Siphonaria Sowerby I, 1823

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 : 19-21

Siphonaria Sowerby I, 1823

Siphonaria Sowerby I 1823 (in 1821–34):126, pl. 122.

Type species: Siphonaria sipho Sowerby I 1823 , by subsequent designation of Gray (1847: 181).—Numerous subsequent references.

Mouretus Blainville 1824b: 161–162 . Type species: Mouretus adansonii Blainville, 1824 , by monotypy.—Iredale 1921: 207; Morrison 1963: 8; 1972: 52–53; Christiaens 1980a: 78 (all as subgenus of Siphonaria ).

Liria Gray 1824: 275 . Unavailable (introduced in synonymy of Siphonaria , and not treated as available before 1961; Art. 11.6 of the Code).

Liriola Dall 1870: 32 . Type species: Siphonaria thersites Carpenter, 1864 , by original designation; as subgenus of Siphonaria .— Dall 1870: 32–33; Suter 1909a: 33, 1913: 601; Dall 1926: 26; Thiele 1931: 472; Hubendick 1945: 60; 1946: 8–9, 18–19; Tomlin 1948: 229; Powell 1951: 65; Morrison 1963: 7; Abbott 1974: 335; Christiaens 1980a: 80; Trew 1983: 3; Redfern 2001: 759.

Liphonaria Brazier 1878: 135. Invalid; incorrect subsequent spelling of Siphonaria .

Patellopsis Nobre 1886: 32 . Type species: Siphonaria algesirae Quoy & Gaimard, 1833 , by monotypy; as subgenus of Siphonaria .— Hubendick 1945: 60, 70; 1946: 9, 12–13, 30– 35 (as section of subgenus Liriola ); Abbott 1974: 335.

Kerguelenia Mabille & Rochebrune in Rochebrune & Mabille 1889: H.27–H.28. Invalid; junior homonym of Kerguelenia Stebbing, 1888 [Crustacea], Kerguelenella is a replacement name.— Suter 1909a: 34; Iredale 1915: 478; Hedley 1916a: 61, 62; 1917b: 96; Dall 1921: 67; May 1921: 89; Gatliff & Gabriel 1922: 56; May 1923: 87; Odhner 1924: 55; Finlay 1927: 442; Powell 1937: 68; 1939: 217 238; Hubendick 1945: 66–67, 70; 1946: 9, 11–12, 18–19, 24, 26–30, 35 (as a section of Liriola ); Powell 1955: 122; Marcus & Marcus 1960: 114 (as subgenus of Siphonaria ); Dell 1964: 290; Abbott 1974: 335 (as subgenus of Siphonaria ); Powell 1979: 293; White & Dayrat 2012: 64–65, 68.

Pugillaria Iredale 1924: 277 . Type species: Siphonaria stowae Verco, 1906 , by original designation.— Thiele 1931: 472 (as subgenus of Siphonaria ); Cotton & Godfrey 1932: 155; Macpherson & Chapple 1951: 142; Cotton 1959: 411; Zilch 1959: 85; Iredale & McMichael 1962: 82; Macpherson & Gabriel 1962: 264; Morrison 1963: 7–8 (as section of Liriola ); Shikama 1964: 6 (as subgenus of Siphonaria ); Macpherson 1966: 256; Trew 1983: 3; White & Dayrat 2012: 61; Jenkins 2018: 276, 279, 281.

Benhamina Finlay 1927: 442 , 478. Type species: Siphonaria obliquata Sowerby I, 1825 , by original designation.— Thiele 1931: 472 (as subgenus of Siphonaria ); Powell 1937: 86; 1939: 217; Hubendick 1945: 60, 66; 1946: 9, 11, 18, 22, 25, 64 (as section of Liriola ); Powell 1946: 91; Borland 1950: 385–393; Knox 1954: 872; 1955: 86; Powell 1955: 120; 1956: 120; 1957a: 114; Marcus & Marcus 1960: 108; Shikama 1964: unnumbered (as subgenus of Siphonaria ); Morton & Miller 1968: 302, 336, 338, 354, 378, figs 121–122, pl. 19, figs 10–10a; Berry 1977: 204, 211; Powell 1979: 293, pl. 54, figs 12–13, Trew 1983: 2; Morley 2004: 130; Russell & Phillips 2009: 579.

Mestosiphon Iredale 1940: 437 , 439. Type species: Mestosiphon eumelas Iredale, 1940 , by original designation.— Habe & Kikuchi 1960: 64; Habe & Kosuge 1966: 113 (as Mestasiphon, incorrect subsequent spelling of Mestosiphon ); Yoo 1967: 89 (as Mastosiphon, incorrect subsequent spelling of Mestosiphon ) (all as subgenus of Siphonaria ); Morrison 1972: 54, 56 (as section of Siphonaria s. str.); Habe & Kohno 1980: 23; Jenkins 1983: 29; White & Dayrat 2012: 63, 64, 66.

Mallorisiphon Iredale 1940: 437 , 440. Type species: Mallorisiphon oppositus Iredale, 1940 , by monotypy.— Morrison 1972: 57 (as synonym of Mestosiphon ); White & Dayrat 2012: 66, 70.

