Siphonaria denticulata Quoy & Gaimard 1833

Siphonaria denticulata Quoy & Gaimard, 1833 (Figs 19G–L, R, 20D–H) Siphonaria denticulata Quoy & Gaimard 1833: 340, pl. 25; figs 19– 20 (type locality: Nouvelle-Hollande, port Western [Western Port, French Island, Vic, Australia]).— Lamarck 1836: 559, 1839: 206; Reeve 1842: pl. 138, fig. 2; Catlow & Reeve 1845: 100; Menke 1852: 38; H. Adams & A. Adams, 1855 (in 1853–58): 271; Reeve 1856 pl. 1, species 4; Hanley 1858b: 151; H. Adams & A. Adams 1863: 271; Angas, 1867: 232; Paetel 1873: 117; 1875: 92; Tenison Woods 1878b: 99; Paetel 1883: 178; Etheridge 1887: 29; Henn & Brazier 1894: 178; Brazier 1889; 25; Paetel 1889: 428; Whitelegge 1889: 117; Adcock 1893: 11; Hedley 1909; 369; 1918: M96; McAlpine 1952: 41; Ewers 1961; Galindo 1977: 416; Crease 1980: 38; Jenkins 1981: 2; Hochlowski et al. 1983: 7413; Quinn 1983: 81; Trew 1983: 2; Jenkins 1983: 29; 1984: 113; Hochlowski & Faulkner 1984: 3839; Short & Potter 1987: 122, pl. 60, fig. 3; Jansen 1995: 89; Davey 1998: 118, text-fig.; Grove et al. 2006: 60; White & Dayrat 2012: 62; Colgan & da Costa 2013: 74. Siphonaria denticula Anton 1838: 26 (invalid; incorrect subsequent spelling of S. denticulata). Siphonaria atra — Angas 1867: 233; Whitelegge 1889: 117. Siphonaria cochlearformis Whitelegge 1889: 117; Hedley 1918: M96 (invalid; incorrect subsequent spelling, not S. cochleariformis Reeve, 1856). Ellsiphon marza Iredale 1940: 438 pl. 34, figs 1, 2 (type locality: Keppel Bay, Queensland).— McAlpine 1952: 42; White & Dayrat 2012: 57, 60, 63. Siphonaria scabra — Iredale 1940: 438 (not S. scabra Reeve, 1856). Siphonaria marza — Hubendick 1946: 61, pl. 5, fig. 27, 28; Galindo 1977: 416. Siphonaria (Ductosiphonaria) diemenensis var. denticulata — Hubendick 1946: 38, pl. 2 fig. 17. Ellsiphon denticulatus — McAlpine 1952: 41, fig. 1; Iredale & McMichael 1962: 82. Siphonaria (Siphonaria) currumbinensis Hubendick 1955: 126– 136, figs 4–8, pl. 1, figs 1–6 (type locality: Currumbin, Qld, Australia).— White & Dayrat 2012: 62, 70. Ductosiphonaria denticulata — Cotton 1959: 411. Siphonaria alternicosta — Cotton 1959: 411 (not S. alternicosta Potiez & Michaud, 1838). Siphonaria ‘ atra group, unit 33’— Dayrat et al. 2014: 264, fig. 5 J (not S. atra Quoy & Gaimard, 1833). Material examined. Type material. Lectotype of Siphonaria denticulata Quoy & Gaimard, 1833, present designation, from ‘partie sud de la Nouvelle-Hollande, au port Western, et probablement aussi au port du Roi-Georges’ [southern part of Australia, at Western Port, Vic, Australia, and probably also at King George Harbour; see comments below] (MNHN IM 2000-35951, Fig. 19G). Two paralectotypes, same data as lectotype (MNHN IM 2000-5062). Probable holotype of Ellsiphon marza Iredale, 1940 from Keppel Bay [Qld, Australia]; coll. H. Bernhard, 1940 (AM C.103715, Fig. 19K). Probable paratypes of Ellsiphon marza Iredale, 1940. Similar data to holotype label (refer Taxonomic remarks) (MV F13843, 3 d); Tampian Heads, Yeppoon, Keppel Bay; coll. H. Bernhard Dec 1938 (AM C.58529, 26 d); Four probable paratypes; coll. 1940 (AM C.106867). Holotype of Siphonaria currumbinensis Hubendick, 1955 from Currumbin [28°07.83′S, 153°29.98’E, Qld, Australia]; coll. R. Endean & R. Kenny, 20-6-1952 (MV F15562, Fig. 19I). Probable paratypes, same