Granata sulcifera (Lamarck, 1822)

Herbert, D. G., 2012, A revision of the Chilodontidae (Gastropoda: Vetigastropoda: Seguenzioidea) of southern Africa and the south-western Indian Ocean, African Invertebrates 53 (2), pp. 381-381 : 484-489

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; 1854 a: 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 ; 1854 a: 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 1854 a: 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).

Description:

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 µm; 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 well-developed 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 ∞+(3–4)+1+(3–4)+∞, 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. Halioti s 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°S 39.3104°E), I.F. Lambert (F8571). MOZAMBIQUE: Bazaruto Is., north reef (21.5197°S 35.4915°E), E. Roscoe, 1 xii.1974 (9647); Santa Carolina Is., Bay north of Battleship Rock (21.6187°S 35.3371°E), found under dead coral by itself, instead of in a colony, E. Roscoe (G2100); Santa Carolina Is., west bay (21.6187°S 35.3371°E), E. Roscoe, 1969–1973 (G99, J9645, J9646, J9648); ditto, living, R. Kilburn, 21.viii.1974 (K3084; L2695); Bazaruto Is., Ponta Gengareme (21.6638°S 35.4313°E), E. Roscoe, 1972–1974 (G2168, J9644); Inhaca Is., living, vii.1969 (L1437); ditto, bay west of lighthouse at Cabo Inhaca (25.9741°S 32.9778°E), living, under stones at low water, R. Kilburn & D. Herbert, x.1993 (L1442); ditto, sheltered west coast (26.0093°S 32.9075°E), living, under rock and dead coral blocks at low water, R. Kilburn & D. Herbert, x.1993 (L1446). MADAGSACAR: Pointe Barrow (25.20333°S 44.32167°E), - 4 m, Exped’n ATIMO VATAE, st’n TA31, 28.v.2010 ( MNHN) ; secteur du Cap Malaimpioka (25.3650°S 44.8367°E), living, - 10–17m, sable et algues, Exped’n ATIMO VATAE, st’n BP36, 8.vi.2010 ( MNHN) ;

