Siphonaria zelandica Quoy & Gaimard, 1833

Siphonaria zelandica Quoy & Gaimard, 1833 View in CoL

( Figs 26F–O View FIGURE 26 , 27C–E View FIGURE 27 )

Siphonaria zelandica Quoy & Gaimard 1833: 344 View in CoL , pl. 25; figs 17–18 (type locality: Nouvelle-Zélande [ New Zealand]).— Lamarck 1839: 205; Catlow & Reeve 1845: 100; H. Adams & A. Adams 1855 (in 1853–58): 271; 1863: 271; Iredale 1915: 478; Hanley 1858b: 153; Jenkins 1983: 4, pls 1a–h, 2a–h, 6a–c. fig. 3a–d; Paul 1984: 28; Wells 1984: 53; Phillips et al. 1984: 78, fig. (unnumbered); Villiers & Hodgson 1987: 135; Short & Potter 1987: 122, pl. 60, fig. 6; Ludbrook & Gowlett-Holmes 1989: 612, fig. 11.29 j, k; Jansen 1995: 90, fig. 376; Wilson 2002: 172, text-fig.; Wells et al. 2003: 289; White & Dayrat 2012: 69; Colgan & da Costa 2013: 74; Dayrat et al. 2014: 268, fig. 5 B, C.

Siphonaria bifurcata View in CoL — Hanley 1858b: 151; Angas 1867: 232; Tenison Woods 1878b: 100; Paetel 1873: 173; 1883: 117; 1889: 428; Whitelegge 1889: 117; Hedley 1917b: M95, 1918: M95; Iredale 1924: 276; Hubendick 1955: 4; Berry 1977: 209, fig. 19; Short & Potter 1987: 122 (not S. bifurcata Reeve, 1856 View in CoL ).

Siphonaria australis View in CoL — Lamarck 1836: 562; Hanley 1858b: 151; Hochlowski & Faulkner 1984: 3838 (not S. australis Quoy & Gaimard, 1833 View in CoL ).

Siphonaria baconi Reeve 1856 View in CoL : pl. 6, species 30 (type locality: Swan River, New Holland [Perth, WA, Australia]).— Hanley 1858b: 151; Paetel 1883: 178; 1889: 428; Angas 1865: 189; Tenison Woods 1878b: 100; Pritchard & Gatliff 1903: 221; Verco 1907: 104; 1912 b: 204; Iredale 1924: 276; Cotton & Godfrey 1932: 154; Macpherson & Chapple 1951: 142; Hubendick 1955: 129; Macpherson & Gabriel 1962: 264, fig. 302; Hodgkin et al. 1966: 27, pl. 8, fig. 4; Galindo 1977: 416; Mapstone 1978: 85; Jenkins 1981: 12, pl. 1 b, c; Roberts & Wells 1981: 72, pl. 30; Trew 1983: 4; Jenkins 1983: 5; Paul 1984: 28; Short & Potter 1987: 122; White & Dayrat 2012: 61.

Siphonaria zealandiae Paetel 1889: 430 (invalid; incorrect subsequent spelling of zelandica View in CoL ).

Siphonaria zebra Reeve 1856 View in CoL : pl. 5, species 22 [not species 21] (type locality: Port Jackson [Sydney Harbour], Australia).— Hanley 1858b: 153; Paetel 1889: 430; Hedley 1918: M95; Iredale,1924: 276; Hubendick 1955: 4; Galindo 1977: 416; Jenkins 1983: 5, 12, pl. 1 e–h; Trew 1983: 7; White & Dayrat 2012: 69.

Siphonaria (Ductosiphonaria) baconi View in CoL — Hubendick 1945: 70; 1946: 37, pl. 6, fig. 10, 11.

Siphonaria luzonica View in CoL — Adcock 1893: 11; Shikama 1964: 6, pl. 2, fig. 7 (not S. luzonica Reeve, 1856 View in CoL ).

Planesiphon elegans Iredale 1940: 441 View in CoL , figs 3–4 (type locality: Keppel Bay, Queensland).— Marcus & Marcus 1960: 122; Jenkins 1983: 5, 12, pl. 1 d, pl. 2 g –h; Short & Potter 1987: 122; White & Dayrat 2012: 63.