Ellsiphon Iredale 1940: 437 , 438. Type species: Ellsiphon marza Iredale, 1940 , by original designation.— McAlpine 1952: 40, 42.

Triellsiphon Iredale 1940: 437 , 439. Type species: Triellsiphon acervus Iredale, 1940 , by monotypy.— Hubendick 1945: 29; White & Dayrat 2012: 60.

Parellsiphon Iredale 1940: 437 , 438. Type species: Parellsiphon zanda Iredale, 1940 , by original designation.— Hubendick 1945: 29; 1946: 30–31, 41, 61; Morrison 1972: 57; Jenkins 1983: 29; White & Dayrat 2012: 61, 64, 67, 69.

Legosiphon Iredale 1940: 437 , 440. Type species: Legosiphon optivus Iredale, 1940 , by original designation.— Hubendick 1955: 7, 132; White & Dayrat 2012: 62, 65–66.

Hebesiphon Iredale 1940: 437 , 441. Type species: Hebesiphon monticulus Iredale, 1940 , by monotypy.— Morrison 1972: 51, 61; White & Dayrat 2012: 65.

Torquisiphon Iredale 1940: 437 . Type species: Torquisiphon percea Iredale, 1940 , by monotypy.— Hubendick 1946: 63; White & Dayrat 2012: 66.

Talisiphon Iredale 1940: 441 , 442. Type species: Siphonaria virgulata Hedley, 1915 , by original designation.— Cotton 1959: 411; Iredale & McMichael 1962: 82; Morrison 1963: 7 (as section of Liriola ); Marincovich 1973: 42; White & Dayrat 2012: 66, 69.

Sacculosiphonaria Hubendick 1945: 61 , 72. Type species: Patella japonica Donovan, 1834 , by original designation; as section of Siphonaria .— Hubendick 1946: 41–43 (as subgenus of Siphonaria ); Knox 1955: 88; Kira 1962: 201; Habe & Kikuchi 1960: 64; Marcus & Burch 1965: 256; Habe 1971: 15; Kuroda et al. 1971: 302; Christiaens 1980b: 466; Inaba 1983: 145; Fukuda et al. 1992: 76; Trew 1983: 9.

Simplisiphonaria Hubendick 1945: 61 , 72. Type species: Siphonaria cookiana Suter, 1909 , by original designation; as section of Siphonaria . — Hubendick 1946: 36; Trew 1983: 8 (as subgenus of Siphonaria ).

Pachysiphonaria Hubendick 1945: 12 , 15, 16, 60, 64, 72. Type species: Siphonaria lessonii Blainville, 1827 , by original designation; as section of Liriola .— Hubendick 1946: 20–22, 24; Powell 1951: 65, 1960: 163; Shikama 1964: 7; Rios 1970: 140; Dell 1971: 214; Trew 1983: 2, 5 (as genus); Ludbrook & Gowlett-Holmes 1989: 612.

Ductosiphonaria Hubendick 1945: 61 , 70. Type species: Siphonaria bifurcata Reeve, 1856 , by original designation; as section of subgenus Siphonaria .— McAlpine 1952: 42; Hubendick 1955: 4–6; Cotton 1959: 411.

Heterosiphonaria Hubendick 1945: 61 . Type species: Siphonaria gigas Sowerby I, 1825 , by original designation; as section of Siphonaria .— Hubendick 1945: 61, 70, 72; 1946: 13, 15, 39–41, 65, 66, 1955: 6; Morrison 1963: 8; Shikama 1964: 6; Keen 1971: 850, 852; Trew 1983: 8–9.

Kerguelenella Powell 1946: 91 (nom. nov. pro Kerguelenia Mabille & Rochebrune, 1889 , non Stebbing, 1888 [Crustacea]). Type species: Siphonaria redimiculum Reeve, 1856 , by original designation.— Powell 1951, 55; Carcelles 1953: 206; Knox 1955: 85; Powell 1955: 120–122; 1957a: 114; 1957b: 137; Boreham 1959: 72; Powell 1960: 163; Dell 1963: 227; 1964: 290; Morton & Miller 1968: 313, 379, 380; Dell 1971: 215; Gaillard 1971: 278; Arnaud 1974: 542; Burn & Bell 1976: 234; Simpson 1977: 126, 132; Powell 1979: 293; Christiaens 1980a: 80 (as subgenus of Siphonaria ); Branch et al. 1991: 55; Forcelli 2000: 132; Griffiths & Waller 2016: 1147.

Anthosiphonaria Kuroda & Habe 1952: 302 . Type species: Siphonaria sirius Pilsbry, 1894 , by original designation.— Christiaens 1980a: 79; Inaba 1983: 145; Habe et al. 1986: 23; Je 1989: 29; Fukuda et al. 1992: 76.

Hubendickula McAlpine 1952: 42 . Type species: Siphonaria diemenensis Quoy&Gaimard,1833 , byoriginaldesignation.— Cotton 1959: 411; Ludbrook & Gowlett-Holmes 1989: 610, fig. 11 (as subgenus of Siphonaria ).

