Siphonaria funiculata Reeve, 1856

Siphonaria funiculata Reeve, 1856 View in CoL

( Figs 33A–H, O–P, S–U View FIGURE 33 , 34A–D View FIGURE 34 )

Siphonaria funiculata Reeve 1856 View in CoL : pl. 2, species 6 (type locality: ‘Van Diemen’s Land’ [Tasmania, Australia]).— Hanley 1858b: 152; Angas 1867: 232; Tenison Woods 1877: 58; 1878b: 99; Whitelegge 1889: 117; Paetel 1889: 428; Henn & Brazier 1894: 179; Tate & May 1901: 419; Pritchard & Gatliff 1903: 220; Hedley 1917b: M96; May 1921: 88; 1923: 87, pl. 41, fig. 3; Iredale 1924: 275; Macpherson & Gabriel 1962: 262, fig. 301; Galindo 1977: 416; Jenkins 1981: 2, pl. 1b; 1983: 29; Quinn 1983: 83; Phillips et al. 1984: 79, text-fig.; Short & Potter 1987: 122; Jansen 1995: 90, fig. 377; Davey 1998: 119, top fig.; Grove et al. 2006: 60, 2011: 62, pl. 29, fig.14; White & Dayrat 2012: 63; Colgan & da Costa 2013: 74; Dayrat et al. 2014: 266, ‘unit 8’, fig. 3 I–J; Gonzàlez-Wevar et al. 2018: 5, fig. 1.

Siphonaria blainvillei Hanley 1858b: 152 View in CoL , 153 (type locality unknown [probably Tasmania, Australia]).— Paetel 1889: 428; Hedley 1915: 752; Jenkins 1981: 2, pl. 1a; Coan & Kabat 2012: 336; Grove et al. 2006: 60; White & Dayrat 2012: 61.

Siphonaria lirata View in CoL — Hanley 1858b: 152; Hedley 1915: 751; Galindo 1977: 416; White & Dayrat 2012: 65; Dayrat et al. 2014: 267 (not S. lirata Reeve, 1856 View in CoL ).

Siphonaria luzonica View in CoL — Angas 1865: 190 (not S. luzonica Reeve, 1856 View in CoL ).

Siphonaria laeviuscula View in CoL — Hutton 1878: 42 (not S. laeviuscula Blainville, 1835 View in CoL ).

Siphonaria zonata View in CoL — Tate & May 1901: 419 (not S. zonata Tenison Woods, 1878 View in CoL ).

Siphonaria virgulata Hedley 1915: 751 View in CoL , pl. 85, figs 96–98 (type locality: Terrigal, Sydney, and Twofold Bay [NSW, Australia]).— Hedley 1917b: M95; Iredale 1924: 276; Hubendick 1943: 4; Macpherson & Chapple 1951: 142; Hubendick 1955: 1, fig 1; Galindo 1977: 416; Crease 1980: 38; Jenkins 1981: 2, pl. 2a–k; Short & Potter 1987: 122; Davey 1998: 119; Grove et al. 2006: 60; Chim & Tan 2009: 269; White & Dayrat 2012: 69.

Talisiphon virgulata — Iredale 1940: 442; Iredale & McMichael 1962: 82.

Siphonaria oblivirgulata Hubendick 1943: 2 View in CoL , figs 2, 6 (type locality: Port Jackson, Australien [ Australia]).— Grove et al. 2006: 60; White & Dayrat 2012: 66.

Siphonaria (Pachysiphonaria) funiculata View in CoL — Hubendick 1945: 12, 15, 16, 66, figs 3, 12, 58; 1946: 23, pl. 1, figs 15–17.

Siphonaria (Benhamina) oblivirgulata View in CoL — Hubendick 1946: 25, pl. 1, figs 26–29.

Talisiphon funiculata — Cotton 1959: 441.

Talisiphon oblivirgulata — Iredale & McMichael 1962: 82.

Siphonaria (Talisiphon) virgulata View in CoL — Morrison 1963: 7.

Pachysiphonaria funiculata — Trew 1983: 2.

Pachysiphonaria virgulata — Trew 1983: 2.

Siphonaria diemenensis View in CoL — Davey 1998: 117 (not S. diemenensis Quoy & Gaimard, 1833 View in CoL ).

