Siphonaria normalis Gould 1846

Siphonaria normalis Gould, 1846 (Figs 31A–I, O, S–U, 32A–D) Siphonaria normalis Gould 1846: 12 (type locality: ‘Sandwich Islands’ [Hawaii]).— Gould 1848: 154; 1852: 359; H. Adams & A. Adams 1855 (in 1853–1858): 271; Gould 1856: 13, fig. 468a–b; Hanley 1858b: 152; H. Adams & A. Adams 1863: 271; Paetel 1883: 178; 1889: 429; Pilsbry 1920b: 379; Edmondson 1946: 188, fig. 102a; Hubendick 1946: 63; 1947b: 2, figs 3a–i, 4a–i, 5, 8–10; Morris 1952: 206, pl. 40, fig. 17; Johnson 1964: 116; Cook 1969: 679; Cernohorsky 1972: 210, pl. 60, fig. 2; Cook 1976: 34; Galindo 1977: 416; Coles 1981: 11; Trew 1983: 8; Jenkins 1984: 117; White & Dayrat 2012: 62, 2014: 266; Gonzàlez-Wevar et al. 2018: 5, fig. 1; Krug et al. 2022: 5. Siphonaria normalis var. lirata — Pilsbry 1920b: 37980 (not S. lirata Reeve, 1856). Planesiphon soranus Iredale 1940: 441, fig. 20, 21 (type locality: Townsville, Queensland).— White & Dayrat 2012: 67. Siphonaria ‘ normalis group, unit 14’ (in part)— Dayrat et al. 2014: 267, fig. 3 P–S. Siphonaria cf normalis (unit 52)— Ossenbrügger et al. 2023: fig. 2c–o. Material examined. Type material. Syntype of Siphonaria normalis Gould, 1846 from Hawaii, United States, North Pacific Ocean; coll. Carpenter (USNM 15346, Fig. 31A). Lectotype of P. soranus Iredale, 1940, present designation, from Townsville Qld, Australia; coll. 1929 (AM C.103709, Fig. 31D). Seven paralectotypes, same data as lectotype (AM C.124989). Two paralectotypes, same data as lectotype (MV F13840). Other, non-type material. Marquesas: Baie des Vierges, Fatu Hiva, 10°27,84’S, 138°39,97’W (MNHN IM-2013-74897 p [M570, SK508], IM-2013-74898 p [M569], IM-2013-74899 p [M565]). Hawaii, Oahu: N end Waikiki Beach, 21°15.743’N, 157°49.307’W (AM C.584887 p [M293]); N end Waikiki, 21°16.646’N, 157°50.048’W (AM C.585622 5p, C.584886 p [SK210], C.585930 p [SK209]). Maui: Hanakao’o Beach, 20°54.564’N, 156°41.310’W (AM C.585929 p [M296]); Makalua-puna Point, 21°54.586’N, 156°41.338’W (AM C.585555 3p); Ho’okipa Beach, 20°56.029’N, 156°21.411’W (AM C.584892 p [SK212]). Guam: Umatac Bay, N end, 13°17.917’N, 144°39.494’E (AM C.585325 p C.584869 p [SK142], C.585493 p [M347], C.585494 p [M348]); Pago Bay, below UoG Marine Lab, 13°25.645’N, 144°47.927’E (AM C.585996 10+p); Tanguisson Beach, S end, 13°32.549’N, 144°48.443’E (AM C.585994 2p, C.584873 p [M343], C.584874 p [M439]). Fiji, Viti Levu: Vuda Point Marina seawall, 17°40.878’S, 177°23.009’E (AM C.585970 20+p, C.584866 p [SK115 protoconch C9], C.584867 p [M286], C.584868 p[M287]). PNG: Sek Is., 05°04,7’S, 145°48,9’E (MNHN IM-2013-13133 p [M558], IM-2013-13135 p [M562]); Wonad I., 05°08,1’S, 145°49,3’E (MNHN IM-2013-15280 p [M563]); Riwo waters, 05°08,9’S, 145°48,2’E (MNHN IM-2013-15250 p [M555] IM-2013- 15251 p [M554]). NC, Lifou: We Baie de Chateaubriand E coast (AM C.585396 10p, C.584946 p [M388], C.584947 p [M389]). S of Pouebo 20°25.950’S, 164°39.251’E NC04-2 (AM C.585006 p [M381], C.585007 p [M382]); Bonhomme de Bourail, La Roche Percee, 21°36.487’S, 165°27.423’E (AM C.585013 p [M370], C.585014 p [M371]); Presqu’ile de Ouano La Foa, 20°51.434’S, 165°48.479’E (AM C.595911 6p). Australia, Qld: Umagico, 10°53.125’S, 142°20.799’E (AM C.585178 p [SK195]); Capt Billy Landing, 11°38.019’S, 142°51.472’E (AM C.585415 10+p, C.584792 p [M180], C.584793 p [M181], C.584794 p [M402]); S of Bathurst Head, 14°17.583’S, 144°11.845’E (AM C.585348 10p); Lizard Is, bch rock, 14°40.730’S, 145°26.838’E (AM C.585643 5p); Lizard Is, 14°40.908’S, 145°27.007’E (AM C.585566 4p, C.585175 p [M030]); Cape Kimberley, 16°16.535’S, 145°28.737’E (AM C.585720 9p, C.585168 p [M041], C.585169 p [M394], C.585170 p [M395], C.585171 p [M397]); Pebbly Beach Yule Reef Trinity Bay, 16°35.031’S, 145°30.823’E (AM C.585703 8p); Gribble Pt Mission Bay Yarrabah, 16°53.781’S, 145°51.852’E (AM C.585347 p); Mourilyan Harbour, 17°35.951’S, 146°07.583’E (AM C.585411 10+p, C.585155 p [M012], C.585156 p [SK123]); W side Kissing Pt Townsville, 19°14.332’S, 146°48.040’E (AM C.585672 6p, C.585146 p [M083], C.585147 p [M186], C.585148 p [M187], C.585149 p [M188], C.585931 p [SK197], C.585932 p [SK075]); Slade Pt Mackay, 21°03.813’S, 149°13.527’E (AM C.585501 2p). Gulf of Carpentaria: Mutee Head, 10°54.682’S, 142°15.204’E (AM C.585416 10p); Weipa, 12°37.795’S, 141°51.853’E, Q52-1 (AM C.585450 12p); Sweers Is: 17°07.029’S, 139°35.805’E (AM C.585645 5p), 17°07.413’S, 139°35.816’E, Q56-2 (AM C.585350 p), Inspection Pt, 17°08.471’S, 139°36.868’E (AM C.585417 15+p). NT: Cape Wirawawoi Nhulunbuy, 12°09.513’S, 136°46.904’E (AM C.585530 20+p); Sandy Is Pt 11°07.862’S, 132°11.187’E (AM C.585406 10+p); Smith Pt 2, 11°07.466’S, 132°08.538’E (AM C.585636 5p); Luxmore Hd Melville Is, 11°20.639’S, 130°23.149’E (AM C.585349 7p); Nightcliff Darwin, 12°22.836’S, 130°50.402’E (AM C.585977 10+p); Cox Peninsula, 12°24.824’S, 130°40.921’E (AM C.585978 10+p); N of Native Pt Dundee Bch, 12°42.182’S, 130°20.881’E (AM C.585668 6p); Native Pt oyster reef Dundee Bch, 12°42.906’S, 130°20.653’E (AM C.585413 8p); Native Pt reef Dundee Bch, 12°42.981’S, 130°20.807’E (AM C.585381 10p). Taxonomic remarks. Siphonaria normalis was subsequently figured by Gould (1856: 13, pl. 30, figs 468, 468a–b). The syntype corresponds well with this figure in shell profile, dimensions and colouration. The description of P. soranus does not contain an original type designation. The figured specimen of P. soranus (Iredale, 1940: pl. 34, figs 20–21) is herein designated as the lectotype for the stabilisation of the name (AM C.103709). Our delineation of this species is based on comparative analyses of the morpho-anatomy and mitochondrial genetics of freshly collected topotypes of S. normalis and P. soranus and geographic series of additional specimens (Table S1). We stablish P. soranus as a junior synonym of S. normalis. Pilsbry (1920b: 379) erroneously considered S. amara, S. nuttallii, S. lirata, S. crebricostata and S. normalis f. chirura as synonyms of S. normalis. This was followed by Cernohorsky (1972: 210). Dayrat et al. (2014: 267) recognised several independent molecular units within the normalis group (i.e., unit 12 from Thailand, unit 13 from Singapore, and unit 14 from Hawaii). These units are herein recognised as three distinct species, S. radiata (unit 12), S. costellata sp. nov. (unit 13), and S. normalis (unit 14). External morphology. Foot sole, foot wall, mantle, cephalic folds and pneumostomal lobe evenly pale grey/cream, paler at edge foot/wall; blotches of black pigmentation on centre of cephalic folds, faintly on foot wall; mantle narrower than width of foot wall, non-translucent, covers exposed inner shell lip, edge