Coluzea spiralis (A. Adams, 1856 )

Harasewych, M. G., 2011, The Living Columbariinae (Gastropoda: Neogastropoda: Turbinellidae) of New Zealand, Zootaxa 2744 (1), pp. 1-33 : 8-15

Coluzea spiralis (A. Adams, 1856)

( Figures 19–35 , 39 )

Synonymy:

Fusus spiralis A. Adams, 1856: 221 ; Hutton, 1880: 50; Tryon, 1881: 68, 227, pl. 85, fig. 593; Hutton, 1884: 227; Hutton, 1893: 40, pl. 6, fig. 9.

Fusus pensum Hutton, 1873: 8 ; Dell, 1956: 48; Marshall, 1996: 24.

Fusinus spiralis Suter, 1913: 357–358 ; Suter, 1915: pl. 41, fig. 4; Bucknill, 1924: 60, pl. 7, fig. 17.

Columbarium suteri E. A. Smith, 1915: 87 , pl. 1, fig. 30; Mestayer, 1916: 126, pl. 12, fig. 8.

Coluzea spiralis Finlay, 1926: 407 ; Powell, 1927: 298, pl. 34, fig. 3; Finlay, 1930b: 268; Powell, 1937: 79, pl. 8, fig. 24; Dell, 1951: 54; Dell, 1956: 47; Cernohorsky, 1977: 99–100, figs. 16–17.

Coluzea espinosa Finlay, 1930b: 268–269 .

Columbarium (Coluzea) spiralis Powell, 1979: 169 , pl. 37, fig. 2.

Type locality. Fusus spiralis , New Zealand; Fusus pensum, Kapiti Island, Cook Strait, New Zealand; Columbarium suteri , near North Cape, New Zealand, in 20–37 m; Coluzea espinosa Blue clays, Petane, New Zealand [Nukumaruan (Late Pliocene – Early Pleistocene)].

Type material. Fusus spiralis , holotype, BMNH 1951.10.16.1. Not illustrated in original publication, subsequently illustrated by Dell (1956, fig. 56) and Cernohorsky (1977: fig. 16).

Fusus pensum , lectotype, NMNZ M.000103. As noted by Marshall (1996: 24), according to Hutton’s handwritten type list, a syntype was deposited in the Canterbury Museum , but has been lost or is now unrecognizable. Dell’s (1956: fig. 57) illustration of the “Type of Fusus pensum ” constitutes a lectotype designation [Article 74.4, 5, 6 ( ICZN, 1999: 82–83)] .

Columbarium suteri , holotype, BMNH 1915.4.18.255. Illustrated by Cernohorsky, 1977: fig. 16.

Coluzea espinosa , holotype, AK 70179. Not illustrated in original publication, subsequently illustrated by Dell (1956, fig. 59).

Diagnosis. Shell moderately large, narrow, fusiform, tan-colored, with tall, acutely stepped, conical spire (spire angle 33–37°), rounded shoulder with multiple spiral threads or cords, nodulose periphery, strong cord anterior to periphery, weak anterior carina, and long, axial siphonal canal. Protoconch of ~1⅓ whorls, first whorl broad, angular, deviated. Teleoconch with up to 8 strongly convex whorls. Axial sculpture of 20–25 weak ribs most evident near periphery, with weak, irregular, brown axial bands present between adjacent ribs.