Siphonaria (Ductosiphonaria) bifurcata View in CoL — Hubendick 1945: 22, 70; 1946: 37, pl. 2, figs 9–13 (not S. bifurcata Reeve, 1856 View in CoL ).

Siphonaria (Siphonaria) elegans View in CoL — Hubendick 1945: 29, 72, fig. 44, 46, 47.

Siphonaria (Siphonaria) australis View in CoL — Hubendick 1945: 72; 1946: 49, pl. 3, fig. 28–31 (not S. australis Quoy & Gaimard, 1833 View in CoL ).

Ductosiphonaria bifurcata — McAlpine 1952: 42, fig. 3 (not S. bifurcata Reeve, 1856 ).

Siphonaria elegans View in CoL — McAlpine 1952: 42.

Siphonaria (Pachysiphonaria) zealandica Hubendick 1955: 129–136 , figs 5–8 (invalid; incorrect subsequent spelling of zelandica View in CoL ).

Ductosiphonaria baconi — Cotton 1959: 411.

Ellsiphon denticulata — Cotton 1959: 411 (not S. denticulata Quoy & Gaimard, 1833 ).

Planesiphon bifurcata — Iredale & McMichael 1962: 82 (not S. bifurcata Reeve, 1856 ).

Siphonaria bifaciata Galindo 1977: 416 (invalid; incorrect subsequent spelling, not S. bifurcata Reeve, 1856 ).

Material examined. Type material. Lectotype of Siphonaria zelandica Quoy & Gaimard, 1833 , from Nouvelle Zealandie [ New Zealand (in error for Australia, see Jenkins, 1983: 5)] ( MNHN IM 2000-5055 , Fig. 26F View FIGURE 26 ). Two paralectotypes, same data as lectotype ( MNHN IM 2000-5135 ).

Lectotype of Siphonaria baconi Reeve, 1856 from Swan River [Perth, WA] ( NHMUK 1979164.1 , Fig. 26J View FIGURE 26 ). Two paralectotypes of Siphonaria baconi Reeve, 1856 . Same data as lectotype ( NHMUK 1979164.2 and 1979164.3).

Lectotype of Siphonaria zebra Reeve, 1856 from Port Jackson [on type label; ‘ Philippine Islands’ in Conch. Icon. incorrect—see Taxonomic remarks below] [Sydney, Australia], ( NHMUK 1979168 /1, Fig. 26H View FIGURE 26 ). Three paralectotypes of Siphonaria zebra Reeve, 1856 . Same data as holotype ( NHMUK 1979168/2-4 ).

Syntype of Planesiphon elegans Iredale, 1940 from Keppel Bay , Qld; coll. H. Bernhard, Sept 1935 and 1936 ( AM C.103710 , Fig. 26L View FIGURE 26 ) .

Syntypes of P. elegans Iredale 1940 . Expanded syntype data ( AM C.108528 , 6d, locations on label of North Keppel Island and Emu Park ; AM C.126707 , 5d, location on label of Emu Park ; MV F13845 , 1d, location on label of Emu Park ) .