Siphonaria tasmanica View in CoL — Davey 1998: 118 (not S. tasmanica Tenison Woods, 1877 View in CoL ).

Material examined. Type material. Lectotype of Siphonaria funiculata Reeve, 1856 , present designation, from Tasmania [ Australia] ( NHMUK MC.197927/1 , Fig. 33A View FIGURE 33 ). Three paralectotypes, same data as lectotype ( NHMUK MC.197927/2-4 ).

Holotype of Siphonaria blainvillei Hanley, 1858 ( NHMUK 1907.10.28.90 , Fig. 33D View FIGURE 33 ).

Holotype of Siphonaria virgulata Hedley, 1915 , from Terrigal [E of Gosford , NSW], Sydney; coll. C. Hedley, 1915 ( AM C.39858 , Fig. 33C View FIGURE 33 ). Twenty paratypes, same data as holotype ( AM C.337311 ).

Holotype of Siphonaria oblivirgulata Hubendick, 1945 , from Port Jackson [Sydney, Australia]; coll. Eugenie Exp., 1851–1853 ( UUZM 1575 , Fig. 33E View FIGURE 33 ).

Other, non-type material. Australia: NSW: Brunswick Heads, 28°32.297’S, 153°33.444’E, NSW12- 1 ( AM C.585592 4p, C.585068 p [M185]); GoogleMaps Cape Byron, 28°38’S, 153°38’E ( AM C.311682 5p); GoogleMaps Sand Point, N of Ballina, 28°50.66’S, 153°36.45’E ( AM C.343608 p); GoogleMaps MinniE, WAter, E of Grafton 29°46.6’S, 153°18’E ( AM C.343604 p); GoogleMaps N of Coffs Harbour, 30°14’S, 153°9’E ( AM C.311683 7p); GoogleMaps Nambucca Heads, 30°38.5’S, 153°1’E AM ( AM C.311681 7p); GoogleMaps Fingal Bay nr Port Stephens , 32°44.990’S, 152°10.481’,E, NSW09-1 ( AM C.585335 p); GoogleMaps Catherine Hill Bay , 33°9.3’S, 151°38’E ( AM C.343605 3p); GoogleMaps S end Catherine Hill Bay , 33°9.5’S, 151°38’E ( AM C.311679 6p); GoogleMaps Broken Head Terrigal, 33°26.796’S, 151°27.030’E, NSW08-1 ( AM C.585665 6p, C.585051 p [M044]); GoogleMaps Terrigal The Skillion, 33°27.008’S, 151°27.122’E, NSW08-2 ( AM C.585405 10+p, C.585055 p [M223], C.585056 p [M224]). GoogleMaps Sydney, Long Reef Collaroy, 33°44.6’S, 151°18.6’E ( AM C.343674 5p); GoogleMaps S side Long Reef Collaroy, 33°44.7’S, 151°19’E ( AM C.343678 3p; AM C.446108 6p); GoogleMaps Fairlight, North Harbour , 33°47.986’S, 151°16.837’E, NSW06-1 ( AM C.585470 16p); GoogleMaps Shelly Beach Headland Manly , 33°48’S, 151°17.5’E ( AM C.311680 5p); GoogleMaps North Harbour , SE side Reef Bay , 33°48.5’S, 151°16.38’E ( AM C.311687 7p). GoogleMaps Middle Harbour, Wy-ar-gine Point, 33°49’S, 151°15’E ( AM C.311678 10+p); GoogleMaps Wy-ar-gine Point, 33°49.159’S, 151°15.195’E, NSW06-5 ( AM C.585664 6p, 3d); GoogleMaps Edwards Beach Balmoral , 33°49.38’S, 151°15’E ( AM C.311890 10+p); GoogleMaps Balmoral, 33°49.7’S, 151°15.02’E ( AM C.343610 2p), GoogleMaps Laings Point Sydney Harbour, 33°50.419’S, 151°16.638’E, NSW06-3 ( AM C.585475 17p, C.585035 p [M162, SK035], C.585036 p [M163], C.585037 d [M164]); GoogleMaps Tamarama