Siphonaria tasmanica Tenison Woods, 1877

Jenkins, Bruce & Köhler, Frank, 2024, Hidden in plain sight: Systematic review of Indo-West Pacific Siphonariidae uncovers extensive cryptic diversity based on comparative morphology and mitochondrial phylogenetics (Mollusca, Gastropoda), Megataxa 13 (1), pp. 1-217 : 111-114

Siphonaria tasmanica Tenison Woods, 1877

( Figs 41H–I, 42A–G, M–N, P, R–S)

Siphonaria denticulata var. tasmanica Tenison Woods 1877: 54 (type locality: Tasmania).— Verco 1907: 105; Hardy 1915: 62.

Siphonaria zonata Tenison Woods 1878b: 99 (type locality: Tasmania).— Pritchard & Gatliff 1903: Verco 1907: 105; Hardy 1915; 23; Hedley 1915: 752; May 1921: 89; May 1923: 87; Galindo 1977: 416; Grove et al. 2006: 61; White & Dayrat 2012: 69.

Siphonaria tasmanica — Iredale 1924: 276; Cotton & Godfrey 1932: 154; Macpherson & Chapple 1951: 142; Macpherson & Gabriel 1962: 262, fig. 300; Galindo 1977: 416; Jenkins 1981: 2; 1983: 29; Quinn 1983: 81; Jenkins 1984: 3; Phillips et al. 1984: 79; Grove et al. 2006: 61; Chim & Tan 2009: 269; Grove 2011: 62, pl. 29, fig. 15; White & Dayrat 2012: 68; Colgan & da Costa 2013: 74.

Talisiphon tasmanicus nereis Iredale 1940: 442 (type locality: Port Fairy, Vic, [ Australia]).— White & Dayrat 2012: 66.

Talisiphon tasmanicus turritus Iredale 1940: 442 (type locality: Macquarie Harbour [Tas, Australia]).

Siphonaria (Pachysiphonaria) tasmanica — Hubendick 1945: 66; 1946: 22, pl. 1, fig. 12–14.

Pachysiphonaria tasmanica — Trew 1983: 2.

Liriola (Pachysiphonaria) tasmanica — Ludbrook & Gowlett-Holmes 1989: 612, fig. 11.

Siphonaria nereis — Grove et al. 2006: 61.

Siphonaria turritus — Grove et al. 2006: 61.

Talisiphon tasmanica turritus — White & Dayrat 2012: 69.

Materialexamined. Typematerial. Neotypeof Siphonaria tasmanica Tenison Woods, 1877 , present designation, from Tasman Arch, Tasmania (Art. 76.1 of the Code); coll. B.W. Jenkins, T03-3, 27 March 2018 ( AM C.585259, Fig. 42A [M114], condition Art. 75.3.7 of the code) .

Seven syntypes of Siphonaria zonata Tenison Woods, 1877 from S Tasmanian coast; coll. J.E. Tenison Woods, 1877 (MV F686, Figs 42B–C, M–N; AM C.103723, 2d).

More than twenty syntypes of Talisiphon tasmanicus nereis Iredale, 1940 from Port Fairy , Vic, [ Australia] coll. R. Bell, 1918–1919 ( AM C.108499, Figs 42D, E) .

Holotype (measurements in text) of Talisiphon tasmanicus turritus Iredale, 1940 from Macquarie Harbour, Tas; coll. A.F. Basset Hull, 1922 ( AM C.595951, Fig. 42F). Thirteen paratypes, same data as holotype ( AM C.53828; labelled ‘syntypes’).