Other, non-type material. Australia, Qld: Keppel Bay, 23°25’S, 150°55’E ( AM C.103710 d); GoogleMaps Emu Park, 23°15’S, 150°49’E ( AM C.126707 5d); GoogleMaps North Keppel Is & Emu Park 2, 3°25’S, 150°55’E ( AM C.108528 6d). GoogleMaps Caloundra Shelly Beach, 26°48’S, 153°9’E ( AM C.391924 d). GoogleMaps NSW: Fingal Bay nr Port Stephens , 32°44.990’S, 152°10.481’E, NSW09-1 ( AM C.585559 3p); GoogleMaps Broken Head Terrigal, 33°26.796’S, 151°27.030’E, NSW08-1 ( AM C.585380 10p, C.585052 p [M098]); GoogleMaps Terrigal The Skillion, 33°27.008’S, 151°27.122’E, NSW08-2 ( AM C.585591 4p, C.585057 p [M222]); GoogleMaps Wy-ar-gine Point Balmoral, 33°49.159’S, 151°15.195’E, NSW06-5 ( AM C.585445 11p, C.585047 p [M155], Fig. 26I View FIGURE 26 , C.585048 p [M156], C.585049 p [M157]); GoogleMaps Bombo Kiama, 34°39.232’S, 150°51.649’E, NSW03-1 ( AM C.585557 3p); GoogleMaps Murunna Point Camel Head, 36°22.720’S, 150°04.766’E, NSW02-1 ( AM C.595960 2p) GoogleMaps . Vic: Bear Gully, 38°53.519’S, 145°59.029’E, V07-3 ( AM C.585572 3p); GoogleMaps Caves, Inverloch 38°39.777’S, 145°40’871’E, V07- 2 ( AM C.585359 p); GoogleMaps San Remo , headland, 38°31.913’S, 145°22.209’E, V07-4 ( AM C.585582 34p); GoogleMaps San Remo , 38°31.489’S, 145°21.858’E, V07-1 ( AM C.585358 p); GoogleMaps Frankston , 38°09.236’S, 145°06.457’E, V06-1 ( AM C.585651 5p); GoogleMaps West Head Flinders , 38°28.883’S, 145°01.727’E, V06-3 0 ( AM C.585717 8p); GoogleMaps Point Lonsdale (nr Queenscliff), 38°17.276’S, 144°36.977’E, V05-1 ( AM C.585608, 4 p); GoogleMaps Roadknight Point , 38°25.707’S, 144°11.102’E, V04-1 ( AM C.585474 16p); GoogleMaps Marengo Rocks Apollo Bay , 38°46.772’S, 143°39.997’E, V03-1 ( AM C.585650 5p); GoogleMaps Crofts Bay , 38°35.363’S, 142°50.633’E, V01-3 ( AM C.585607 4p, C.585284 p [M177]); GoogleMaps Armstrong Bay , 38°21.012’S, 142°21.633’E, V01-2 ( AM C.585726 9p). GoogleMaps SA: Cape Northumberland , 38°03.503’S, 140°40.378’E, SA15-1 ( AM C.585351 p); GoogleMaps Cape Northumberland Port Macdonnell , 38°03.308’S, 140°39.398’E, SA15-2 ( AM C.585723 9p, C.585223 p [M200]); GoogleMaps Cape Thomas , 37°04.461’S, 139°44.659’E, SA14-1( AM C.585766 7p,C.584893d [R3032], C.584935 p [SK038], C.585218 p [M199]); GoogleMaps Fisheries Bay Lands End , 35°37.999’S, 138°06.921’E, SA13-2 ( AM C.585713 8p, C.584900 p [SK509], C.585216 p [SK024]); GoogleMaps Groper Bay nr West Cape , 35°14.108’S, 136°49.883’E, SA10- 1 ( AM C.585601 4p); GoogleMaps Pondalowie Bay , 35°13.989’S, 136°49.892’E, SA10-2 ( AM C.585473 16p, C.585492 p); GoogleMaps Port Neill , 34°07.102’S, 136°21.271’E, SA06-1 ( AM C.585710 8p); GoogleMaps Port Moonta , 34°03.273’S, 137°33.592’E, SA09-1 ( AM C.585647 5p); GoogleMaps Salmon Point , 33°38.547’S, 134°51.916’E, SA04-2 ( AM C.585688 7p); GoogleMaps Whyalla , 33°02.539’S, 137°35.511’E, SA07-1 ( AM C.585675 6p); GoogleMaps Rocky Point , 32°12.250’S, 133°14.861’E, SA02-4 ( AM C.5856877p); GoogleMaps Ceduna , 32°07.438’S, 133°40.260’E,SA03- 2 ( AM C.585707 8p); GoogleMaps Denial Bay Ceduna , 32°05.886’S, 133°34.817’E, SA03-1 ( AM C.585762 10p); GoogleMaps Port Le Hunte Point Sinclair , 32°05.554’S, 132°59.476’E, SA02- 2 ( AM C.585706 8p); GoogleMaps Wandrilla Beach , nr Cape Nuyts 32°01.894’S, 132°16.052’E, SA01-1 ( AM C.585522, 20 p). GoogleMaps WA: Cave Point , 35°06.965’S, 117°54.080’E, WA62- 1 ( AM C.584768 5p, WAM S74186 5p); GoogleMaps Waterbay Point , 35°05.540’S, 117°56.963’E, WA62-2 ( AM C.584684 2p, WAM S74187 View Materials 2p); GoogleMaps Peaceful Bay , 35°02.989’S, 116°55.769’E, WA60-7 ( AM C.584744 4p, WAM S74183 4p); GoogleMaps 35°02.865’S, 116°55.722’E, WA60-8 ( AM C.585579 8p); GoogleMaps Wilson Head Ocean Beach, 35°02.250’S, 117°19.894’E, WA61-1 ( AM C.584730 10p, WAM S74185 5p); GoogleMaps Cape Riche 34°36.213’S, 118°45.401’E, WA62-5 ( AM C.584714 5p, WAM S74188 5p); GoogleMaps Bremer Bay Boat Harbour, 34°25.613’S, 119°23.818’E, WA63-2 ( AM C.584773 5p, WAM S74189 5p); GoogleMaps Sarge Bay Cape Leeuwin, 34°22.091’S, 115°08.820’E, WA60-4 ( AM C.585371 p); GoogleMaps Augusta, 34°20.451’S, 115°10.069’E, WA60-5 ( AM C.584699 3p, WAM S74182 3p); GoogleMaps Alexander Bay 2, 33°53.467’S, 122°44.995’E, WA64-4 ( AM C.584776 5p, 74191 5p); GoogleMaps Alexander Bay , 33°53.374’S, 122°44.922’E, WA64-3 ( AM C.584774 5p, 74190 5p); GoogleMaps Salmon Beach Esperance , 33°53.254’S, 121°50.381’E, WA64-2 ( AM C.585580 3p); GoogleMaps Blue Haven Beach Esperance , 33°53.105’S, 121°51.687’E, WA64-1 ( AM C.585314 p [M097]); GoogleMaps Cowaramup Point, 33°51.934’S, 114°58.904’E, WA60-3 ( AM C.585521 2p); GoogleMaps Yallingup, 33°38.358’S, 115°01.481’E, WA60-9 ( AM C.584696 3p, WAM S74184 2p); GoogleMaps Point Dalling Dunsborough, 33°35.955’S, 115°06.315’E, WA59-4 ( AM C.584750 5p, C.595950 p [M129], WAM S74180 View Materials 5p); GoogleMaps Point Casuarina Bunbury, 33°18.544’S, 115°38.201’E, WA59- 3 ( AM C.585578 3p); GoogleMaps groyne nr Robert Point Mandurah, 32°31.270’S, 115°42.409’E, WA59-1 ( AM C.585437 5p, WAM S74179 5p); GoogleMaps Fremantle Hbr breakwater 32°03.342’S, 115°43.987’E, WA58-5 ( AM C.585612 4p); GoogleMaps Longreach Bay point, Rottnest Is, 31°59.333’S, 115°32.063’E RI01 ( AM C.584783 4p, WAM S74164 4p); GoogleMaps Point Brown Swan River , 32°02.344’S, 115°45.471’E, WA59-5 ( AM C.584709 4p, C.584899 p [SK039], C.585935 p [M127, SK243], WAM S74181 4p); GoogleMaps Grey, 30°39.968’S, 115°08.072’E, WA58-2 ( AM C.585367 p); GoogleMaps Jurien Bay 30°17.244’S, 115°02.482’E, WA58-1 ( AM C.585576 3p); GoogleMaps Cambawarra Head Green Head, 30°04.136’S, 114°57.830’E, WA57-6 ( AM C.585518 2p); GoogleMaps Illawong, 29°42.254’S, 114°57.542’E, WA57-5 ( AM C.584708 4p, WAM S74178 View Materials 4p); GoogleMaps Illawong bch rocks, 29°42.198’S, 114°57.551’E, WA57-4 ( AM C.584705 4p, WAM S74177 View Materials 3p); GoogleMaps S end Leander Point Port Denison , 29°16.725’S, 114°54.918’E, WA57-2 ( AM C.584682 2p, WAM S74176 View Materials 2p); GoogleMaps Leander Point Port Denison , 29°16.568’S, 114°54.858’E, WA57-1 ( AM C.584692 3p, WAM S74175 View Materials 2p); GoogleMaps Cape Burney Geraldton, 28°52.084’S, 114°38.056’E, WA54-1 ( AM C.584748 3p, WAM S74174 View Materials 3p); GoogleMaps Turtle Bay East Wallabi Is , 28°25.804’S, 113°44.538’E, WA55-1 ( AM C.585658 5p); GoogleMaps Horrocks, 28°21.469’S, 114°24.751’E, WA53-1 ( AM C.584681 2p, WAM S74173 View Materials 2p); GoogleMaps Red Bluff, 27°44.627’S, 114°08.576’E, WA52-2 ( AM C.584741 10p, C.595943 p [M124], WAM S74172 View Materials 10p); GoogleMaps Chinamans Rock Kalbarri, 27°42.776’S, 114°09.361’E, WA52-1 ( AM C.584740 10p, WAM S74171 10p); GoogleMaps Pepper Point (Zuytdorp), 26°23.826’S, 113°18.268’E, WA51-1 ( AM C.584756 5p, WAM S74170 View Materials 5p); GoogleMaps Bottle Bay Cape Peron, 25°32.566’S, 113°29.467’E, WA49-1 ( AM C.585364 p); GoogleMaps Point Quobba, 24°29.124’S, 113°24.501’E, WA45-1 ( AM C.584707 4p, WAM S74169 View Materials 4p); GoogleMaps Point Maud, 23°08.322’S, 113°46.294’E, WA44-1 ( AM C.584723 5p, WAM S74168 View Materials 4p); GoogleMaps Pt S of Bruboodjoo Pt Bateman Bay, 23°02.991’S, 113°49.371’E, WA43- 1 ( AM C.584733 12p, WAM S74167 6p); GoogleMaps Tantabiddi, 21°54.739’S, 113°58.706’E, WA42-1 ( AM C.584691 3p, C.585121 p [SK188], C.585310 p [M207], C.585311 p [M208], WAM S74166 2p); GoogleMaps NW Cape Exmouth, 21°48.360’S, 114°07.665’E, WA41-1 ( AM C.585363 p); GoogleMaps Cape Keraudren, 19°57.393’S, 119°46.358’E, WA29-2 ( AM C.584706 4p, WAM S74165 View Materials 4p); GoogleMaps Cape Latouche Treville nr Gourdon Bay , 18°27.457’S, 121°48.725’E, WA27-1 ( AM C.584342 p [SK067],C.584927 p [SK066]); GoogleMaps Catamaran Bay formal, 16°27.622’S, 123°00.242’E, WA22-3 ( AM C.585304 p [M045]) GoogleMaps .