data as holotype (MV F13953, 2 d, 3 p; MV F13948, 2 d, 3 p). Other, non-type material. Australia; Qld: Wreck Pt Yeppoon, 23°08.736’S, 150°45.865’E, Q08-4 (AM C.585471 3p, C.584743 p [SK030 protoconch B7 jaw], C.585133 p [M081], C.585134 p [SK073]); Double Head Yeppoon, 23°09.908’S, 150°47.638’E, Q08-3 (AM C.585561 3p); Bagara Hervey Bay, 24°49.180’S, 152°28.011’E, Q06-1 (AM C.585961 p); Alexandra Mooloolaba, 26°40.342’S, 153°06.822’E, Q04-1 (AM C.585383 10p); Nth Reef Scarborough, 27°11.432’S, 153°06.755’E, Q03-5 (AM C.585560 4p); Drury Pt Scarborough, 27°11.836’S, 153°06.955’E, Q03-7 (AM C.585408 10+p); Scarborough Pt Scarborough, 27°12.168’S, 153°06.980’E, Q03-8 (AM C.585638 5p); Currumbin Point, 28°07.523’S, 153°29.203’E, Q02-2 (AM C.585536 20+p, C.585122 p [M073], C.585123 p [M074], C.585124 p [M183], C.585125 p [M184]). NSW: Brunswick Heads, 28°32.297’S, 153°33.444’E, NSW12- 1 (AM C.585480 16p, C.585066 p [M077], C.585067 p [M079]); Evans River at South Evans Head, 29°6.82’S, 153°25.93’E (AM C.339263 p); Wooli Wooli River at Wooli, 29°53.27’S, 153°15.98’E (AM C.339285 4p; AM C.339287 2p); Redbank (Corindi) River at Red Rock, 29°59.03’S, 153°13.55’E (AM C.339288 3p); Urunga mangroves on S side of town just beyond footbridge, 30°29.92’S, 153°1.16’E (AM C.355024 8p); Camden Haven Inlet, 31°38.37’S, 152°49.63’E (AM C.339289 p); Fingal Bay nr Port Stephens, 32°44.087’S, 152°10.402’E, NSW09-2(AMC. 5856837 p), 32°44.990’S, 152°10.481’E, NSW09-1 (AM C.585065 p [M151]); Bolton Point Lake Macquarie, 33°0.389’S, 151°36.889’E, NSW08-3 (AM C.585737 2p, C.585058 p [M452]; C.585059 p [M453]); Bateau Bay Beach, 33°23.003’S, 151°29.197’E, NSW08- 6 (AM C.595940 p [SK520); Crackneck, Bateau Bay, 33°23.316’S, 151°29.139’E, NSW08-5 (AM C.585063 p [SK295], C.585064 p [SK296]); Broken Head Terrigal, 33°26.796’S, 151°27.030’E, NSW08-1 (AM C.585528 20+p, C.585050 p [M001]); Terrigal The Skillion, 33°27.008’S, 151°27.122’E, NSW08-2 (AM C.585558 3p, C.585053 p [M226], C.585054 p [M225]); Brisbane, WAter, 33°29.82’S, 151°18.18’E (AM C.355025 8p); Fairlight Sydney Harbour, 33°47.971’S, 151°15.990’E, NSW06-4 (AM C.585590 4p); Fairlight, North Harbour, 33°47.986’S, 151°16.837’E, NSW06-1 (AM C.585589 6p); 100m NW of Spit Bridge, Port Jackson, 33°48.210’S, 151°14.664’E (AM C.546770 2p); Spit Bridge, Port Jackson, 33°48.270’S, 151°14.520’E (AM C.546768 p); Wy-ar-gine Point Balmoral, 33°49.159’S, 151°15.195’E, NSW06-5 (AM C.585476 17p, C.585043 p [M158], C.585044 p [M159], C.585045 p [M160], C.585046 p [M161], C.595923 p [SK031]); Laings Point Sydney Harbour, 33°50.419’S, 151°16.638’E, NSW06-3 (AM C.585589 5p, C.585034 p [M193], C.585041 p [M165], C.585042 p [M167]); Bombo Kiama 34°39.232’S, 150°51.649’E, NSW03-1 (AM C.585403 10+p, C.585333 p [SK043]); Murunna Point Camel Head, 36°22.720’S, 150°04.766’E, NSW02-1 (AM C.585401 10+p). Victoria: Bastion Head Mallacoota, 37°34.429’S, 149°45.927’E, V09-1 (AM C.585787 5p, C.585294 p [M194], C.585295 p [M195]); Cape Conran, 37°48.798’S, 148°43.608’E, V08-2 (AM C.585292 p [M197]). Tas: T01-1 S of Granite Point Bridport, 40°59.739’S, 147°23.468’E (AM C.584834p [M173]). Taxonomic remarks. The largest syntype with the clearest external sculpture (Fig. 19G) is herein designated as the lectotype of S. denticulata for the stabilisation