Description. Shell ( Figures 19–27 ) large (to 107.2 mm), moderately thin, with very tall, acutely stepped, conical spire (spire angle 33–37˚), small, ovate aperture, and long, stout, axial siphonal canal. Protoconch ( Figures 30– 32 ) with broad, angular, tilted first whorl initially increasing in diameter from 410 µm to 1.53 mm, then decreasing slightly to 1.48 mm after 1⅓ whorls. First whorl deviated from coiling axis of shell by ~60˚. Transition to teleoconch marked by abrupt onset of strong shoulder ( Figures 31, 32 arrows) followed within ¼ whorl by formation of fine spiral threads between suture and shoulder, and of short, broad spines along shell periphery. Spines angled posteriorly on early whorls, becoming laterally directed elements of peripheral keel by third postnuclear whorl. Teleoconch of up to 8 convex whorls. Suture ( Figure 19 , s) adpressed against anterior carina ( Figure 19 , ac) of previous whorl. Spiral sculpture of: 3–5 cords between suture and nodulose peripheral keel ( Figure 19 , pk) increasing in prominence toward periphery; 1 strong cord, which may be flanked by a weaker cord on each side, situated between periphery and weak anterior carina; 3–5 weaker cords between carina and siphonal canal; 9–14 weak cords along proximal third of siphonal canal. Fine to very fine threads on or between adjacent spiral cords. Axial sculpture of weak ribs (8 on first teleoconch whorl, 22–25 on last whorl) most conspicuous along periphery and adjacent spiral cords, especially those anterior to periphery. Growth striae weak, orthocline to slightly sinuous. Aperture broadly ovate, tapering anteriorly. Aperture narrower in juveniles, becoming more rounded with increasing shell size ( Figures 26 →23→19), deflected from shell axis by 24–28˚. Outer lip with porcelaneous glaze furrowed beneath spiral cords, most deeply beneath peripheral keel. Inner lip smooth, with thin glaze adhering tightly along parietal region, but flaring slightly at base of columella and proximal portion of siphonal canal, producing a slightly raised lamina. Siphonal canal very long (~½ shell length), narrow, axial, with stout, straight proximal half, weakly spiral distal half. Base color of shell white to cream, with irregular axial bands of light tan most conspicuous between axial ribs near periphery and along siphonal canal. Nodes along periphery, adjacent spiral cords white, internodal areas darker tan. Periostracum moderately thin, brownish yellow, lamellose. Operculum ( Figures 28, 29 ) thin, ovate, broadly rounded posteriorly, tapering anteriorly, with broadly ovate attachment area, anterior ⅓ free. Operculum weakly corrugated, with lateral edges thickened, nucleus and free edges worn, abraded.

External anatomy. Preserved, male specimen (NMNZ M.297523, Figure 26 , SL = 61.5 mm) comprised> 2½ whorls, with mantle cavity spanning ⅜ whorl, kidney ¼ whorl, and digestive gland and gonad 1+ whorls. Columellar muscle long, broad, attached to shell ⅔ whorl beyond mantle edge. Contracted foot broadly rectangular (L/W ≈ 2) with deep propodial groove along its anterior edge. Body color yellowish, without discernable pattern. Tentacles short, blunt, with large, round black eyes at their bases. Mantle edge thickened, papillated, siphonal canal long, narrow, muscular, with papillae along its proximal half.

Mantle cavity. Mantle cavity broad, deep, with organs situated as in most neogastropods. Osphradium, bipectinate, long (8 mm), narrow (L/W ≈ 2.4), with ~75 leaflets above broad osphradial nerve, ~65 leaflets below. Ctenidium 1¼ times as long, ⅔ as wide as osphradium, with ~120 thin, triangular leaflets. Ctenidium extending from just behind thickened mantle edge to rear of mantle cavity, its posterior edge adjacent to broad pericardium. Pericardium containing small auricle, large ventricle, lined by narrow nephridial gland. Hypobranchial gland transversely pleated, spanning roof of mantle cavity, thicker along right side, which partially covers rectum. Renal organ fairly short, with primary and secondary lamellae not interdigitated [termed Meronephridiens ( Perrier 1889)], forming right rear wall of mantle cavity.

Alimentary system. Proboscis extremely long (95 mm, ~1½ shell length), narrow (<1 mm in diameter), greatly convoluted when retracted into thin-walled, non-evertable proboscis sheath broadest near rear of cephalic haemocoel. Mouth small, situated at tip of rounded distal end of proboscis, opening into small buccal cavity. Buccal mass narrow, slightly shorter that radula. Radular ribbon ( Figures 33, 34 ) 5.25 mm long, comprised of 135 rows of teeth. Rachidian teeth 134.6 µm wide, with a basal plate with U-shaped central section flanked by broad, rectangular lateral edges. Central of 3 cusps (54.1 µm), 1¼ times as long as flanking cusps, its distal end resting in dimple in base of corresponding cusp of posteriorly adjacent tooth. Inner edge of right cusp with serrated edge ( Figure 34 , arrow), an anomalous feature not previously seen in radulae of Coluzea spiralis (see Harasewych, 1991:fig. 21) or any other species of Columbariinae . Lateral teeth, each with a single, long (98 µm), recurved, scythe-shaped cusp, attached to radular ribbon by thin band along widest portion (54 µm) of basal plate ( Figure 33 , bp). Anterior esophagus narrow, extending length of proboscis, then anteriorly from rear of proboscis sheath, expanding slightly to form small valve of Leiblein just prior to passing through nerve ring. Esophagus runs posteriorly beneath broad, flattened, posteriorly tapered gland of Leiblein, leaving cephalic hemocoel to enter U-shaped stomach lining anterior portion of digestive gland. Intestine enters mantle cavity, running anteriorly along its right side to form a narrow, tubular rectum, terminating in simple anus and small anal gland. Two ascinous salivary glands, the right gland larger, compacted in anterior end of cephalic hemocoel, emptying laterally into the buccal mass via long, narrow ducts passing beneath dorsal folds of esophagus.