Taxonomic remarks. The original description of P. elegans contains no type designation. Hence, we consider all types as syntypes. Jenkins (1983) designated the lectotypes of S. zelandica (5, pl. 1, fig. a), S. baconi (10, pl. 1, fig. b) and S. zebra (11, pl. 1, fig. e). Anatomical and molecular analyses of freshly preserved topotypic specimens matching type specimens of each nominal species and geographical series validate the identity and occurrence of S. zelandica and confirm that S. zebra , Planesiphon elegans and S. baconi are junior synonyms. The descriptions of S. zebra and S. bifurcata were mixed up in Reeve’s monograph: The types of ‘ S. zebra’ match the description of S. bifurcata (species 22, figure 21) while the types of ‘ S. bifurcata’ match the description of S. zebra (species 21, figure 22) (see Jenkins, 1983: 28; White & Dayrat, 2012: 60). As a result, the correct type locality of ‘ S. zebra’ is Port Jackson instead of the Philippines Islands. The labelling errors in Reeve (1856) confused many subsequent authors. Iredale (1924: 276) also noted the transposition and correctly considered that S. zebra referred to the same species as S. baconi . Siphonaria zebra has been treated as a synonym of S. sipho by Reeve (1856), repeated in Schrenck (1867: 306). Hutton (1883: 143) refrained from accepting Reeve’s (1856) synonymy. The record of ‘ S. luzonica ’ in Adcock (1893, 11, SA) and listing of S. stellata , S. exigua , S. baconi and S. kurracheensis as its synonyms are incorrect. Herein we demonstrate that ‘ S. luzonica ’ (= S. sipho ) and S. zelandica are distinct species with different distributions.