Point, 33°54’S, 151°16’E ( AM C.311684 7p; AM C.343615 2p). GoogleMaps Bombo Kiama, 34°39.232’S, 150°51.649’E, NSW03-1 ( AM C.585455 13p, C.584884 p [SK384], C.585280 p [SK048]); GoogleMaps Ulladulla, Wardens Head, 35°21’S, 150°29’E ( AM C.311677 7p); GoogleMaps Batemans Bay , Batehaven, 35°44’S, 150°12.5’E ( AM C.311685 5p); GoogleMaps Wimbie Beach, 35°47’S, 150°14’E ( AM C.343613 2p); GoogleMaps Eurobodalla Shire, Pretty Point SE facing, 35°48.28’S, 150°14’E ( AM C.343611 p); GoogleMaps Burrewarra Point, 35°50’S, 150°13.5’E ( AM C.343606 3p); GoogleMaps Mullimburra Point SE facing, 35°59.75’S, 150°9.58’E ( AM C.343607 5p); GoogleMaps Murunna Point Camel Head, 36°22.720’S, 150°04.766’E, NSW02- 1 ( AM C.585718 9p, C.585030 p [SK025], C.585332, d [SK026]); GoogleMaps Bermagui, 36°25.18’S, 150°3.78’E ( AM C.343612 2p); GoogleMaps Wapengo Lagoon estuary S side near Bithry Inlet, 36°37.7’S, 150°59’E ( AM C.395918 8p); GoogleMaps Aslings Beach, N end Twofold Bay , 37°3.1’S, 149°55’E ( AM C.148856 p); GoogleMaps Oman Point Eden, 37°04.634’S, 149°53.445’E, NSW01-1 ( AM C.585629 5p). GoogleMaps Twofold Bay , Murrumbulga Pt, 37°4.75’S, 149°53.06’E ( AM C.343622 2p, C.343623 p, C.343624 p, AM C.311688 4p); GoogleMaps Red Point, 37°6.083’S, 149°57.1’E ( AM C.343625 p); GoogleMaps Munganno Point, 37°6.2’S, 149°55.48’E ( AM C.343616 p, C.311689 4p, C.343617 3p, C.343618 2p, C.343619 3p, C.343621 3p, C.150599 p); GoogleMaps Fisheries Beach, 37°6.78’S, 149°55.6’E ( AM C.343620 p). GoogleMaps Green Cape, 37°15.8’S, 150°3’E ( AM C.343614 p, C.311676 5p); GoogleMaps Wonboyn Beach, Disaster Bay , 37°16’S, 149°57’E ( AM C.343733 2p); GoogleMaps Nadgee Fauna Reserve , N of Little River, 37°24’S, 149°57’E ( AM C.343735 3p). GoogleMaps Vic: Bastion Head Mallacoota, 37°34.429’S, 149°45.927’E, V09-1 ( AM C.585459 13p); GoogleMaps Cape Conran, 37°48.798’S, 148°43.608’E, V08-2 ( AM C.585610 4p); GoogleMaps Wilsons Promontory, 39°S, 146°22’E ( AM C.311674, 3 p); GoogleMaps Bear Gully, 38°53.519’S, 145°59.029’E, V07-3 ( AM C.585652 5p); GoogleMaps Flat Rock, Inverloch, 38°38.877’S, 145°41.638’E, V07-6 ( AM C.585573 3p); GoogleMaps Blow Hole, Western Port Bay nr. Flinders, 38°29’S, 145°1’E ( AM C.311673 2p); GoogleMaps Cape Schanck, 38°29.951’S, 144°53.369’E, V06- 4 ( AM C.585515 p [SK548]); GoogleMaps Port Phillip , Portsea, 38°19’S, 144°43’E ( AM C.311675 2p); GoogleMaps Point Lonsdale (nr Queenscliff), 38°17.276’S, 144°36.977’E, V05-1 ( AM C.585514 2p); GoogleMaps Roadknight Point, 38°25.707’S, 144°11.102’E, V04-1 ( AM C.585457 13p); GoogleMaps Loutit Bay Lorne, 38°31.190’S, 143°59.429’E, V03-2 ( AM C.585465 15p). GoogleMaps Tas: Flinders Island , Northeast River , 39°43.8’S, 147°57.6’E ( TMAGE542122 