Male reproductive system. Testes line inner wall of digestive gland, forming a thick- walled, highly convoluted seminal vesicle, embedded within anterior wall of digestive gland. Testicular duct runs from its anterior end, along pericardium, entering mantle cavity along its right wall, running ventral and parallel to the rectum before descending to floor of mantle cavity. Duct joins muscular, open groove lined with prostatic tissue that leads to base of broad, dorsoventrally flattened penis, along its outer edge to tip of a long, narrow, tapering papilla.

Female reproductive system. Partially preserved female specimen (NMNZ M.061102, SL = 79.3 mm) with oviduct extending anteriorly from ovary, embedded in right wall of digestive gland, running along kidney and pericardium before entering rear of mantle cavity. Pallial oviduct linear, consisting of a thin-walled albumen gland, a cylindrical, glandular capsule gland, and an anterior, muscular bursa copulatrix with a terminal female opening. Ingesting gland not found, possibly due to poor preservation.

Material examined. NORTH ISLAND: 1 NMNZ M.086706, N of North Cape [34°20.0’S, 173°06.6’E], 163– 168 m, 27 Jan. 1981, RV TANGAROA; 1 NMNZ M.074903, off Parengarenga Harbour [34°32.0’S, 173°06.0’E], 102– 93 m, foraminiferal ooze, 27 Jan. 1981, RV TANGAROA; 2 NMNZ M.076035, NW of Ahipara [34°50.0’S, 172°46.1’E], 90 m, sandy mud, 10 Jan. 1981, RV TANGAROA; 1 NMNZ M.059528, off Ninety Mile Beach [34°59.4’S, 172°35.5’E], 156–160 m, 19 Nov. 1978, RV JAMES COOK; 2 NMNZ M.043632, Off Takou Bay , N of Kerikeri [35°10’S, 174°11’E], 80 m, mud, sand and sponge bottom, 16 Feb. 1974, RV ACHERON; 3 NMNZ M.035884, W of Poor Knights Islands [35°32.0’S, 174°41.0’E], 121– 113 m, mud, 15 Feb. 1974, RV ACHERON ;

2 NMNZ M.066727, SE of Alderman Islands [37°00.8’S, 176°12.3’E], 178–248 m , mud, 23 Jan. 1979, RV TAN- GAROA; 7 NMNZ M.060111 and 5 NMNZ M.066561, NW of Mayor Island [37°11.5’S, 176°10.0’E], 198–273 m , mud, 22 Jan. 1979, RV TANGAROA; 1 NMNZ M.060104, c. 11 km NW of Mayor Island [37°13.1’S, 176°07.8’E], 119–169 m , mud, 22 Jan. 1979, RV TANGAROA; 2 USNM 896320 [37°35’S, 178°46’E], 128–146 m , 28 May 1966; 4 NMNZ M.060581, c. 9 km N of Motuhora Island [37°45.8’S, 177°00.8’E], 72–84 m , mud, 20 Jan. 1979, RV TANGAROA; 4 NMNZ M.297523, off Opotiki [37°53’S, 177°17’E], 37 m , 1967, RV IKATERE; 2 NMNZ M.060132, E of Mahia Peninsula [39°17.2’S, 178°12.7’E], 258–306 m , mud, 27 Jan. 1979, RV TANGAROA; 4 NMNZ M.061102, E of Mahia Peninsula [39°18.7’S, 178°04.6’E], 127–134 m , mud, 27 Jan. 1979, RV TANGAROA; 2 NMNZ M.053738, off Foxton [40°30.5’S, 174°53.5’E], 101 m , 1 Mar. 1976, RV ACHERON; 4 NMNZ M.052731, off Foxton [40°33.5’S, 174°59.5’E], 86–88 m , mud, 2 Mar. 1976, RV ACHERON; 1 DNMH 73077, Whale Island , 137 m ; 1 DMNH 48393 , Off Kawau Id.; 1 USNM 681696 , off Plate Island , Bay of Plenty, 73 m , 1968; 1 USNM 22859 , New Zealand. COOK STRAIT: 1 NMNZ M.049935, Midway between Cape Jackson and Mana Island , Cook Strait narrows [41°02’S, 174°33’E], 256– 186 m , rock bottom, 6 Mar. 1976, RV ACHERON. SOUTH ISLAND: 7 NMNZ M.053084, c. 24 miles NE of Stephens Island [40°24’S, 174°17’E], 110 m , mud, 4 Mar. 1976, RV ACHERON; 1 NMNZ M.053490, N of base of Farewell Spit [40°28’S, 172°48’E] in 51 m , on sand. 10 Mar 1976, RV ACHERON; 6 NMNZ M.053432, N tip of Farewell Spit [40°31.5’S, 173°03’E], 44 m , 10 Mar. 1976, RV ACHERON; 3 NMNZ M.059020, off Kahurangi Shoals , NW of Kahurangi Point [40°42.5’S, 172°07’E], 91 m , stones and subfossil shells, 10 Mar. 1976, RV ACHERON; 1 NMNZ M. 297041, between Titi Island and Alligator Head [40°57’S, 174°08’E], 84–88 m , mud and shell bottom, 30 Aug. 1975, RV ACHERON; 1 USNM 877100 , off Canterbury, in 366 m.