Paetel (1883: 178, 1889: 430) and Galindo (1977: 416) incorrectly recorded S. zebra from the Philippines. Verco (1907: 104) incorrectly considered Trimusculus albida (Angas, 1878) as a possible white form of S. baconi . Hubendick (1946: 37) pointed out that Adcock’s (1893: 11) synonymisation of S. baconi with S. luzonica was questionable. The record of ‘ S. zebra ’ in Kuroda (1941: 137, pl. 3, figs 49–50) is a misidentification of S. rucuana . Hubendick (1946: 49) treated S. zelandica Quoy & Gaimard, 1833 as a synonym of S. australis Quoy & Gaimard, 1833 . McAlpine (1952: 42) compared the conchology and anatomy of P. elegans , S. diemenensis and S. ‘ bifurcata’ and species interpretations by Iredale and Hubendick, from which Hubendick (1955: 4–5) subsequently indicated P. elegans was a synonym of S. zelandica . Confirmed in Jenkins (1983: 4–5, 28) reviewing the type specimens and redescribing the anatomy and distributions. Hubendick (1946: 69) uncertain of the identity of S. zebra , suggested it may be identical with S. plana . Morrison (1972: 56–58) assigned S. elegans , along with 29 other nominal species ( S. zelandica not mentioned), to S. laciniosa without specific explanation other than based on similarity to a ‘ laciniosa ’ shell form and ‘common reproductive development’. These synonymies are not supported by examination of type specimens and morpho-anatomy. ‘ S. zelandica’ in Berry (1977: 204–5, pl. 1), based on Hutton (1882: 343, correctly as S. australis ) refers to S. australis . Trew (1983: 7) considered S. zebra a synonym of S. kurracheensis , probably following Hubendick (1946: 54), however, is presently indeterminable as location of the material in the collection is not indicated. Jenkins (1983: 5, 10–11, 28) and Wells (1984: 52), treated S. baconi as a synonym of S. zelandica . The taxon was redescribed by Jenkins (1983: 6, fig. 1C). Here we utilise and expanded the information provided there.

