Siphonaria denticulata Quoy & Gaimard 1833
Creators: name: Jenkins, Bruce
name: Köhler, Frank
Publisher: name: Zenodo
Year: 2024
Subjects: Biodiversity
Taxonomy
Animalia
Mollusca
Gastropoda
Siphonariida
Siphonariidae
Siphonaria
Siphonaria denticulata

description: 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 S. scabra dark chocolate to tan-brown, S. diemenensis and S. pravitas sp. nov. are more golden brown.
Reproductive system (Figs 20D, F, G; n = 3). Positioned within coelom under the respiratory cavity, over foot muscle, against right side of foot wall; epiphallic parts positioned between BM and RAM; single GP inconspicuous, opening from GA through foot wall posterior to right cephalic fold. Join of AO, GA and ED distinct, all whitish fibrous muscular tissue; AO elongated, bluntly pointed, folded, larger than GA, thicker than ED; ED relatively long, thick and twisted; EG broad, lobed, bluntly pointed with two curved, broad, flagellum (F1 and F2; F2 twice length of F1); BD and CD connect close together into GA between ED, AO and GP; BD noticeably longer and thinner than CD with a prominent twisted loop immediately before BC (no distal loop or MA), both ducts smooth, featureless, pass together through RAM connecting into soft curved folds of MG, BD dorsal to CD; BC large, bulbous, thin test, internally embedded into folds of MG; HD short thick, coiled, links AG to elongated narrow yellowish granulated HD; HG larger than AG, MG and AG folded, soft white tissue; SV embedded on left side of AG under BC; AG and HG sides match curvature of inner foot wall at right posterior quarter of coelom.
Spermatophore (Figs 20E, H). Head section over half of total length (head length = 8.08 ± 0.42 mm; ~ 43% of SPM length; head width = 119 ± 3 μm; flagellum width = 33 ± 1 μm, n = 3), cylindrically elongate, tip bluntly rounded, containing a white gelatinous mass; tapers quickly into consistently filamentous transparent flagellum; both sections smooth, translucent, test thin, featureless. multiple tightly coiled SPM in bursa, embedded in brown gelatinous mass. Seven and two SPM found in BC of two specimens [SK043, SK030].
Radula and jaw (Figs 83E–H). Mean dentition formula 31:1:31 (SD = 8.1), mean transverse rows 138 (SD = 20.1, n = 3, AM C.201775, C.201765, C.316303); single central rachidian tooth flanked squarely by 31 half row laterals, 0–4 are inner (SD = 4.98), 12–14 mid (SD = 4.98) and 15–19 outer (SD = 3.58) laterals (n = 5); central tooth with narrow unicuspid mesocone less than half length of basal plate, lower than laterals; inner laterals absent or few (Figs 83E–F), mid lateral mesocones generally unicuspid, however, maybe irregularly bicuspidate (Figs 83E–F), mid laterals with pointed ectocone; ectocone strong, protrudes at acute angle halfway along the tooth’s length; outer laterals typically with a ‘chisel’ shaped mesocone flanked by small, pointed single ecto and endocones, endocone positioned higher on side of tooth than ectocone, angle of separation of each cone from the mesocone varies (Fig. 83G). Jaw located inside front of buccal cavity, orange-brown, arch shaped with ‘shingle’- arranged cone-like rods, ~ 140 rods wide (~ 1.9 mm) by ~ 16 rods deep (mean length = 50 μm, SD = 6, mean width = 12 μm, SD = 3, n = 15); tip bluntly rounded [SK030].
Comparative remarks. Siphonaria denticulata (atra group, unit 33) is the sister taxon of a clade containing two species, S. opposita (unit 34) and S. plana (unit 35) (Figs 1–4). It differs from these species by COI distances of ≥ 14.8% (unit 35) and ≥ 15.8% (unit 34) (Table S3).
We found eight congeners with sympatric occurrences in SE Australia. For comparisons with S. atra and S. diemenensis refer to comparative remarks under these species. Siphonaria stowae has a smaller, more elongate, paler shell with a strongly offset apex, less prominent ribbing and edge scalloping, a smaller AO, shorter ED larger BC, and shorter SPM. Siphonaria emergens has a smaller, elongate, paler brown shell with a strongly offset apex, less prominent ribbing and edge scalloping. Siphonaria funiculata has a paler, taller shell with less raised ribs broadening to shell edge, a fainter scalloped edge, a larger AO, shorter BD and ED, absent to smaller F1, and larger, drop-like SPM. Siphonaria pravitas sp. nov. has a lower shell with greater raised ribbing and edge scalloping, no BD distal loop, a larger BC, wider F1, and shorter SPM. Siphonaria scabra has a paler shell with finer ribbing and coarser exterior, a longer AO, BC and F1, and a bursal loop. Siphonaria zelandica has a paler, lower shell with less prominent and finer narrower ribbing, a shorter ED and BD, and a short. drop-like SPM.
Menke (1851: 38) referred to the name S. denticulata for a specimen from Mexico. This reference was subsequently considered as a misidentification of S. lecanium var. palmata Carpenter, 1857 (Carpenter 1857: 183). Hubendick (1946: 40) treated this taxon as a junior synonym and subspecies of S. maura Sowerby I, 1835.
Erroneous records of ‘ S. denticulata ’ from NZ (Hutton, 1873: 55) and Mozambique (Braga, 1956: 7, pl. 1, fig. 2) are herein attributed to S. propria Jenkins, 1983 and possibly S. ferruginea Reeve, 1856, respectively. We have not located any samples of this species from NZ nor Mozambique in any of the examined museum collections. The treatment of S. denticulata as a synonym or variety of S. diemenensis by several authors is refuted herein and both species are accepted as distinct species. Hedley (1915; 1918; M96) and Angas (1867: 232) incorrectly recorded S. denticulata from Port Jackson (= Sydney) and NSW.
A specimen figured as ‘ S. marza’ in Hubendick (1946: 61, pl. 5, figs 27–28) from Port Jackson [Sydney Harbour]) is herein identified as S. denticulata. McAlpine (1952: 40, 42, fig. 1) misidentified specimens of S. scabra as ‘E llsiphen ’ (sic Ellsiphon) denticulatus. The RS figured herein (Figs 20D, G) matches closely the figure in McAlpine (1952: 43, fig. 1; as Ellsiphon denticulatus) and that shown in (Fig. 20F, from Currumbin) closely matches the distal RS figure of ‘ S. currumbinensis ’ from in Hubendick (1955: fig. 4). A specimen from NZ labelled as ‘ S. denticulata ’ in Davey (1998: 117) is herein identified as S. funiculata. Specimens stated to resemble either S. diemenensis, S. denticulata or S. australis by Dayrat et al. (2014) as ‘unit 7’ are herein identified as S. diemenensis. By contrast, the figured specimens of ‘ atra group, unit 33’ in Dayrat et al. (2014: fig. 5J, K) are morphologically consistent with S. denticulata.
Distribution and habitat. Endemic to temperate eastern and southeastern Australia, from Keppel Bay, Qld, south to Cape Conran, Vic, and Bridport, Tas (Fig. 16). In this study found in sheltered to moderately exposed sites on hard substrates (e.g., rocky shores, wooden pilings); upper and mid littoral levels; home scars prominent.
description: Published as part of 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), pp. 1-217 in Megataxa 13 (1) on pages 55-58, DOI: 10.11646/megataxa.13.1.1, http://zenodo.org/record/14984186