Siphonaria restis, B. W. Jenkins & Köhler, 2024

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 : 177-179

publication ID	https://doi.org/10.11646/megataxa.13.1.1
DOI	https://doi.org/10.5281/zenodo.14989366
persistent identifier	https://treatment.plazi.org/id/0D49832F-FF2D-82AE-FF68-FF02FC00FC36
treatment provided by	Plazi (2025-03-05 09:04:49, last updated 2025-03-07 14:54:03)
scientific name	Siphonaria restis
status	sp. nov.

Treatment

Siphonaria restis View in CoL sp. nov.

( Figs 71D–H, O–P View FIGURE 71 , 72B–D View FIGURE 72 )

Siphonaria kurracheensis View in CoL — Knox 1955: 88; Roberts & Wells 1981: 72, pl. 30, figs 142, 143; Black & Johnson 1981: 79; Jenkins 1983: 28; 1984: 117; Wells 1984: 54; Johnson & Black 1997: 104; Chim & Tan 2009: 269; Willan et al. 2015: 330 (not S. kurracheensis Reeve, 1856 View in CoL ).

Siphonaria luzonica View in CoL — Hubendick 1955: 3, fig 3; Hodgkin et al. 1966: 27, pl. 8, fig. 3 (not S. luzonica Reeve, 1856 View in CoL ).

Siphonaria sp. — Verco 1912 b: 205; Wells & Bryce 1997: 379.

Material examined. Type material. Holotype, from Chinamans Rock , 27°42.776’S, 114°09.361’E, Kalbarri, WA; coll. B.W. Jenkins, WA52-1, 20 Nov 2017 ( WAM S74049 [M400], Fig. 71D View FIGURE 71 ). GoogleMaps

Paratypes same as holotype ( WAM S74079 6p, S74048 p [M100], Fig. 71E View FIGURE 71 , AM C.584731 11p, C.585200 p [M313], Fig. 71F, C.585919 View FIGURE 71 p [SK150], Fig. 71G View FIGURE 71 ) .