External morphology ( Fig. 26M View FIGURE 26 ). Foot sole evenly dark grey, foot wall, foot edge mantle, cephalic folds and pneumostome lobe evenly yellow; cephalic folds large; mantle wider than foot wall, thin, translucent; mantle edge unthickened, black edge bands absent; faint irregular black blotches on foot wall and centre of cephalic folds, touch and extend over the mouth region; two small black subepithelial eye spots are centrally located cephalic folds.

Shell ( Figs 26F–O View FIGURE 26 ; Table S9). small to medium sized, ovate, (max sl mean = 17.2 mm, SD = 1.6 mm, n = 12), height low; apex offset posterior and left, apical sides straight to concave; shell whorl dextral, protoconch direction homostrophic (n = 2; Fig. 26O View FIGURE 26 ); growth striae fine to inconspicuous; thickness fine; outer lip is uneven, often broken and finely scalloped with primary ribs extending past the outer lip; rib count (mean 48, SD = 5.1, n = 12), white primary ribs slightly raised, mainly broad, apically continuous, radiate from the apex with black to light brown interstices finely striated with four to six apically discontinuous secondary ribs; exteriorly the siphonal ridge is indistinct; pneumostomal margin of the shell slightly raised and the outer edge extends weakly out to the right side; interior smooth with brown to white spatula, slightly swollen margin; brown apical rays are often presented extending from the spatula to slightly swollen lip margin corresponding with exterior rib interstices; siphonal groove (fig. 1C, sg) is shallow, smooth, of a similar colouration to the spatula; CMS (fig. 1C, cms) is shallow and weakly concave to straight; ADM scars are similarly shallow, may be darker or lighter brown than the spatula. Juvenile shells are miniature adults displaying similar ribbing but with paler interiors and exteriors.

Reproductive system ( Figs 27C–D; n View FIGURE 27 = 5). HG (ovotestis), large yellow, granular and AG both located in posterior region of coelom, against inner foot wall and under the respiratory system; HG joined to anterior of soft folds of AG by a short pink lobed HD, passes and between the translucent soft white semicircular folds of the MG and the white AG; thin white CD emerges from MG. SV small ovate pinkish, connected via a short thin duct to the junction of the HD and CD. BD long thin white, passes alongside but dorsal to the much thicker CD through the adductor muscle and both enter adjacently into the GA; both CD and BD smooth, non-looped, featureless; BC small, brown, appears spherical when holding SPM but deflated when empty; ED long, thick, white and may be centrally looped ( Jenkins, 1984: fig 1b), enters the GA almost opposite the BD and CD juxtaposed points of entry, no AO present; EG, cream, soft, lobed, longer than ED; single F1, short, broad, white, branching from the junction of the EG and ED. GP small, opens from GA through the foot wall, under the mantle and posterior to the right cephalic fold.