Additional material at NMNZ. NORTH ISLAND: 1 NMNZ M.148196, off Tom Bowling Bay [34º19.43'S, 172º54.57'E], 94 m, 26 Jan. 1999 , RV KAHAROA; 1 NMNZ M.153113, off Cape Reinga [34º22.80'S, 172º24.60'E], 121 m, bryozoa and shell bottom, 2 Feb. 1981 , RV TANGAROA; 4 NMNZ M.297040, E of North Cape [34º 25.00' S, 173º 13.10' E], 257–327 m, foraminiferal sand, 27 Jan. 1981 , RV TANGAROA; 1 NMNZ M.002522, 15’S of Great Island , Three Kings Islands [34°25.1’S, 172°09.0’E], 179 m, 29 May 1914 , SS HINEMOA; 1 NMNZ M.153114, Great Exhibition Bay [34º33.60'S, 173º 04.90'E], 66 m, 27 Jan. 1981 , RV TANGAROA; 1 NMNZ M.153115, NW of Ahipara [34º50.00'S, 172º46.10' E], 90 m, sandy mud, 10 Jan. 1981 , RV TANGAROA; 2 NMNZ M.083862, off Ninety Mile Beach [34º53.00'S, 172º55.00'E]; 1 NMNZ M.025800, off Doubtless Bay [34º55.00'S, 173º30.00'E], 47 m, 18 Sep. 1971 ; 1 NMNZ M.035741, off Cape Brett [35º08.00'S, 174º12.50'E], 80 m, mud, 16 Feb. 1974 , RV ACHERON; 8 NMNZ M.028457, NE of Ninepin Rock , Bay of Islands [35º08.80'S, 174º10.90'E], 79 m, 01 Dec. 1971 , MV KOKINGA; 2 NMNZ M.030757, NE of Ninepin Rock , Bay of Islands [35º08.80'S, 174º10.30'E], 75 m, 1 Dec. 1971 , MV KOKINGA; 1 NMNZ M.112109, off Deep Water Cove , Bay of Islands [35º11.70'S, 174º17.40'E], 55 m, Mar. 1973 ; 4 NMNZ M.035093, N of Poor Knights Islands [35º22.00'S, 174º43.00'E], 146 m, sandy mud, 15 Feb. 1974 , RV ACHERON; 1 NMNZ M.016233, SE of Poor Knights Islands [35º38.00'S, 174º56.00'E], 165 m, 20 Nov. 1962 , RV IKATERE; 1 NMNZ M.297068, off Peach Cove , Bream Bay [35º 53.367'S, 174º 33.00' E], 77 m, 26 Feb. 1998 ; 1 NMNZ M.021045, off Miner's Head , Great Barrier Island [35º55.00'S, 175º20.50'E], 73–82 m; 1 NMNZ M.153116, Centre Reef , N of Little Barrier Island, Hauraki Gulf [36º00.00'S, 175º07.50'E], 73 m, May 1967 ; 3 NMNZ M.083865, off Hauraki Gulf; 1 NMNZ M.090206, Hauraki Gulf , 50–60 m, 1986; 1 NMNZ M.083864, off Miner's Head , Great Barrier Island [36º04.00'S, 175º21.00'E], 1950; 6 NMNZ M.130835, SE of Te Arai Point , Mangawhai , N of Warkworth [36º10.50'S, 174º42.00'E], 35–40 m, sand and shell suction dredged for beach replenishment at Mission Bay, Auckland, 3 Mar. 1996 ; 1 NMNZ M.083863, off Horn Rock , Great Barrier Island [36º15.00'S, 175º11.00'E]; 1 NMNZ M.016220, N of Cuvier Island [36º17.00'S, 175º46.00'E], 110 m, 25 Nov. 1962 , RV IKATERE; 1 NMNZ M.136053, Omaha Bay , N of Auckland [36º20.48'S, 174º49.98'E], 32 m, mud, 30 Nov. 1995 ; 1 NMNZ M.136133, Omaha Bay , N of Auckland [36º20.87'S, 174º50.98'E], 34 m, mud, 30 Nov. 1995 ; 1 NMNZ M.021048, off Whale Rock , Mercury Islands, Whitianga [36º42.00'S, 175º54.50'E], 73–91 m, 1967; 1 NMNZ M.021049, off Mercury Bay , Whitianga [36º47.00'S, 175º50.00'E], 73 m, 11 Apr.1967 ; 1 NMNZ M.153111, N of the Aldermen Islands [36º47.50'S, 176º 0.00'E], 108–113 m, mud, 24 Jan. 1981 RV TANGAROA; 1 NMNZ M.044393, off