d); GoogleMaps PalanaBeach, 39°45.6’S, 147°52.8’E ( TMAG E54211 View Materials d); GoogleMaps Port Davies & Cave Beach , 40°0.6’S, 147°52.8’E ( TMAG E27150 View Materials d; TMAG E27142 View Materials 4d). GoogleMaps Boat Harbour, 40°57’S, 145°38’E ( AM C.311672 3p); GoogleMaps West Point , 40°57’S, 144°36’E AM ( AM C.311668 3p); GoogleMaps Little Peggs Beach, 40°51’S, 145°21.6’E ( TMAG E41987 View Materials d); GoogleMaps Rocky Cape: Picnic Beach & rocks to S, 40°52.2’S, 145°28.8’E ( TMAG E41989 View Materials d); GoogleMaps S of Granite Point Bridport, 40°59.739’S, 147°23.468’E, T01- 1 ( AM C.585251 p [M174]); GoogleMaps Bridport: beach, 41°24.6’S, 147°23.4’E ( TMAG E41986 View Materials d); GoogleMaps Bellingham, 41°0.6’S, 147°9.6’E ( TMAG E54215 View Materials 4d); GoogleMaps Greens Beach, 41°4.8’S, 146°45’E ( TMAG E54216 View Materials d). GoogleMaps The Gardens, Seatons Cove, 41°12.6’S, 148°16.8’E ( TMAG E41988 View Materials 3d); GoogleMaps Swimcart Beach, 41°13.8’S, 148°17.4’E ( TMAG E27411 4d, TMAG E54214 View Materials 5d, TMAG E41975 View Materials 3d). GoogleMaps Beaumaris, Shelly Point, 41°26.4’S, 148°16.8’E ( TMAG E41981 View Materials d); GoogleMaps Steels Beach, 41°28.2’S, 148°16.2’E ( TMAG E54217 View Materials d); GoogleMaps Bicheno, 41°52.837’S, 148°18.525’E, T02-1 ( AM C.585693 7p); GoogleMaps Bicheno, S end Redbill Beach, 41°53’S, 148°18’E ( AM C.311669 2p); GoogleMaps Maria Island , Darlington Bay , 42°34.8’S, 148°3.6’E ( TMAG E41984 View Materials 3d); GoogleMaps Dodges Ferry, 42°51.083’S, 147°36.981’E, T03-1 ( AM C.585462 14p); GoogleMaps Park Beach Dodges Ferry , 42°51.716’S, 147°36.665’E, T03-4 ( AM C.584883 p [SK138], C.585660 p [M111], C.585262 p [SK139]); GoogleMaps Lagoon Bch (near Saltwater River ), 42°56.903’S, 147°39.962’E, T03-2 ( AM C.585648 5p, C.595922 p [SK551]); GoogleMaps Blackmans Bay , 43°0.6’S, 147°19.8’E ( TMAG E15879 10+d); GoogleMaps Tasman Arch, 43°02.033’S, 147°56.963’E, T03- 3 ( AM C.585772 12p, C.585258 p [M116]); GoogleMaps Huon Point d’Entrecasteaux Channel, 43°17.471’S, 147°05.778’E, T04-1 ( AM C.585678 6p, C.585267 p [M106], C.585268 p [M135]); GoogleMaps South Bruny Island: Coal Point, 43°19.8’S, 147°19.8’E ( TMAG E35240 p); GoogleMaps Moss Glen, 43°31.910’S, 146°53.641’E, T05-1 ( AM C.585569 3p); GoogleMaps Flensing Rock, 43°34.291’S, 146°54.856’E, T05-2 ( AM C.585605 4p). GoogleMaps King Island, Little Porky Beach, 39°51’S, 143°51.6’E ( TMAG E41980 View Materials 2d), GoogleMaps Naracoopa, foreshore, 39°55.2’S, 144°7.2’E ( TMAG E41978 View Materials d), GoogleMaps Currie Harbour, 39°55.8’S, 143°50.4’E ( TMAG E41985 View Materials 10d), GoogleMaps Red Hut Point, 40°6’S, 144°6’E ( TMAG E41983 View Materials d) GoogleMaps .