Other, non-type material. Australia, WA: Chinamans Rock, Kalbarri, 21°54.739’S, 113°58.706’E; coll. B.W. Jenkins, WA52-1, 20 Nov 2017 ( WAM S74047 View Materials p [M099], Fig. 54D, S View FIGURE 54 4078 p [SK050]); GoogleMaps Thompson Bay , Rottnest Is, RI 012 ( AM C.585201 p [SK381], Fig. 54E, C View FIGURE 54 .585202 d [SK382], C.595909 d [R2004]); GoogleMaps Tantabiddi, 21°54.739’S, 113°58.706’E, WA42-1 ( AM C.585656 5p, WAM S74045 View Materials p [SK036]); GoogleMaps Cape Latouche Treville nr Gourdon Bay , 18°27.101’S, 121°48.911’E, WA27-2 ( AM C.584687 5p, WAM S74135 5p); GoogleMaps 18°27.457’S, 121°48.725’E, WA27-1 ( AM C.584711 5p, WAM S74136 2p); GoogleMaps Cape Keraudren, 19°57.393’S, 119°46.358’E, WA29-2 ( AM C.584704 5p, WAM S74138 View Materials 2p); GoogleMaps Withnell Bay Dampier, 20°35.106’S, 116°47.252’E, WA33-1 ( AM C.584752 5p, WAM S74141 5p); GoogleMaps Point Samson, 20°36.655’S, 117°11.472’E, WA32- 2 ( AM C.584781 8p, WAM S74140 View Materials 8p); GoogleMaps 20°36.684’S, 117°11.303’E, WA32-1 ( AM C.584751 5p, WAM S74139 5p); GoogleMaps Dampier Hbr, 20°39.920’S, 116°42.134’E, WA33- 3 ( AM C.585575 3p); GoogleMaps Gnoorea Point, 1 20°50.560’S, 116°21.804’E, WA36-1 ( AM C.584689 3p, WAM S74142 View Materials 2p); GoogleMaps Beardon Pt Onslow, 21°37.860’S, 115°06.573’E, WA37-1 ( AM C.584722 5p, WAM S74143 View Materials 4p); GoogleMaps NW Cape Exmouth, 21°48.360’S, 114°07.665’E, WA41-1 ( AM C.584698 6p, WAM S74144 3p); GoogleMaps Fly Island , Great Sandy Islands , 21°48’19.548’‘S, 114°33’9.36’‘E ( WAM S97276 View Materials p); GoogleMaps Tantabiddi, 21°54.739’S, 113°58.706’E, WA42-1 ( AM C.584690 5p, d; WAM S74145 View Materials 2p); GoogleMaps Pt S of Bruboodjoo Pt Bateman Bay, 23°02.991’S, 113°49.371’E, WA43- 1 ( AM C.584732 12p, WAM S74146 5p); GoogleMaps Point Maud, 23°08.322’S, 113°46.294’E, WA44-1 ( AM C.584753 5p, C.585919 p [SK150], WAM S74147 5p); GoogleMaps Coral Bay , S of Exmouth Gulf, 23°08.322’S, 113°46.294’E ( AM C.595919 p [SK004], C.959956 p [SK001]); GoogleMaps N of Point Quobba, 24°26.288’S, 113°24.204’E, WA45-2 ( AM C.584727 7p, WAM S74149 View Materials 5p); GoogleMaps Point Quobba , 24°29.124’S, 113°24.501’E, WA45-1 ( AM C.584754 5p, WAM S74148 View Materials 5p); GoogleMaps Bottle Bay Cape Peron, 25°32.566’S, 113°29.467’E, WA49-1 ( AM C.584755 5p, WAM S74150 5p); GoogleMaps Islet off Eagle Bluff, Shark Bay , Peron Peninsula, 26°05.5’S, 113°34.4’E ( WAM S72339 6p); GoogleMaps Whalebone , 26°07.835’S, 113°38.391’E, WA47-2 ( AM C.585657 5p); GoogleMaps Kells Rock Shark Bay , 26°10.473’S, 113°12.415’E, WA50-3 ( AM C.584734 12p, WAM S74151 View Materials 6p); GoogleMaps Pepper Point (Zuytdorp), 26°23.826’S, 113°18.268’E, WA51-1 ( AM C.584725 6p, d, WAM S74152 View Materials 6p); GoogleMaps Chinamans Rock , Kalbarri , 27°42.776’S, 114°09.361’E ( AM C.585489 p [M053], C.585490 p [M398], C.585491 p [M399], C.585920 p [SK151]); GoogleMaps Red Bluff , 27°44.627’S, 114°08.576’E, WA52-2 ( AM C.595908 p [M125], C.585365 p [M126]); GoogleMaps Horrocks , 28°21.469’S, 114°24.751’E, WA53-1 ( AM C.584680 2p; WAM S74153 View Materials 2p); GoogleMaps Turtle Bay East Wallabi Is , 28°25.804’S, 113°44.538’E, WA55-1 ( AM C.584758 5p; WAM S74155 5p); GoogleMaps Houtman Abrolhos Islands , 28°25.804’S, 113°44.538’E ( AM C.595971 2p), GoogleMaps S of Fish Point East Wallabi Is, WA55-2 28°25.816’S, 113°44.633’E ( AM C.584720 7p; WAM S74156 2p); GoogleMaps Cape Burney Geraldton , 28°52.084’S, 114°38.056’E, WA54-1 ( AM C.584749 5p; WAM S74154 View Materials 5p); GoogleMaps Leander Point Port Denison , 29°16.568’S, 114°54.858’E, WA57-1 ( AM C.584760 5p, WAM S74157 View Materials 5p); GoogleMaps S end Leander Point Port Denison , 29°16.725’S, 114°54.918’E, WA57-2 ( AM C.585517 d); GoogleMaps Freshwater Point , 29°36.256’S, 114°58.464’E, WA57-3 ( AM C.584693 3p, C.585009 p [SK063], WAM S74158 View Materials 2p); GoogleMaps Illawong Bch rocks, 29°42.198’S, 114°57.551’E, WA57-4 ( AM C.584683 2p, WAM S74159 2p); GoogleMaps Illawong , 29°42.254’S, 114°57.542’E, WA57-5 ( AM C.584694 3p; WAM S74160 2p); GoogleMaps Jurien Bay , 30°17.244’S, 115°02.482’E, WA58-1 ( AM C.584679 2p; WAM S74161 p); GoogleMaps Grey , 30°39.968’S, 115°08.072’E, WA58- 2 ( AM C.585366 p); GoogleMaps Cape Leschenault , 31°17.508’S, 115°27.089’E, WA58-3 ( AM C.584712 5p, WAM S74162 View Materials 5p); GoogleMaps Quinns Rock , 31°39.822’S, 115°41.345’E, WA58- 4 ( AM C.585368 p); GoogleMaps Longreach Bay Point , Rottnest Is , 31°59.333’S, 115°32.063’E RI01 ( AM C.585600 4p, C.585198 p [M311], C.585199 p [M312], C.585200 p [M313], C.584943 p [SK154], Fig. 71H View FIGURE 71 ); GoogleMaps Point Brown Swan River , 32°02.344’S, 115°45.471’E, WA59-5 ( AM C.585369 p [M128]); GoogleMaps Fremantle Hbr , breakwater, 32°03.342’S, 115°43.987’E, WA58-5 ( AM C.585519 d); GoogleMaps Yallingup , 33°38.358’S, 115°01.481’E, WA60-9 ( AM C.584724 6p, WAM S74163 5p); GoogleMaps Cowaramup Point , 33°51.934’S, 114°58.904’E, WA60-3 ( AM C.585370 p); GoogleMaps Sarge Bay Cape Leeuwin, WA60-4 ( AM C.585809 p); GoogleMaps Yanchep, sheltered limestone cliffs, 34°32.783’S, 115°37.962’E ( AM C.595972 p) GoogleMaps .