of the name and to ensure the unambiguous identity of this taxon (Art. 74.1 of the Code). Our delineation of this species is based on comparative analyses of the morpho-anatomy and mitochondrial genetics of freshly collected topotypes of S. denticulata (AM C.585303 [SK043], C.585044 [M159], C.585053 [M226]), Fig. 19H), Ellsiphon marza (AM C.584743 [SK030], C.585133 [M081], Fig. 19L) and S. currumbinensis (AM C.585123 [M074], Fig. 19J) and geographic series of additional specimens (Table S1). Quoy & Gaimard’s (1833: 340) statement ‘probably also at King George Harbour’ is not considered as part of the type locality. While there is also a King George Harbour in Tas, the King George Harbour referred to by Quoy & Gaimard (1833) is probably that near Albany, WA. However, we have not found this species in WA. The type locality for E. marza is Keppel Bay and Caloundra as stated in the original description, yet some paratypes are from additional sites (Tampian Heads and Yeppoon). Noosa, Qld is mentioned by Hubendick as a site for ‘ S. currumbinensis ’ The locality on the holotype label (MV F15562) is misspelled as Corrumbin (sic Currumbin) and the name is misspelled as S. corrumbinensis. Hutton (1873: 55) stated that S. denticulata and S. diemenensis occur in NZ; however, this is likely based on misidentifications of S. australis and/or S. propria. External morphology (Fig. 19R). Foot sole smooth, evenly pale yellow grey; foot wall pale yellow grey with evenly spread white subepithelial pustules becoming more vivid and dense around pneumostomal lobe; foot wall shows black blotches; fringing mantle translucent, covers shell mantle, outer edge lobed, thickened, with black pigmented blotches aligning with rib interstices; pneumostomallobewithinmantlebetweentherightADMs, closes the pneumostomal and anal openings at the mantle edge; two small black epithelial eye spots centralised on two thick brownish yellow centrally touching cephalic folds; genital pore inconspicuous, located on foot wall to right anterior of right cephalic fold. Shell (Figs 19G–L; Table S9). Medium to large sized (max sl mean = 24.46 mm, SD = 2.04 mm, n = 20), ovate; height medium; apex offset sightly posterior and central (usually eroded), apical sides strongly convex, protoconch direction undetermined, shell whorl dextral; growth striae prominent in bands, shell thickness thick; rib count (mean = 42.6, SD = 4.5, n = 20), primary ribs pale white, fairly straight, increasingly raised and protrude beyond shell lip to unevenly scallop and corrugate the edge; 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 dark brown to tan, white rays align on shell margin under primary/secondary ribs, siphonal groove distinct, same colour as shell edge, points to right anterior; spatula dark chocolate brown to mottled tan even whitish; ADM scar distinct, CMS straight, paler than shell lip; thickening of shell lip translucent, infills and reduces lip scalloping, spatula becomes whitened. The shells of S. denticulata, S. scabra, S. pravitas sp. nov. and S. diemenensis may appear similar and may be misidentified; external ribbing is similar. Differences exist in the sharpness and intensity of primary ribs and rib interstice colouration. In addition, the spatula colouration differs; S. denticulata and