Taxonomic remarks. The lectotype of S. funiculata has been referred to as the holotype by Jenkins (1981: 2), an act that qualifies as the designation of the lectotype. The lectotype ( Fig. 33A View FIGURE 33 ) closely matches the figure in the original description ( Reeve, 1856: pl. 2, fig. 6a–b). Reeve (1856: pl. 7, fig. 35a–b) erroneously labelled species 35 (= S. lirata ) as ‘ S. funiculata ’ (see erratum in the appendix). Our delineation of this species is based on comparative analyses of the morpho-anatomy and mitochondrial genetics of freshly collected topotypes of S. funiculata ( Fig. 33B View FIGURE 33 ), S. blainvillei ( Fig. 33P View FIGURE 33 ), S. virgulata ( Figs 33H, O View FIGURE 33 ), and S. oblivirgulata ( Fig. 33F View FIGURE 33 ) and a geographic series of additional specimens (Table S1).

Tenison Woods (1877: 58) incorrectly treated S. funiculata as a variety of S. diemenensis . Hutton (1878: 42) treated S. funiculata as a synonym of S. laeviuscula , but Hubendick (1946: 23) thought that Hutton’s (1878: 42) reference to S. laeviuscula was a misidentification of S. funiculata . Hedley (1915: 751) described S. virgulata indicating that Angas’ (1867: 232) record of ‘ S. funiculata ’ was in fact a misidentification of this new species. This conclusion was subsequently upheld by Hedley (1917b) and Hubendick (1946: 23), the latter also including a record of ‘ S. funiculata ’ by Adam & Leloup (1939: pl. 2, fig. 2a–b) from Pisang Island, PNG. Hedley (1915) and Jenkins (1981: pl. 1, fig. a) observed that S. blainvillei (based on one specimen) was a tall and broad ribbed specimen of S. funiculata . Hubendick (1946: 23) also treated S. blainvillei as a synonym of S. funiculata . Several tall specimens of S. funiculata have been collected as part of this work with a shell geometry resembling the type specimen of S. blainvillei and analyses of these specimens confirm this conclusion (e.g., AM C.585660, AM C.585251). Similarly tall individuals occur in other Siphonaria species, such as S. radiata , S. plicata , S. radians , S. sipho and S. monticulus .

Iredale (1924: 275–276) and Jenkins (1981) considered S. virgulata as a geographical variant of S. funiculata . Hubendick (1947b: 1) incorrectly considered S. funiculata Reeve 1856 as a synonym of S. pisangensis Hubendick 1947b (type locality Pisang Island, PNG). Grove et al. (2006: 60) correctly listed S. blainvillei , S. virgulata and S. oblivirgulata in the synonymy of S. funiculata . No records of S. funiculata are available from NZ based on our examinations of museum samples. The description below is based on the redescription in Jenkins (1981), which is expanded here for completeness and consistency.

External morphology ( Fig. 33T View FIGURE 33 ). Foot sole smooth, greyish yellow centrally, fading to become paler at foot edge; foot wall dark grey with evenly spread white subepithelial pustules becoming more vivid and dense close to the foot sole; genital pore inconspicuous, located on foot wall posterior to right cephalic fold; two small black epithelial eye spots centralised on two thick centrally touching dark grey to blueish cephalic folds, folds fade to a paler cream to yellowish at outer edge, covered with clustered white mucous cells similar (but smaller) to those of the foot wall tissue; fringing whitish mantle around top of foot wall, extends and increasingly transparent to shell edge, mantle edge thickened, lobed with an outer band of cream and brown pigmentation reflecting corrugations and inner colouration of shell lip and ribs; thin whitish pneumostomal lobe part of the mantle, between the right ADMs, closes the pneumostome and anus at the mantle edge.