External morphology ( Fig. 71O View FIGURE 71 ). Foot wall usually without irregular black blotches, may be weakly present in some individuals; edge, pneumostomal lobe and cephalic folds all evenly brownish yellow, without any black/darker pigmentation markings, foot sole maybe dark brown/grey, foot edge paler; mantle translucent to transparent, as wide as foot wall, weakly lobed, aligns with undulations of primary shell ribs; the mantle edge thickened with yellowish band, may show dark pigmentation markings aligned with rib interstices; pneumostome wide between right adductor muscles and within mantle; cephalic folds thickened.

Shell ( Figs 71D–H, P View FIGURE 71 ; Table S9). Medium sized (max sl mean = 17.5 mm SD = 2.7 mm, n = 19); height low to medium; ovate; apex central, apical sides weakly convex; protoconch direction weakly homostrophic (n = 1, Fig. 71P View FIGURE 71 ), shell whorl dextral, protoconch area distinctly darker brown; growth lines distinct; rib count (mean = 42, SD = 8.7, n = 19), weakly raised pale white to cream, weakly extend beyond shell edge, rib interstices dark brown; 10–12 evenly spread primary ribs; siphonal ridge prominent raised rounded, formed by 3 primary ribs; shell lip uneven, scalloped. Interior shell lip cream with brown splashes aligning with rib interstices, paler than shell margin, ADM scar prominent; spatula cream sometimes bluish; shell lip often golden brown with dark brown markings flanking shallow golden brown siphonal groove; CMS weakly concave, similar but darker colouration to spatula and shell margin; thickening of inner shell lip and spatula occurs in larger specimens, whitening covers brown colouration of inner shell lip.