the Aldermen Islands [36º59.00'S, 176º12.00'E], 165 m, 10 Jun. 1970 , RV IKATERE; 1 NMNZ M.039421, off Slipper Island [37º01.00'S, 175º57.00'E], 37 m, 8 Apr. 1970 , RV IKATERE; 1 NMNZ M.039453, off Slipper Island [37º03.00'S, 175º59.00'E], 55 m, 2 Dec. 1969 , RV IKATERE; 3 NMNZ M.083861, off Mayor Island, 91 m; 1 NMNZ M.119093, off Lottin Point [37º26.10'S, 178º13.10'E], 255 m, 29 Jan. 1995 , RV KAHAROA; 1 NMNZ M.039528, off Plate Island [37º35.00'S, 176º36.00'E], 146 m, 9 Jun. 1970 , RV IKATERE; 1 NMNZ M.039537, off Plate Island [37º35.00' S, 176º36.00'E], 155 m, 7 Apr. 1970 , RV IKATERE; 1 NMNZ M.083866, off Plate Island ; 5 NMNZ M.153112, off East Cape , southeastern slope of Ranfurly Bank [37º38.50'S, 178º56.40'E], 153 m, mixed Recent and fossil fauna, 22 Jan. 1981 , RV TANGAROA; 3 NMNZ M.112411, Between Motuhora and White Islands [37º40.20'S, 178º53.60'E], 117 m, mud, shell and pumice, 2 May 1990 , RV AKADE- MIK ALEXANDER NESMEYANOV; 3 NMNZ M.039988, off Whale Island (Motuhora) [37º51.00'S, 176º58.00'E], 18– 55 m, 1967; 2 NMNZ M.083709, off Whale ( Motuhora ) Island; 3 NMNZ M.088299, SW of Kawhia Harbour [38º21.50'S, 174º17.80'E], 83 m, 13 Jan. 1981 , RV TANGAROA; 1 NMNZ M.028428, off New Plymouth [39º07.00'S, 173º48.00'E], 97 m, 21 May 1964 ; 1 NMNZ M.050583, S of Waverley [40º10.00'S, 174º40.00'E], 82 m, bryozoa and shell bottom, 1 Mar. 1976 , RV ACHERON; 2 NMNZ M.053335, W of Rangitikei River mouth [40º16.00'S, 174º58.50'E], 75 m, 2 Mar. 1976 , RV ACHERON; 1 NMNZ M.052615, W of Rangitikei River mouth [40º18.00'S, 174º59.00'E], 82 m, mud, 2 Mar. 1976 , RV ACHERON; 2 NMNZ M.053811, NW of Manawatu River mouth [40º22.50'S, 174º59.50’E], 86 m, mud, 2 Mar. 1976 , RV ACHERON; 2 NMNZ M.052912, N of Kapiti Island [40º38.00'S, 174º54.50'E], 91 m, 1 Mar. 1976 , RV ACHERON; 1 NMNZ M.000103, Kapiti Island ; 1 NMNZ M.048946, off Whakatane, 1966; 1 NMNZ M.083867, off Cape Runaway. COOK STRAIT: 1 NMNZ M.013779, Nicholson Canyon , off Turakirae Head [41º28.50'S, 174º 50.00'E], 274 m, mud, gravel and shell bottom, 21 Mar. 1954 , MV ALERT. SOUTH ISLAND: 2 NMNZ M.090257, NE of Cape Farewell [40º06.00'S, 172º12.00'E], 102–137 m, 11 Nov. 1987 , FV UNZEN MARU; 1 NMNZ M.053752, NE of Stephens Island [40º31.00'S, 174º15.00'E], 117– 119 m, mud, 4 Mar. 1976 , RV ACHERON; 1 NMNZ M.053040, NE of Stephens Island [40º33.00'S, 174º07.00'E], 132 m, mud, 4 Mar. 1976 , RV ACHERON; 1 NMNZ M.032979, W of Cape Farewell [40º40.00'S, 171º33.00'E], 265 m, 15 Jan. 1971 , RV JAMES COOK; 4 NMNZ M.053599, off Seal Bay , S of Kahurangi Point [40º52.00'S, 172º04.00'E], 64 m, 10 Mar. 1976 , RV ACHERON; 2 NMNZ M.055072, Admiralty Bay [40º54.00'S, 173º56.00'E], 44 m, mud, 5 Mar. 1976 , RV ACHERON; 1 NMNZ M.050558, NW of Wekakura Point, between Kahurangi Point and Heaphy River [40º54.00'S, 172º04.00'E], 55 m, sandy mud, 10 Mar. 1976 , RV ACHERON.