Spermatophore ( Fig. 27E View FIGURE 27 ). Drop-like with short, hooked flagellum (length = 1.12 mm, n = 1); head section bulbous, round tip (head length = 1.0 mm, head ~86% of SPM length; head width = 150 μm; flagellum width = 34 μm), body and flagellum test opaque, thick, tapering to a pointed end; both sections featureless; 1 SPM in brown gelatinous mass in one BC.

Radula (in Jenkins, 1983: pl. 6, fig. a–c). Wide intraspecific variation exists in the number of transverse rows and the number of inner, mid and outer lateral teeth in longitudinal rows. The mean dentition formula is 28:1:28 ( SD = 3.8, n = 7) with 122 parallel and slightly curved (anteriorly convex) transverse rows ( SD = 6.5, n = 8). Of the 28 half row laterals, 10 ( SD =2.4) are inner, 2 ( SD = 0.8) mid and 16 ( SD = 1.5) outer teeth means respectively (n = 8). In each longitudinal row a gradual reduction in tooth size and an increase in distance between transverse rows occurs from the central tooth to the outer laterals. All teeth are bluntly concave posteriorly. The central tooth has a lower profile than the flanking laterals and a short pointed mesocone less than half the base length. The anterior is forked and wider than the notched posterior. The narrow base interlocks and articulates with the adjacent central teeth. Lateral teeth have broad bases. The mesocone of the inner laterals is either pointed or bicuspidate. The inner cusp is always longer and usually overlapping the posterior of the inner lateral in front. Ecto and endocones are absent. The mid laterals have an ectocone and a bluntly bicuspidate mesocone with the inner cusp being longer. Outer laterals have a square shaped base with a broad blunt mesocone flanked by short pointed ecto and endocones. The clefts between the mesocones and the side denticles of the outer laterals are widely variable in width and angle of separation. Around the third or fourth outer longitudinal row, often aberrant outer lateral teeth appear as fused teeth with double mesocones.

Comparative remarks. Siphonaria acmaeoides from Japan is the closely related sister species of S. zelandica ( atra group, unit 26) ( Figs 1 View FIGURE 1 , 2 View FIGURE 2 ). Both species differ from each other by a genetic distance of at least of 5.8% (Table S4). The next closely related species are S. restis sp. nov. (distance = 23%) and S. stowae (distance = 17.7%) (Table S4). Examination of the type specimens and topotypic specimens of S. zelandica and S. acmaeoides revealed differences in shell geometry, the spacing of ribs, secondary ribbing, colouration, and reproductive features. Both taxa occur in splendid isolation from each other and are maintained as distinct species. Throughout the range of S. zelandica , we found seventeen congeners with partly sympatric distributions. Five species are sympatric in southern Qld. For comparisons with S. atra , S. denticulata , and S. viridis refer to comparative remarks under these species. Siphonaria hienghenensis sp. nov. has a larger, darker shell with more raised ribbing, a more prominent siphonal ridge, darker interstices, and interior colouration, a larger AO and BC, BD with distal loop, and a thread-like SPM. Siphonaria opposita has a larger shell with a more prominent and flared siphonal ridge, and stronger edge scalloping, a larger AO, ED and BC, and a thread-like SPM. Siphonaria scabra has a larger, taller, darker shell with more raised ribbing, rougher exterior, stronger edge scalloping and a darker interior, larger AO and BC, longer ED, and a long thread-like SPM.

Seven congeners are sympatric in NSW (along with S. denticulata and S. scabra ). For comparison with S. diemenensis refer to comparative remarks there. Siphonaria emergens has a much smaller, orange-brown shell with less raised ribbing and an apex more posteriorly offset. Siphonaria funiculata has a shell with paler ribs and with dark brown interstices, a larger AO and BC, shorter BD, and a longer ED. Siphonaria pravitas sp. nov. has a taller shell with more raised prominent ribbing and siphonal ridge, stronger edge scalloping, a larger BC, and a thread-like SPM. Siphonaria stowae has a taller and much smaller, yellowish-cream shell, a larger BC, and a thread-like SPM.