Reproductive system ( Figs 72B, D; n View FIGURE 72 = 2). Positioned within right side of coelom,against foot wall on foot muscle, under the respiratory cavity; epiphallic parts positioned between RAM and BM. GA relatively small, with singular GP through foot wall; AO medium sized, broad, bluntly pointed, central bend, joined to upper GA; ED elongated, broad, centrally twisted, joins to side of GA; GA, AO, ED all white muscular fibrous tissue; EG medium sized, soft whitish tissue, slightly folded, joins at junction of end of ED and extension of single broad long flagellum (F1), end often looped; BD and CD connect closely together into GA between ED join and GP, both ducts long, slightly bent, smooth, narrow, whitish, featureless, pass closely together through outer side of RAM ( BD over CD) into soft white folded tissues of MG; MG / AG complex large; CD connecting to ducts, BD without distal loop, often with loop or fold immediately in front of BC; BC embedded in MG folds, close to embedded SV; BC medium, thin whitish translucent test; HD distinct, short, coiled, links ducts in soft white folded tissues of AG to yellowish granulated HG; outer edge of MG lobbed; AG and HG of similar size, sides match curvature of inner foot wall.

Spermatophore ( Fig. 72C View FIGURE 72 ). Thread-like, test thin, translucent (length = 6.38 ± 3.71, n = 3), flagella incomplete, head section cylindrical, tip bulbous bluntly rounded, containing a white gelatinous core, tapers into the filamentous transparent flagellum, both sections smooth, featureless, head much thicker than flagellum (length = 5.90 ± 3.02, n = 3, ~71% of SPM length, head width = 119 ± 33 μm, n = 3), two SPM embedded in brown gelatinous mass in one bursa ( AM C.584943).

Radula ( Figs 83I–L View FIGURE 83 ). Dentition formula 32:1:32 (n = 1, AM C.320123); single central rachidian tooth short broad, flanked squarely by 32 half row laterals, 10 are inner, 6 mid and 16 outer laterals; number of transverse rows not counted; central tooth short wide with short unicuspid mesocone; inner laterals (without endo or ectocones) prominent, mesocones of inner and mid laterals bicuspidate ( Figs 83I–L View FIGURE 83 ), mid laterals with broad pointed ectocone; outer, laterals typically with a ‘chisel’ shaped mesocone, often weakly bicuspidate, flanked by small, pointed single ecto and endocones, angle of separation of each cone from the mesocone varies and maybe inwardly curved ( Fig. 83K View FIGURE 83 ).

Comparative remarks. Siphonaria restis sp. nov. ( atra group, unit 54) forms a well-differentiated lineage in clade G of the mitochondrial tree ( Figs 1 View FIGURE 1 , 2 View FIGURE 2 ). It differs from other species by COI distances of ≥ 21.8% (Table S4). Throughout its range, we found S. restis sp. nov. in sympatry with seven congeners. Two are species are sympatric in south-western WA: For comparisons with S. stowae and S. jeanae refer to comparative remarks under these species. Three are sympatric in northern WA: For comparisons with S. atra , S. viridis , and S. gemina sp. nov. refer to comparative remarks under these species One species is sympatric in western WA: Refer to comparative remarks under S. zelandica . The RS figure of ‘ S. luzonica’ in Hubendick (1955: 3,fig 3) from Rottnest Is, corresponds well with the RS of S. restis sp. nov. shown here (i.e., parallel junction of BD and CD into GA, relative size of AO and twist in BD close to BC, Fig. 72B View FIGURE 72 ), and not that of S. sipho , the senior synonym of S. luzonica .

Distribution and habitat. Recorded as endemic to western and northern coasts of WA, Indian Ocean ( Fig. 73 View FIGURE 73 ). In this study, commonly found in sheltered positions on moderately exposed rocky shores, at upper and mid littoral levels.

Etymology. From the Latin ‘restis’ meaning ‘cord’— referring to the cord-like primary ribs on the shell of this species.

References

Black, R. & Johnson, M. S. (1981) Genetic differentiation independent of intertidal gradients in the pulmonate limpet Siphonaria kurracheensi s. Marine Biology, 64, 79-84. https://doi.org/10.1007/BF00394083

Chim, C. K. & Tan, K. S. (2009) Vertical distribution, spawning and recruitment of Siphonaria guamensis (Gastropoda: Pulmonata) on a seawall in Singapore. Raffles Bulletin of Zoology, Suppl. 22, 269-278.