Other, non-type material. Australia, Vic: Cape Schanck , 38°29.951’S, 144°53.369’E, V06-4 ( AM C.585458 13p); Point Lonsdale (nr Queenscliff), 38°17.276’S, 144°36.977’E, V05-1 ( AM C.585730 p [M120]) . Tas: Lagoon River: mouth, 41°29.4’S, 144°49.2’E ( TMAG E.41995 2d); Bicheno, Redbill Beach, 41°51.6’S, 148°17.4’E ( TMAG E.42002 p, d); Maria Island, Howells Point & Painted Cliffs, 42°35.796’S, 148°2.886’E ( TMAG E.42001 p, d); Maria Island , Trigonia Corner, 42°41.196’S, 148°4.404’E ( TMAG E.41993 d, p); Marion Bay , northern beaches 42°45.048’S, 147°53.592’E ( TMAG E.42004 d, p); Park Beach Dodges Ferry , 42°51.716’S, 147°36.665’E, T03- 4 ( AM C.585429 10+p, C.585266 p [SK549 protoconch D6]); Carlton Beach, Spectacle Island , 42°52.044’S, 147°36.024’E ( TMAG E.41994 d, p); Taroona, Dixons Beach, 42°56.358’S, 147°21.414’E ( TMAG E.41999 d, p); Lagoon Bch (near Saltwater River ), 42°56.903’S, 147°39.962’E, T03-2 ( AM C.585659 4p, C.585876 d); Taroona Beach; 42°57’S, 147°21’E ( TMAG E03651 3p); 42°57.18’S, 147°21’E ( TMAG E.42006 4d, 4p): Kingston Beach , 42°58.8’S, 147°19.2’E ( TMAG E.02013 p); Eaglehawk Neck , eastern side, 43°0.444’S, 147°56.082’E ( TMAG E.42003 d, p), ( TMAG E.01659 2p); Blackmans Bay 43°0.6’S, 147°19.8’E ( TMAG E.41998, 3 d / 3 p), ( TMAG E.15880 10p); Calverts Beach & Goats Bluff, 43°1.65’S, 147°29.07’E ( TMAG E.42005 3d, 3p); South Arm–Hope Beach , southern end, 43°1.8’S, 147°27.6’E ( TMAG E.01245 3p); Tasman Arch , 43°02.033’S, 147°56.963’E, T03-3 ( AM C.585728 9p, C.585255 p [SK020], C.585259 p [M114]); Tinderbox Beach , 43°3.6’S, 147°19.8’E ( TMAG E.05224 p); North Bruny Island , Dennes Point , 43°3.87’S, 147°21.066’E ( TMAG E.42000 4p, 4d); Nubeena , Parsons Bay , 43°6’S, 147°6’E ( TMAG E.05996 2p); Nubeena , White Beach , 43°7.2’S, 147°43.8’E ( TMAG E.06015 p); Fortescue Bay , 43°8.4’S, 147°57.6’E ( TMAG E.05580 6p); Port Arthur , 43°9’S, 147°52.2’E ( TMAG E.05550 4p); Three Hut Point d’Entrecasteaux Channel , 43°16.195’S, 147°14.414’E, T04-3 ( AM C.595913 2p); South Bruny Island : Simpsons Bay , 43°17.4’S, 147°18.6’E ( TMAG E.04867 4p), Cloudy Beaches –eastern beach, 43°26.352’S, 147°14.202’E ( TMAG E.41996 20d, 20p), ( TMAG E.25061 d); Peaches Point , 43°34.122’S, 146°55.037’E, T05-3 ( AM C.585606 4p); Flensing Rock , 43°34.291’S, 146°54.856’E, T05- 2 ( AM C.585460 13p, C.585512 p [SK080], C.585877 p [M119]); Cockle Creek Bay , 43°34.8’S, 146°53.4’E ( TMAG E.32706 d); Pancake Bay , 43°34.673’S, 146°55.293’E, T05-5 ( AM C.585715 8p); Trial Harbour , 41°55.758’S, 145°10.434’E ( TMAG E.41997 5d, 5p); Lucas Point , Pilot Bay , Macquarie Harbour , 42°12.241’S, 145°12.005’E,T06-1 ( AM C.585483 18p;C.585538 20+p, C.585878 p [M171], C.585879 p [M172]) . SA: Fishery Bay Cape Wiles, 34°55.107’S, 135°41.086’E, SA05- 1 ( AM C.585689 7p); Haleys Beach Gibson Peninsula , 32°45.084’S, 134°05.490’E, SA03-4 ( AM C.585467 20+p); Wandrilla Beach, nr Cape Nuyts, 32°01.894’S, 132°16.052’E, SA01-1 ( AM C.585705 10+p, C.585208 p [SK019]) .