Fossil specimens at NMNZ. NORTH ISLAND: 4 NMNZ M.001418, Castlecliff , Wanganui , Lower Beds, Pleistocene (Castlecliffian), 1924; 2 NMNZ M.017770, Castlecliff , Wanganui , Blue mudstone. Pleistocene (Castlecliffian) 26 Jan 1909 ; 13 NMNZ M.026406, Castlecliff , Wanganui, Pleistocene (Castlecliffian); 1 NMNZ M.026407, Shakespeare Cliff , Wanganui ( GS206 ), Pleistocene (Castlecliffian); 2 NMNZ M.026408, Castlecliff , Wanganui , Blue mudstone. Pleistocene (Castlecliffian). 29 Jul 1909 ; 1 NMNZ M.028184, Castlecliff , Wanganui, Pleistocene (Castlecliffian); 4 NMNZ M.071116, Buttress , Tainui Shellbed , Wanganui, Pleistocene (Castlecliffian) 17 Jul 1974 .

Distribution ( Figures 35 , 39 ). Coluzea spiralis inhabits mud, sand and rubble bottoms on the mid- to outer continental shelf and uppermost continental slope along the northern, western, and northeastern coasts of North Island, the Cook Strait, and off Cape Farewell, South Island. Excluding one questionable 366 m [“200 fm”] record originating from a private collection, the species has been reported from depths ranging between 32 and 327 m, with a mean station depth [n = 74] of 111.9 m, and a confirmed bathymetric range of 32– 274 m. The distribution of Coluzea spiralis is confined to the area north of the biogeographic disjunction south of the Cook Strait at 42°S latitude that has been documented for several shallow water species (see Apte & Gardner 2002: fig. 1; Ayers & Waters 2005: fig. 1).

Remarks. Coluzea spiralis is the shallowest dwelling member of the subfamily Columbariinae , with the majority of records sampled from mid to outer continental shelf depths. Nearly all of the shallowest records (<50 m) are from the northern coast of North Island, an area with coastal upwelling and bottom water temperatures approaching 15°C ( Longdill & Healy 2008: figs. 5, 6). Deepest records (> 200 m) are from steep slopes near the various capes at the corners of North Island, the Mahia Peninsula and Cape Farewell on South Island. The fossil record of Coluzea spiralis extends to the Nukumaruan (Pliocene / Pleistocene) and Castlecliffian (Pleistocene) deposits on North Island ( Beu et al. 1990: 356).