Hodgkin, E. P., Kendrick, G., Marsh, L. & Slack-Smith, S. (1966) The shelled gastropoda of south-western Australia. Western Australia Naturalists' Club, Perth, 59 pp.

Hubendick, B. (1955) On a small quantity of Siphonaria material from Queensland. Memoirs of the National Museum of Victoria, 19, 126-136. https://doi.org/10.24199/j.mmv.1955.19.04

Jenkins, B. W. (1983) Redescriptions and relationship of Siphonaria zelandica Quoy and Gaimard to S. australis Quoy and Gaimard with a description of S. propria sp. nov. (Mollusca: Pulmonata: Siphonariidae). Journal of the Malacological Society of Australia, 6 (1 - 2), 1-35. https://doi.org/10.1080/00852988.1983.10673952

Jenkins, B. W. (1984) A new siphonariid (Mollusca: Pulmonata) from south-western Australia. Journal of the Malacological Society of Australia, 6 (3 - 4), 113-123. https://doi.org/10.1080/00852988.1984.10673964

Johnson, M. S. & Black, R. (1997) Distributions of high-intertidal gastropods in the Houtman Abrolhos Islands. In: Wells, F. (Ed.), The Marine Flora and Fauna of the Houtman Abrolhos Islands, Western Australia. Vol 1. Proceedings of the Seventh International Marine Biological Workshop. Western Australian Museum, 566 pp.

Knox, G. A. (1955) The Development of Kergulenella stewartiana (Powell) (Gastropoda: Siphonariidae). Pacific Science, 9 (1), 85-91.

Reeve, L. A. (1856) Monograph of the genus Siphonaria. In: Reeve, L. A. (Ed.), Conchologia Iconica, or, illustrations of the shells of molluscous animals, vol. 9. L. Reeve, London, unpaginated text, pls. 1-7.

Roberts, D. & Wells, F. (1981) Seashells of Western Australia. Creative Research, 91 pp.

Verco, J. C. (1912) Notes on the marine shells of Western Australia, with descriptions of new species. Part II. Transactions and Proceedings of the Royal Society of South Australia, 36, 192-205.

Wells, F. E. (1984) A guide to common molluscs of South-Western Australian estuaries. Western Australian Museum, Perth, 111 pp.

Wells, F. E. & Bryce, C. W. (1997) A preliminary checklist of the marine macromolluscs of the Houtman Abrolhos Islands, Western Australia. In: Wells, F. E. (Ed.), Proceedings of the Seventh International Marine Biological Workshop; The marine flora and fauna of the Houtman Abrolhos Islands, Western Australia. Western Australian Museum, Perth, pp. 363-383.

Figures

FIGURE 1. Maximum Likelihood phylogram based on analyses of a concatenated sequence data set of 16S and COI. Branches are collapsed at the species level. Branch labels give unit numbers and accepted species names. Numbers on branches indicate branch support employing 10,000 ultrafast bootstraps.Available genus-group names are shown next to their type species. Scale bar indicating modelled sequence divergence.

FIGURE 2. Maximum Likelihood phylogram (partial, species not collapsed). Clades G–I (atra group) of the tree shown in Fig. 1. Branch labels give specimen identifiers for new sequences or Genbank accession numbers for imported sequences from other studies and geographic regions (seeTables S1–S2 for details). Identical haplotypes are merged into single tips. Numbers on branches indicate branch support by employing 10,000 ultrafast bootstraps. Clade names give unit numbers and accepted species names. Scale bar indicating modelled sequence divergence. Figure spread over two pages.

