilis          150-250 m, living
7    Ascetostoma               50-85 m, living
8    Euchelus asper            no data
9    Euchelus atratus          no data
10   Herpetopoma (s.s.)        near shore, living
11   Herpetopoma (s.s.)        near shore, living
12   Herpetopoma (s.s.)        no data
13   Herpetopoma (s.s.)        near shore, living
14   Herpetopoma (s.l.)        70-180 m, living
15   Herpetopoma (s.l.)        280-375 m, dead
       ?naokoae                  shells
16   Herpetopoma (s.l.)        near shore to 247 m,
       xeniolum                dead shells
17   Perrinia angulifera       50-65 m, living
18   Perrinia konos            55-110 m, living
19   Perrinia stellata         18-50 m, living
20   Pholidotrope gloriosa     35-250 m, dead shells
21   Vaceuchelus cretaceus     near shore to 70 m,
22   Vaceuchelus jayorum       near shore, living
23   Vaceuchelus gemmula       low shore to 115 m,
24   Vaceuchelus natalensis    low shore to 84 m,
25   Vaceuchelus               near shore, living
26   Granata cumingii          low shore, living
27   Granata sulcifera         low shore to 18 m,

     Species                   Principle habitat in

1    Clypeostoma reticulatum   no data
2    Clypeostoma salpinx       sponge communities
3    Clypeostoma meteorae      no data
4    Clypeostoma nortoni       no data
5    Danilia boucheti          no data
6    Danilia textilis          sponge communities
7    Ascetostoma               sandstone and coral
       providentiae            rubble
8    Euchelus asper            no data
9    Euchelus atratus          no data
10   Herpetopoma (s.s.)        coral rubble
11   Herpetopoma (s.s.)        coral rubble
12   Herpetopoma (s.s.)        no data
13   Herpetopoma (s.s.)        fringing reef systems
14   Herpetopoma (s.l.)        varied hard substrata
15   Herpetopoma (s.l.)        no data
16   Herpetopoma (s.l.)        no data
17   Perrinia angulifera       coral rubble and
                               lithothamnion pebbles
18   Perrinia konos            sponge and coral
19   Perrinia stellata         inconclusive
20   Pholidotrope gloriosa     rocky substrata
21   Vaceuchelus cretaceus     coral rubble
22   Vaceuchelus jayorum       rocky substrata
23   Vaceuchelus gemmula       rocky substrata
24   Vaceuchelus natalensis    rocky substrata
25   Vaceuchelus               rocky substrata
26   Granata cumingii          under rocks
27   Granata sulcifera         under rocks and coral
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Article Details
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Title Annotation:p. 461-502
Author:Herbert, D.G.
Publication:African Invertebrates
Article Type:Report
Geographic Code:0INDI
Date:Dec 1, 2012
Previous Article:A revision of the chilodontidae (Gastropoda: Vetigastropoda: Seguenzioidea) of Southern Africa and the south-western Indian ocean.
Next Article:Revision of Durbadnus Pasteels, 1954, with notes on other Afrotropical Blennocampinae and Allantinae (Hymenoptera: Symphyta: Tenthredinidae).

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