Shell ( Fig. 33A–H, O, P, U View FIGURE 33 ; Table S9). Small to medium sized (max sl mean = 23.3 mm, SD = 4.45 mm, n = 30); height medium to tall; shell thickness medium; ovate; apical sides weakly convex, apex often eroded creating a white spot; protoconch direction homostrophic (n = 3; Fig. 33U View FIGURE 33 ), shell whorl dextral; apex offset weakly posterior and central; exterior sculpture finely costate with irregularly spaced radial ribs and growth striae; ribbing flat, broad with rib widths at shell lip ranging from 0.39 to 1.89 mm (mean = 0.82 mm, SD = 0.02 mm, n = 30), white axial ribs, often bistriate, chocolate coloured interstices, narrow and curve adapically; number of ribs variable, rib count (range 28–94; mean = 63, SD = 10.5, n = 60); siphonal ridge is weakly visible with fine, clustered, brown radial striae above a slight fold in the marginal lip; interior is polished and purplish brown with a white to blue spatula (colouration extends into the shallow siphonal groove) fading to a chocolate brown zone above the brown ADM impression and tan margin; marginal lip is shallowly scalloped with alternating chocolate brown and white radial markings, restricted to the lip margin and reflecting the exterior ribbing; ADM impression is horseshoe-shaped with a thin, lightly convex, anterior attachment area and a bare siphonal groove flanked by broad, ovate muscle impressions. Juvenile specimens have fine radial ribs, a dark brown interior and exterior (often obvious around the apex of adult specimens), with a white spatula.

Reproductive system ( Figs 34A, B, D; n View FIGURE 34 = 3). Epiphallic parts fill the region between RAM and BM, ED, GA, AO, EG white, smoothly rounded, possessing thick fibrous layers of tissue, ED larger than EG, often elongated; F1 very stubby, inconspicuous; GA large, bulbous, opening below the mantle on the side of the foot, behind right cephalic fold, anterior to pneumostome; AO very large, sack-like, joins GA; ED very short thick, joins to side of GA, EG larger than AO or GA; BD and CD enter GA very close together and AO, both pass through RAM ( BD above CD) are of similar length (although BD may be shorter); HG at posterior right quarter of coelom over the foot muscle tissue; HG usually yellow, granulated, linked by thin duct to the pinkish white, lobed, coiled HD, which in turn links to CD; SV partly lobed, uncoiled, pink to white, connected via thin duct alongside the AG to CD; AG / MG complex yellow to white, folded, lobed, closely attached to HG; SV embedded in folds; BC ovate, brown, patterned, tissue often expanded and stretched or collapsed and wrinkled; test thin, enclosing granulated, brown gelatinous mass.

Spermatophore ( Fig. 34C View FIGURE 34 ). Elongated drop shaped, test thin, smooth, featureless, translucent (length = 1.12 mm, n = 1), head spherical; flagellum very short; both sections smooth, featureless; head much larger than flagellum (head length = 0.93 mm, head ~83 % length of SPM, head width = 0.79 mm, flagellum width = 0.103 mm, n = 1); single SPM found in one BC ( AM C.584848). SPM matches SPM depicted in Jenkins (1983: fig. 3f).

Radula and jaw (figured in Jenkins 1983: 10, pl. 3a–h). Radula with a central tooth and longitudinally variable number of inner, mid and outer lateral teeth in longitudinal rows. Mean dentition formula 43:1:43 (SD = 7.9, n = 17) with around 120 transverse rows (SD = 14.9). These rows are parallel and slightly curved (anteriorly convex) Jenkins (1981: 9, pl.3a–h). Of the 43 half row laterals, 7 (SD = 4.2) are inner, 17 (SD = 7.6) mid and 18 (SD = 3.1) outer lateral teeth, respectively. The total number of lateral teeth appears related to the length of the shell (max. 54:1:54, shell length = 22.0 mm, mean 43:1:43, shell length = 18.1 mm, min 34:1:34, shell length = 15.1mm). However, the numbers of inner, mid and outer lateral teeth vary independently of animal dimensions and distributions. All teeth are bluntly concave posteriorly. The central tooth is narrow and weakly bicuspidate (often pointed) with a lower profile than the flanking laterals. The base is broad with adjacent central teeth. Mid and inner lateral teeth interlock with posteriorly and anteriorly aligned laterals. Outer laterals do not interlock between transverse rows. The space between rows increases to the ribbon edges associated with a gradual decrease in tooth size ( Jenkins 1981: 10, pl.3c, g). The space varies between individuals ( Jenkins, 1981: 10, pl.3d, h) as well as posterior and anterior areas of the ribbon. All lateral teeth are broad based and bicuspidate on the mesocone with a longer inner cusp. Outer lateral teeth are often multicuspidate. Increasing side denticle numbers, less elongated shape and increasingly stunted mesocones are transverse row features less accentuated from the central to the outer lateral teeth. Inner lateral teeth are elongated without flanking endo and ecto cones (inner and outer side denticles respectively). The more numerous outer lateral teeth have both ecto and endo cones while the mid laterals possess only an ectocone. The angle of separation from the mesocone of these side denticles is widely variable. Both side cones curve either towards or away from the mesocone ( Jenkins, 1981: 10, pl. 3 g, h). The length and width of the separation cleft is also widely variable, both generally increase towards the ribbon edge ( Jenkins, 1981: 10, pl.3c, d mid half ribbon area, pl. 3g, h ribbon edge). Aberrant outer lateral teeth are often present on both sides of the ribbon appearing as fused teeth with double mesocones. Not all individuals have inner lateral teeth, most have increased numbers of mid laterals. The number is independent of the number of lateral teeth, for example, of two radulae with 54 half row laterals, one had no inner laterals while the other had 18. The same variability was noted for radulae with fewer numbers of lateral teeth. Inner laterals do not possess endo or ectocones.