Taxonomic remarks. The type locality was not explicitly stated in the original designation of S. tasmanica but is evident from the title of the work (“on Tasmanian Patellidae ”). Originally, Tenison Woods (1877: 45) described S. tasmanica as a variety of S. denticulata . Subsequently, Tenison Woods (1878b: 99) described the same taxon again as S. zonata without mentioning the earlier introduced name S. diemenensis var. tasmanica . No original types of S. tasmanica are known to exist and we could not locate any in the collection of the AM. The neotype of S. tasmanica (Fig. 42A) is designated herein to clarify the taxonomic status of this taxon (Art. 75.3.1 of the Code). Our delineation of this species is based on comparative analyses of the morpho-anatomy and mitochondrial genetics of freshly collected topotypes and geographic series of additional specimens (Table S1). These analyses confirm the synonymy of S. zonata (Fig. 42B–C, M–N) and establish Talisiphon tasmanicus nereis and T. tasmanicus turritus (Fig. 42D–F) as new synonyms. The name ‘ Siphonaria zonata (‘Schub. et Wagn’)’ in Deshayes (1843: 31, pl. 62, fig. 17–18) is misapplication for Patella zonata Schubert & Wagner, 1829 . The figured specimens in Schubert & Wagner (1829) and Deshayes (1843) differ from one another, neither showing a species of Siphonaria . Christiaens (1975: 91) listed Patella zonata as a synonym of Scurria scurria (Lesson, 1830) , Lottiidae . Iredale (1924: 276) was the first to recognize the synonymy of S. zonata and S. tasmanica transferring the species to Talisiphon . Tate & May (1901: 419) incorrectly considered S. zonata as a synonym of S. funiculata .

External morphology (Fig. 42S). Foot sole and foot wall evenly dark grey, paler at foot/wall edge; foot wall and mantle blue-green-grey; fringing mantle narrow, unlobed, translucent, covers exposed inner shell lip; genital pore inconspicuous, 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 small, thin, part of the mantle, between the right anterior and right posterior ADMs, closes the pneumostome at the mantle edge.

Shell (Figs 42A–G, M–N, P, R; Table S9). small to medium sized (max sl mean = 13.8 mm, SD = 2.4 mm, n = 12), height tall; apex offset weakly to posterior and left, often eroded and appearing as a white spot, apical sides convex; protoconch direction homostrophic (n = 2; Fig. 42R), shell whorl dextral; growth striae indistinct; rib count (mean = 48.3, SD = 9.6, n = 11), primary ribs fairly straight, unraised, flattened, broad, few secondary ribs; rib interstices distinct dark brown narrow lines extending to shell lip. Exterior shell colouration very distinct and unlike any other siphonariid; 3 prominent colour bands align with shell growth dividing shell height unevenly into thirds; top band upper half of shell dark brown, mid band widest and pale blue, bottom band greenish blue, limited to shell margin (Figs 42A, G). Internal colouration banded and variable; spatula blueish or cream/white, shell lip markings white aligned under primary ribs, reddish brown aligned under rib interstices; siphonal groove same colour as spatula, ADM scar dark brown, shell margins immediately above and below are paler; CMS straight. Shell thickening not observed. Shell height may be variable; e.g. lower in S. t. turritus (Fig. 42G) and taller in S. blainvillei (Fig. 42F) forms.

The neotype (Fig. 42A). Shell medium sized (sl = 17.5, sw = 14.3, sh = 8 mm), circular ovate, tall; medium thickness, apex offset strongly to posterior and weakly to left, ~52 mainly primary ribs, few secondary ribs, interior dark brown, spatula and shallow siphonal groove white to bluish; taller and slightly darker interior shell form of S. tasmanica . Neotype specimen grouped within unit 76 ( S. tasmanica ).

Reproductive system ( Figs 41H; n = 2). Positioned against inside of foot wall and over foot sole on the right posterior quarter within coelom, under the respiratory cavity. GA, EG and ED positioned in coelom between BM and RAM. GA very large, smoothly bulbous, with singular GP; AO absent; ED very short, broad, curved, unfolded, joins to side of GA; GA and ED white muscular fibrous tissue; EG soft whitish, slightly folded, smaller than GA; flagellum absent; BD and CD relatively short (BD longer), slightly curved, smooth, featureless, connect to GA together, pass together through RAM (BD over CD), CD connecting into MG/AG; BC small, elongated, bulbous, thin test, embedded in lower folds of MG/AG; HD short, narrow, coiled, links smallish AG to a large elongated yellowish granulated HD; MG/AG complex small, soft white tissue folds, enveloping SV close to embedded BC, AG larger than HG, HG side reflects curvature of inner foot wall. AL= 13.52 mm.