Dell (1956: 48) commented on the occurrence of northern and southern forms of Coluzea spiralis . The northern form, which he considered to be exemplified by the holotype of Fusus spiralis , has a single, prominent keel on the last whorl. A southern form, occurring from Kapiti Island southward and exemplified by Suter’s (1913: pl. 41, fig. 4) illustration (based on a photograph of the holotype of F. pensum ), is distinguished by the presence of two keels on the antepenultimate and penultimate whorls. Finlay (1930b: 269) speculated that perhaps Suter’s drawing was based on a fossil (Nukumaruan) specimen, and went on to describe that phenotype as C. espinosa . Dell (1956: 48) reported that Castlecliff (Castlecliffian) fossils have a single keel and are indistinguishable from the northern form, and went on to note that specimens from the northern portion of the range may also develop two keels (see Figures 19, 23, 27 ), but do so at a larger size than southern specimens.

Coluzea spiralis may be discerned from other Recent New Zealand species of Coluzea by its stout shell, tall spire with stepped, angular whorls, its strongly angular protoconch, nodular peripheral keel, weak anterior carina, and the presence of a strong, often nodular spiral cord between the peripheral keel and anterior carina. Brownish pigment is often present in areas between nodules along the periphery and the subperipheral spiral cord, especially in larger specimens. The peripheral keel is more posteriorly situated along the whorls than in other living species of New Zealand Coluzea . The aperture is more ovate and less rounded than in C. wormaldi or C. altocanalis . The radula of C. spiralis differs from that of other Recent species of New Zealand Coluzea in having the three cusps span a proportionally narrower section of a broader basal plate.

Although Coluzea spiralis is the only species of Columbariinae inhabiting the western coast of New Zealand, its geographic range overlaps with that of Columbarium viridicum and Coluzea wormaldi along the northern shore of North Island, and with Coluzea wormaldi and Coluzea altocanalis along the eastern shore of North Island. However, the confirmed bathymetric range of C. spiralis [32–274 m] does not overlap with the ranges of Columbarium veridicum [622–818 m], Coluzea wormaldi [337–685 m]; or C. altocanalis [403–984 m].

Figures

FIGURES 19–34. Coluzea spiralis (A. Adams, 1856). 19, Apertural 20, lateral, 21, dorsal, and 22, apical views of specimen from NMNZ M.074903, off Parengarenga Harbour North Island [34°32.0’S, 173°06.0’E], in 102–93 m, on foraminiferal ooze. 23, Apertural, 24, lateral and 25, dorsal views of specimen from NMNZ M.059528, off Ninety Mile Beach, North Island [34°59.4’S, 172°35.5’E], in 156–160 m. 26, Apertural view of a male specimen from NMNZ M.297523, off Opotiki, North Island [37°53’S, 177°17’E], in 37 m. 27, Dorsal view of shell from NMNZ M.060132, E of Mahia Peninsula, North Island [39°17.2’S, 178°12.7’E], in 258–306 m, on mud. 2 cm scale bar applies to figures 19–27. 28, Inner and 29, outer surfaces of operculum of specimen in figure 26. 30–31, Lateral and 32, apical views of protoconch of specimen from NMNZ M.035884, W of Poor Knights Islands, North Island [35°32.0’S, 174°41.0’E], in 121–113 m, on mud. Arrows indicate transition to teleoconch. 33–34. Radular ribbon of specimen in figure 26. 33, oblique view of radula, 34, dorsal view with lateral teeth expanded. 100 mm scale bar applies to figures 33–34. Arrow indicates atypical denticles on cusp or rachidian tooth. ac, anterior carina; bp, basal plate of lateral tooth; pk, peripheral keel; s, suture.

FIGURES 35–38. Geographic distributions of species of New Zealand Coluzea. 35, Coluzea spiralis (A. Adams, 1856). The type locality is New Zealand. 36, Coluzea wormaldi Powell, 1971. 37, Coluzea altocanalis Dell, 1956. 38, Coluzea mariae Powell, 1952. = type locality. = station records.

FIGURES 39–42. Bathymetric distributions of species of New Zealand Coluzea. 39, Coluzea spiralis (A. Adams, 1856). 40, Coluzea wormaldi Powell, 1971. 41, Coluzea altocanalis Dell, 1956. 42, Coluzea mariae Powell, 1952.