secteur de Lavanono (25.440°S 44.935°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°S 44.9400°E), living, - 12–14 m, fond rocheux et cailloux, Exped’n ATIMO VATAE, st’n BB01, 25.v.2010 ( MNHN);Ambatomainty (25.4383°S 44.9417°E), living, intertidal platier a galets basaltiques, Exped’n ATIMO VATAE, st’n BM03, 25.v.2010 ( MNHN); Ambatobe, pres Soamanitse (25.4567°S 44.9567°E), living, rochers et sable grossier, Exped’n ATIMO VATAE, st’n BM02, v–vi.2010 ( MNHN); Ambatobe, Bavarama (25.465°S 44.960°E), living, intertidal platier a galets basaltiques, Exped’n ATIMO VATAE, st’n BM06, 28–29.v.2010 ( MNHN);Andramara (25.48000°S 44.97167°E), living, intertidal roche basaltique, dalles sableuses, Exped’n ATIMO VATAE, st’n BM10, 2.vi.2010 ( MNHN); Cap Sainte Marie (25.58167°S 45.12667°E), living, - 15 m, dalles sableuses et blocs, Exped’n ATIMO VATAE, st’n BV20, 10.vi.2010 ( MNHN); Cap Sainte Marie (25.6050°S 45.1617°E), intertidal platier calcaire, Exped’n ATIMO VATAE, st’n BM16, 10.vi.2010 ( MNHN); Faux-Cap (25.5700°S 45.53167°E), - 1–10 m, piscine de sable fin, protegée, Exped’n ATIMO VATAE, st’n BV09, 4.vi.2010 ( MNHN); entrée Est Baie des Galions (25.155°S 46.755°E), living, - 10 m, brossage sur rares cailloux, Exped’n ATIMO VATAE, st’n TB10, 11.v.2010 ( MNHN); Cap Ranavalona (25.0717°S 46.9617°E), living, intertidal platier gréseux et algues, Exped’n ATIMO VATAE, st’n TM02, 27.iv.2010 ( MNHN); Plage Libanona (25.0417°S 46.9950°E), - 4–5 m, fond rocheux corallien, Exped’n ATIMO VATAE, st’n TB07, 9.v.2010 ( MNHN); Cap d’Antsirabe (25.0433°S 46.9967°E), living, intertidal platier rocheux, Exped’n ATIMO VATAE, st’n TM 14, 6.v.2010 ( MNHN); Plage Monseigneur (25.0350°S 46.9983°E), living, intertidal platier rocheux avec algues, Exped’n ATIMO VATAE, st’n TM01, iv–v.2010 ( MNHN); Pointe Flacourt (25.025°S 47.000°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°S 47.0000°E), living, - 2–4 m, cailloux sur sable, Exped’n ATIMO VATAE, st’n TV21, 15.v.2010 ( MNHN); Pointe Evatra, crique (24.9683°S 47.1017°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°S 47.1067°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°S 47.1183°E), intertidal sable, mode battu, Exped’n ATIMO VATAE, st’n TM05, 30.iv.2010 ( MNHN); Sainte Luce, sud Ilot Souillac (24.76333°S 47.20667°E), living, - 4–7 m, Exped’n ATIMO VATAE, st’n TA37, 5–6.vi.2010 ( MNHN). RÉUNION: 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°S 32.90°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°S 32.62°E), - 9–14 m, mixed algal and coral reef, D. Herbert & NPB, dived, 13.v.1998 (E2520); Umhlali (29.50°S 31.23°E), H. Burnup coll’n (8655); Umhlali, Thompson’s Bay , Charles Pool (29.5229°S 31.2278°E), R. & E. Kilburn, J. Marais 1972–1979 (9332, 9517, B6720, S9043); Tongaat (29.583°S 31.133°E), H. Burnup coll’n (6369); Umdloti (29.6760°S 31.1158°E), W. Falcon coll’n (6366, 8656); Umdloti (29.6829°S 31.1127°E), rocky intertidal zone, low shore pools D. Herbert, L. Davis & T. Nangammbi, 27.ii.2005 (W2790); Durban (29.85°S 31.02°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°S 31.0559°E), dredgings B.J. Young, don. xi.1976 –1979 (A5187, B2306, B2307); Durban, Vetch’s pier (29.866975°S 31.052026°E), living, R. & E. Kilburn, 6.iv.1970 (V3925); Durban area , Reunion Rocks (29.9860°S 30.9648°E), living, rocky intertidal zone, D. Herbert, 19.iii.2003 (W546); Isipingo tidal pool and adjacent rocks (29.9996°S 30.9476°E), R. Kilburn, D. Herbert & R. Fregona, 25.iii.1985 (9023, D609); Widenham, intertidal rocks (30.216829°S 30. 798644°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°S 30.8333°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, 30.vi.1991 (S8016); Park Rynie, Rocky Bay (30.3364°S 30.7353°E), J. Marais (S9056); Mtwalume (30.4833°S 30.6333°E), living, intertidal rock pools, R. Kilburn & D. Herbert, 12.viii.1984 (B8666, V3927); Port Shepstone (30.75°S 30.45°E), H. Burnup coll’n (A4556, A4557); Port Shepstone area , Shelly Beach (30.817°S 31.658°E), W.G. Rump, ii.1930 (6367); Port Edward area , Leisure Bay (31.0214°S 30.2485°E), J.P. Marais (S9042). Eastern Cape: Pondoland Coast, A. Filmer, ex Transvaal Mus. 1978, H. Becker coll’n (B6845); Mzamba (31.08°S 30.20°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°S 29.9682°E), R. Kilburn et al., viii.1983 (C5503); Port Grosvenor (31.38°S 29.90°E), R. Kilburn, 30.iv.1976 (A4852); Mbotyi (31.465°S 29.736°E), living, R. Kilburn & D. Herbert, v–vi.1985 (C8270); Mbotyi east (31.4588°S 29.7484°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°S 29.266°E), R. Kilburn & R. Fregona, vii.1981 (C31); Coffee Bay (31.98°S 29.15°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°N: 78.2255°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°S 49.283°E), Decary (Dautzenberg 1932); Île Ste-Marie, entre l’Île aux Nattes et Ilampy (17.071°S 49.836°E) ( Dautzenberg 1929); Lambetabe [Lambelabe] (24.783°S 43.933°E) ( Dautzenberg 1929); Faux-Cap (25.567°S 45.517°E), Decary (Dautzenberg 1932); Cap Ste-Marie (25.599°S 45.137°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°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).