Spermatophore ( Fig. 41I). Length very short compared to other congenors; broad head with short flagellum (length = 1.03 mm, n = 1); head section cylindrical, bulbous, rounded tip (head length = 0.64 mm, ~62% of SPM length; head width = 159 μm; flagellum width = 34 μm), contains whitish core, test thin, translucent encasing a white opaque coiled core; flagellum transparent, tapering to a thread-like end; both sections uneven, featureless; 1 SPM in brown gelatinous mass of one BC [SK080].

Radula. ( Fig. 83M–P) Mean dentition formula 43:1:43 (SD = 4.8) with 115 (SD = 26.6), mean transverse rows 115 (SD = 26.6, n = 5); single central rachidian tooth flanked squarely by 43 half row laterals, 0–4 are inner ( Figs 83M–N), 14–20 mid and 15 outer laterals ( Fig. 83O); central tooth relatively long (half basal length) with narrow unicuspid mesocone; inner laterals (without endo or ectocones) may occur, mesocones of inner and mid laterals single pointed, mid laterals with broad pointed ectocone protruding at an acute angle halfway along the tooth’s length; outer laterals typically with a weakly bicuspidate ‘chisel’ shaped mesocone flanked by small, pointed single ecto and endocones, angle of separation of each cone from the mesocone varies ( Fig. 83O).

Comparative remarks. Siphonaria tasmanica ( lateralis group, unit 76) is most closely related to S. lessonii from the Southern Ocean (not revised herein, see Güller et al. (2015:81), S. funiculata and S. obliquata ( Figs 1, 4). It differs from S. lessonii by COI distances of ≥ 9.4% and from S. funiculata of ≥ 8.5% (Table S8). Throughout its range, S. tasmanica has been found in sympatry with five congeners in southeastern Australia. For comparisons with S. diemenensis , S. funiculata , and S. zelandica refer to comparative remarks under these species. Siphonaria jeanae has a lower, grey-blue shell with more prominent

FIGURE 42. Shells of S. tasmanica and S. incerta . A–G, M–N, P–R, S. S. tasmanica , A. Neotype AM C.585259 [M114]. B–C, M–N. Syntypes of S. zonata, MV F. 686. D–E. Largest syntypes of S. nereis AM C. 108499. F. Largest syntype of S. turrita AM C. 53828. G. Vic, Pt Lonsdale, AM C.585730 [M120]. P. Tas, Macquarie Harbour, in situ. R. Protoconch, AM C.585208 [SK019]. S. Tas, animal. H–L, O, Q, T– U. S. incerta , H. Neotype MNHN IM-2000-35954 [M263]. I. Réunion, Cap de la Houssaye, AM C.585204 [M260]. J. AM C.585205 [M261]. K. CI, Flying Fish Cove, AM C.584841 [M321]. L. CI, AM C.584843 [M309]. O. CI, TS, AM C.584840 [M320], Q. Réunion, in situ. T. Réunion, animal. U. Protoconch, AM C.585906 [M259]. Scale bars = 10 mm.

unraised brown ribs, a more scalloped edge, a wider ED, larger BC, and more thread-like SPM. Siphonaria stowae has smaller, lower, paler shell with an apex strongly offset to posterior, more prominent ribbing, a larger AO and BC, a smaller ED and a more thread-like SPM. Overall, the combination of shell geometry, size and colouration render S. tasmanica a rather distinctive species.

Tasmanian records of ‘ S. tristensis Sowerby I, 1823 ’ in Tate & May (1901) are incorrect and based on misidentified specimens of S. tasmanica . A record of ‘ S. tasmanica ’ from Percy Island, Qld ( Singleton, 1937: 396) is likely a misidentification of S. normalis . A specimen figured as S. tasmanica in Davey (1998: 118) is a specimen of S. funiculata .

Distribution and habitat. Endemic to cool temperate coasts of southern Australia, between Mallacoota, Vic, and Gibson Peninsula, SA, Tas ( Fig. 37). Found in sheltered positions (e.g., rock hollows, crevices, cracks) on very exposed rocky shores, mid littoral level (Fig. 42P); frequently associated with black mussels; home scars prominent.