FIGURE 54. Shells of S. campestra sp. nov., S. caubianensis sp. nov., S. christmasensis sp. nov. and S. costellata sp. nov. A–C, M–N. S. campestra sp. nov., Timor-Leste, Dili. A. Holotype AM C.584823 [M447, SK230]. B. Paratype AM C.585354 [SK266]. C. Paratype AM C.585355 [SK267]. M. In situ. N. Animal. D–E, O–P. S. camura sp. nov., Okinawa, Tancha Bay. D. Holotype AM C.585614 [M491, SK310]. E. Paratype AM C.584920 [M490, SK309]. O. In situ. P. Protoconch, AM C.585613 [SK510]. F–G. S. caubianensis sp. nov., F. Philippines, Caubian Is., holotype AM C.595933 [SK560]. G. Philippines, NW Polillo Is, WAM S113803 [M600, SK557]. H–J, Q–R. S. christmasensis sp. nov., CI, Flying Fish Cove. H. Holotype AM C.595957 [M298]. I. Paratype WAM S74040 [SK069]. J. Paratype, Ethel Bch AM C.584846 [M308]. Q. Protoconch AM C.585319 [SK071], R. CI, in situ. K–L, S–T. S. costellata sp. nov. K. Singapore, Lazarus Island, Holotype AM C.585236 [M416, SK100]. L. Paratype AM C.585226 [M338]. S. Protoconch, AM C.585233 [SK099]. T. In situ. Unlabelled scale bars 10 mm.

FIGURE 71. Shells of S. pravitas sp. nov., S. recurva sp. nov., S. restis sp. nov., S. rodriguensis sp. nov. and S. striata sp. nov. A–B, M–N, S. S. pravitas sp. nov. A. Holotype AM C.585040 [M192, SK118]. B. Paratype AM C.585038 [M153]. M. Sydney Harbour, in situ. N. Sydney Harbour, animal, S. Protoconch, AM C.546766 [SK432]. C. Holotype of S. recurva NMHN IM-2013- 55336 [M534]. D–H, O–P. S. restis sp. nov., WA, Kalbarri. D. Holotype WAM S74049 [M400]. E. Paratype WAM S74048 [M100]. F. Paratype AM C.585200 [M313]. G. Paratype AM C.585919 [SK150]. H. Rottnest Is, AM C.584943 [SK154]. O. Animal. P. Protoconch, AM C.585009 [SK063]. I, Q–R. S. rodriguensis sp. nov. I. Holotype AM C.585197 [SK330]. Q. Protoconch, AM C.585196 [M427]. R. In situ. J–L, T. S. striata sp. nov., Madagascar, Itampolo. J. Holotype AM C.584952 [M264]. K. Paratype AM C.584953. L. Paratype AM C.584954. T. Protoconch, paratype AM C.584953 [M265]. Scale bars = 10 mm.

FIGURE 72. Reproductive morphology of S. recurva sp. nov., S. restis sp. nov., S. rodriguensis sp. nov. and S. striata sp. nov. A. Holotype of S. recurva sp. nov. MNHN IM-2013-55336 [M534]. B–D. S. restis sp. nov. B. WA, Kalbarri, holotype WAM S74049 [M400]. C. WA, Rottnest Is, AM C.584943 [SK154]. D. WA, Point Maud, AM C.585919 [SK150]. E–F. Holotype of S. rodriguensis sp. nov. AM C.585197 [SK330]. G–H. Holotype of S. striata sp. nov. AM C.584952 [M264]. Scale bars = 1 mm.

FIGURE 73. Known occurrence records of S. pravitas sp. nov., S. recurva sp. nov., S. restis sp. nov., S. rodriguensis sp. nov., S. striata sp. nov., S. tagaqaensis sp. nov., S. tanchaensis sp. nov. and S. tanguissonensis sp. nov.

FIGURE 83. Radula morphology of S. diemenensis, S. denticulata, S. restis sp. nov. and S. tasmanica. A–D. S. diemenensis, Tas, Recherche Bay, AM C.201772. A. Inner central area x256. B. central area x585. C. Outer laterals x256. D. Shell. E–H. S. denticulata, NSW, Sydney, AM C.315783 x520. E. Inner central area. F. Central area. G. Outer laterals. H. Shell. I–L. S. restis sp. nov. WA, Rottnest Is, AM C.320123. I. Inner central area x234. J. Central area x466. K. Outer laterals x466. L. Shell. M–P. S. tasmanica, Tas, Eaglehawk Neck, AM C.311615. M. Inner central area x505. N. Central area x505. O. Outer laterals x650. P. Shell. Unlabelled scale bars 1 mm.