Six congeners are sympatric in Vic and SA Australia (along with S. denticulata , S. diemenensis , S. funiculata and S. stowae ). Siphonaria jeanae has a smaller, darker, grey blue shell with brown ribs and purplish spatula, a very small to absent AO, broader and twisted ED, and a bulbous SPM. Siphonaria tasmanica has a taller, blue grey shell with a more central apex and less distinct siphonal ridge, a shorter BD, and smaller BC.

Six congeners are sympatric in WA (along with S. atra , S. jeanae and S. stowae ). Siphonaria alba has a larger shell with central apex, raised ribbing and stronger edge scalloping, a larger AO and BC, longer ED, and a longer, thread-like SPM. Siphonaria gemina sp. nov. has taller darker shell with more prominent siphonal ridge and stronger edge scalloping, a larger ED and BC, shorter BD, and a thread-like SPM. Siphonaria restis sp. nov. has a taller shell with more prominent and raised ribbing and siphonal ridge, stronger edge scalloping, a larger AO, BC and ED, long narrow F1, and a thread-like SPM.

Siphonaria zelandica displays a shell morphology like other species in the plicata group, but the structure of RS in each of these species differs (mainly size of BC; epiphallus parts of ED and F1).

Hubendick (1945: 29, 72, figs 44, 47; 1946: 56–57, pl. 4, figs 32–34) treated S. elegans as an accepted species. However, he mis-identified specimens of either S. sipho or S. bifurcate S. zelandica . The RS of ‘ S. elegans’ figured in Hubendick (1945: 30–31, figs 44, 47) corresponds to S. zelandica depicted herein ( Fig. 27C–D View FIGURE 27 ). The RS of ‘ D. bifurcata ’ figured by McAlpine (1952: 43, fig. 3) does also closely resemble that of S. zelandica .

The records of S. zelandica by Lamarck (1836: 558), Hutton (1873: 55, 1878: 41; 1880: 36; 1883: 143, pl. 17, figs H–M; 1904: 68), Suter (1904: 68; 1913: 600; 1915), Iredale (1915), Oliver (1923: 498), Odhner (1924: 55), Bucknill (1924: 84), Powell (1933: 186; 1937: 86; 1939: 217; 1946: 91), Borland (1950: 386), Knox (1955: 86), Powell (1955: 120; 1957a: 114), Dell (1960: 147, 1963: 227), Morton & Miller (1968: 83, pl. 19, fig. 8, 8a), Galindo (1977: 416), Powell (1979: 292, pl. 54, fig. 10), Trew (1983:7) and Wells & Wong (1978: 417) are all misidentifications of S. australis (refer Jenkins, 1983: 1).

Hubendick (1946) was uncertain about the identities of S baconi and S. zebra ; probably because of the mix-up of the figures in Reeve’s descriptions of both species). Hubendick (1945: 80; 1946: 37) treated ‘ S. baconi’ as accepted, but Hubendick (1946: 37, pl. 2, figs 9–13) tentatively placed it under ‘ S. bifurcata ’. The figures of ‘ S. baconi ’ in Hubendick (1946: 92, pl. 4, figs 33–34) are specimens of S. zelandica . Hubendick (1946: 54) also tentatively placed S. zebra in synonymy of ‘ S. kurracheensis ’ and listed the variant of S. kurracheensis var. zebra (pl. 2, fig. 37). This figure is attributed here to S. belcheri . Figures of ‘ S. elegans ’ in Hubendick (1946: pl. 4, fig. 33–34) from ‘Kupang, Timor and Java Sea’ are likely specimens of S. viridis . Specimens figured as ‘ S. baconi’ in Hubendick (1946: 91, pl. 6, fig. 10–12) are specimens of S. zelandica . The SPM depicted herein ( Fig. 27E View FIGURE 27 ) resembles that figured in Hubendick (1955: 10, fig.8) of ‘ S. zealandica’ apart from a shorter flagellum (possibly broken). The record of S. zelandica in Davey (1998: 117) is a specimen of S. denticulata . Figured specimens ‘unit 26’ in Dayrat et al. 2014: fig. 5B, C) correspond well with S. zelandica .

Distribution and habitat. Endemic toAustralia, from Keppel Bay, Qld, south along NSW, Victorian and SA coasts to Broome, WA ( Fig. 28 View FIGURE 28 ). Reasonably common in sheltered positions, often within runoff areas and shallow platform pools ( Fig. 26D View FIGURE 26 ), on exposed to moderately exposed rocky intertidal marine shores across upper and mid littoral levels.