Comparative remarks. Siphonaria funiculata ( lateralis group, unit 8) is the sister species of S. lessonii , both together representing the sister lineage of S.tasmanica ( Figs 1 View FIGURE 1 , 4 View FIGURE 4 ). Siphonaria funiculata differs from S. lessonii by COI distances of ≥ 12% and from S. tasmanica by ≥ 8.5% (Table S8). Throughout its range, S. funiculata has been found in sympatry with nine congeners. For comparisons with S. diemenensis , S.denticulata , S. scabra , and S. zelandica refer to comparative remarks under these species. Siphonaria emergens has a much smaller, paler, orange-brown shell with less prominent ribbing, stronger edge scalloping and a strongly offset apex. Siphonaria pravitas sp. nov. has much paler brown shell with a more prominent siphonal ridge, raised ribbing, a darker spatula, a smaller AO, and a narrower, thread-like SPM. Siphonaria stowae has a much smaller, paler, yellowish cream shell with less prominent ribbing, stronger edge scalloping, a strongly offset apex, a smaller AO and BC, a shorter ED, and a narrower, thread-like SPM. Siphonaria jeanae has a smaller, grey-blue shell with slightly more raised brown ribbing, purplish spatula, an indistinct AO, narrower BD, and a more bulbous SPM. Siphonaria tasmanica has a grey-blue shell with a less distinct siphonal ridge, fainter ribbing, a smaller AO and BC.

A record of S. funiculata from NZ ( Hutton, 1873: 55) has been attributed subsequently to S. australis ( Jenkins, 1983: 13) . Based on similarity in reproductive anatomy (i.e., closeness of the duct joint, smallness of the genital atrium and greatly swollen epiphallus duct), Hubendick (1946: 23) assigned S. funiculata to Pachysiphonaria . The RS shown herein ( Figs 34A, B, D View FIGURE 34 ) correspond well with illustrations of the RS figures of S. funiculata elsewhere (e.g., Hubendick 1945: figs 3, 12; 1946: fig. 5, as ‘ S. virgulata ’; 1955: figs 1–2, as ‘ S. virgulata ’; Jenkins, 1983: figs 3a–c). The mean radula dentition formula observed herein is consistent with the 39:1:39 count given by Hubendick (1946: 23). A specimen figured as ‘ S. funiculata ’ from Lakshadweep, India by Ravinesh & Biju Kumar (2015: 38) is a misidentification of an unidentified species. Specimens depicted as ‘ S. funiculata ’ from Kelsi Coast, India in Vakani & Rahul Kundu (2021: 134, figs 2d, 3d) are misidentifications and are likely specimens of S. incerta sp. nov.

Distribution and habitat. Endemic to eastern and southeastern coasts of Australia, from Brunswick Heads, northern NSW, south to west of Lorne, Vic, including Tas ( Fig. 25 View FIGURE 25 ). In this study found to be common in sheltered places, such as crevices and vertical faces, on exposed rocky shores, upper to mid littoral levels, often associated with barnacles ( Fig. 33S View FIGURE 33 ).