Figures

Fig. 4. Operculum morphology in the Chilodontidae: (A) Littorina littorea (Linnaeus, 1758), Cancale, Brittany,France,max.diameter 10.2mm (NMSA G778);(B) Euchelus asper (Gmelin,1791),Bombay, India,max.diameter 9.0 mm (NMSA J8066);(C) Granata sulcifera, Mtwalume,KZN, max.diameter 3.58 mm (NMSA V3927); (D) Clypeostoma salpinx, off Whale Rock, Eastern Cape, max. diameter 2.59 mm (NMSA C9502); (E) Danilia textilis, max. diameter 2.97 mm (holotype, NMSA E7756/ T2595); (F) Ascetostoma providentiae, off Boteler Point, KZN, max. diameter 2.52mm (NMSA S4904); (G) Herpetopoma (s.s.) instrictum, Low Isles, GreatBarrier Reef, Queensland,max.diameter 3.17 mm (AMS C.304699); (H) H. (s.s.) scabriusculum, Bradleys Head, Sydney Harbour,Australia, max. diameter 1.78 mm (AMS C.30695); (I) H. (s.l.) helix, off Umzinto, KZN, max. diameter 1.01 mm (NMSA D5427); (J) Vaceuchelus cretaceus, SE of Kosi Bay, KZN, max. diameter 2.52 mm (paratype, NMSA S4006/T2638); (K) V. natalensis, off Umzinto, KZN, max. diameter 0.99mm (NMSA D5428); (L) V.semilugubris, Réunion, max. diameter 1.28 mm (M.Jay coll’n MNHN); (M) Perrinia angulifera, off Boteler Point, KZN, max. diameter 4.44 mm (NMSA D9209); (N) P.konos, off Port Grosvenor, Eastern Cape, max. diameter 1.23 mm (NMSA E177); (O) P.stellata, Kosi Bay main coral reef, KZN, max. diameter 1.14 mm (NMSA S1985).

Fig.6. External anatomy of Chilodontidae.Schematic illustrations of the external features of the head-foot, viewed from above:(A)Ascetostoma providentiae (Melvill,1909);(B) Clypeostoma salpinx (Barnard, 1964); (C) Danilia textilis sp.n.; (D) Granata sulcifera (Lamarck,1822); (E) Perrinia angulifera (A. Adams, 1853); (F) Vaceuchelus cretaceus sp. n.; (G) V. natalensis (Smith, 1906).

Fig.65. Distribution of Granata species in the south-western Indian Ocean. Granata cumingii (circles) and G. sulcifera (triangles). Each symbol represents one or more records.

Fig.66.Granata sulcifera (Lamarck, 1822):(A–C)Santa Carolina Is., Bazaruto Archipelago,Mozambique, maximum diameter 21.5 mm (NMSA J9645); (D, E) specimen with spotted colour pattern, Durban Bay,maximum diameter 18.8 mm (NMSA 5708);(F–H) lectotype ofStomatella articulata A.Adams, 1850, maximum diameter 19.0 mm (NHMUK 1968113).

Fig. 67. Granata sulcifera (Lamarck, 1822): (A) scratch-like adult microsculpture, Inhaca Is., southern Mozambique, bar = 100 µm (NMSA L1437); (B) protoconch, Mtwalume, KZN, bar = 100 µm (NMSA B8666); (C, D) radula, Santa Carolina Is., central Mozambique, bars = 100 µm and 25 µm respectively (NMSA K3084).