thickened, lobed, vertical bands of black pigmentation aligned with shell rib interstices; genital pore indistinct, located on foot wall to right anterior of right cephalic fold; two small black epithelial eye spots centralised on two centrally touching cephalic folds; pneumostomal lobe long under the mantle between the right ADMs. Shell (Figs 31A–I, O, S–U; Table S9). Small sized (max sl mean = 12.56 mm, SD = 2.02 mm, n = 13), circular ovate; height medium to tall; apex offset posterior and weakly left, apical sides straight to weakly convex, shell edge; protoconch direction heterostrophic, initially hooked (n = 4; Fig. 31S–T), shell whorl dextral; growth striae prominent, radial colour banding often present, shell thickness thick; rib count (mean = 33, SD = 0.87, n = 13), primary ribs distinct, white to pale, usually radially evenly spread, fairly straight, broaden to and align with fairly flat faintly scalloped shell edge; ribs may be raised or flat,; 1–2 interspersed pale white finer secondary ribs, rib interstices darker; paired primary ribs on siphonal ridge, no more prominent than other primary ribs. Interior shell margin varies from pale tan to dark brown; white rays extend from the shell lip to over the shell margin fading to the spatula, align under primary/secondary ribs, spatula varies from pale yellow (Fig. 31E), pale tan (Fig. 31B) to dark chocolate brown (Fig. 31F); siphonal groove distinct, paler than shell margin or spatula; ADM scar distinct, CMS straight, paler than shell lip; thickening of shell lip common, margin becomes whitened (Fig. 31E). Reproductive system (Figs 32A, C; n = 9). Positioned within coelom under the respiratory cavity, hermaphroditic glands positioned to posterior against right foot wall and over foot sole, epiphallic parts positioned to anterior between BM and RAM; GP small, singular, positioned through foot wall behind right cephalic fold, GA small, prominent; AO very small, short, narrow, blunt, slightly bent centrally, joins to top of GA in conjunction with ED; ED short, broad, joins to back side of GA alongside AO; single short broad blunt flagellum (F1), longer than and same width as ED, join of F1 to ED indistinct, marked by connection of very small white folded EG; AO, GA and ED all muscular white tissue; BD and CD connect closely side-by-side into GA between connections of ED and AO, both ducts narrow smooth featureless, pass together through RAM connecting into MG (BD above CD), BD longer than CD, often loop immediately in front of BC; BC large to medium, spherical, embedded in folds of MG, test translucent; SV embedded on left side of AG; HD short, thick coils, links AG to smaller yellowish granular HG; MG and AG folded, soft white tissue; sides match curvature of inner foot wall on right posterior of coelom; outer edge of MG lobed. Spermatophore (Figs 32B, D). Broad head with short flagellum (length = 2.47 ± 0.195 mm, n = 3); head section cylindrical, bulbous, centrally bent, rounded tip; test thin, smooth, featureless, translucent encasing a white opaque central core; short tapering section (often looped) merges head to filamentous flagellum; head slightly shorter, wider than translucent flagellum (head length = 1.23 ± 0.45 mm, flagellum length = 1.24 ± 0.34 mm, SPM head ~ 49% of total lenght; head width = 110 ± 13 μm, flagellum w