Hubendicula Hubendick 1955: 5–6. Invalid; incorrect subsequent spelling of Hubendickula .

Hubendiculata Iredale & McMichael 1962: 82. Invalid; incorrect subsequent spelling of Hubendickula .

Siphonacmea Habe 1958b: 35 . Type species: Acmaea oblongata Yokoyama, 1926 , by original designation.— Habe 1962: 96, pl. 44, fig. 15; 1964: 144, pl. 44, fig. 14; Toyohara et al. 2001: 27–35.

Planesiphon Zilch 1959: 86 . Type species: Planesiphon elegans Iredale, 1940 , by original designation (as subgenus of Siphonaria ).— Hubendick 1946: 23; Marcus & Marcus 1960: 122; Iredale & McMichael 1962: 82; Kuroda et al. 1971: 303; Jenkins 1983: 5, 12; Inaba 1983: 145; Habe et al. 1986: 23; Je 1989: 29; White & Dayrat 2012: 63, 67.

Sacculosiphon Je 1989: 29. Invalid; incorrect subsequent spelling of Sacculosiphonaria .

Perellsiphon Fukuda 1994: 50, 809. Invalid; incorrect subsequent spelling of Parellsiphon .

Taxonomic remarks. The genus Siphonaria was originally described to contain four species, S. sipho , S. concinna , S. tristensis and S. exigua , without an original type species designation. Sowerby referred to two species as ‘typical species of the genus’ (“Adanson’s Mouret” (Le. Lepas mouret Adanson, 1757 ; unfigured) and “ Patella Sipho ”). However, none of these were available names. Gray (1847: 181) subsequently designated S. sipho as the type species. Anton (1838: 26) designated Siphonaria sowerbyi Michelin, 1832 and Hubendick (1946: 19) designated Siphonaria laciniosa (Linnè, 1758) as type species, but none of these taxa were originally included species rendering these subsequent type designations invalid (Art. 74.1 of the Code).

Several subsequent authors incorrectly attributed the genus name Siphonaria to Blainville, 1824, but Blainville (1824a: 268) explicitly attributed Siphonaria to Sowerby. Consequently, Blainville did not establish a junior homonym of Sowerby’s name. Hubendick (1946: 18) treated Iredale’s taxa Ellsiphon , Legosiphon , Hebesiphon , Mallorisiphon , Mestosiphon , Planesiphon (although not valid), Parellsiphon , Torquisiphon , Triellsiphon , and Pugillaria as junior synonyms of Siphonaria as these were essentially based on conchological attributes. Several authors treated Liriola as a subgenus of Siphonaria (e.g., Suter 1913: 601; Hubendick 1945: 60, 1946: 8–9, 18–19; Morrison 1963: 7; Abbott 1974: 335). By contrast, Fischer and Crosse (1900: 108) treated Liriola as a distinct genus. Contrary to Hubendick (1945: 15, fig. 1, 1946: 8, fig. 3), two determinant anatomical characters specified for the subgenus Liriola appear not to be diagnostic: The ‘epiphallus and spermoviduct’ (i.e. ED and CD) do not open separately (i.e., the genital pore is monaulic) in several species and a ‘spermatheca’ (i.e., BC) is present in the type species S. thersites as well as other species occasionally associated with this taxon ( S. obliquata , S. tasmanica , S. funiculata , S. virgulata and S. obvirgulata , S. stowae , S. lateralis , S. acmaeoides , S. capensis , S. belcheri , and S. carbo ). Indeed, all species studied herein possess a monoaulic genital opening and a BC as part of the RS. Hubendick (1946: 30) assigned Patellopsis Nobre, 1886 as a section to Liriola ; however, incorrectly stated the type species to be S. pectinata (Linnè, 1758) . However, the type species is S. algesirae Quoy & Gaimard, 1833 by original designation (Nobre, 1886: 32) although subsequently treated as a synonym of S. pectinata by Hubendick (1946: 43), Morrison (1972: 53), and Giribet & Kawauchi (2015: 5). The subsequent designation of S. lateralis Gould, 1848 as the type species for Kerguelenia Mabille & Rochebrune, 1889 by Hubendick (1945: 60) is invalid because S. lateralis was not originally included in Kerguelenia (Art. 67.2, 69.1 of the Code). Siphonaria redimiculum Reeve, 1856 and S. macgillivrayi Reeve, 1856 are the only originally included species. Powell (1946: 91) subsequently designated S. redimiculum Reeve, 1856 as the type species of Kerguelenia . However, Kerguelenia Mabille & Rochebrune, 1889 is preoccupied by Kerguelenia Stebbing, 1888 (Crustacea: Amphipoda ). Finlay (1927: 442) noted the similarity of S. obliquata with Kerguenella, but for being conchologically “quite aberrant” placed it in a new genus, Benhamina . Torquisiphon has been treated as a junior synonym of Hebesiphon by Morrison (1972: 60). Hubendick (1946: 22) mentioned Talisiphon tasmanicus as a synonym of Siphonaria tasmanica implying that he likely considered Talisiphon as a junior synonym of Siphonaria . By contrast, Morrison (1963: 7) treated Talisiphon as a section of the subgenus Liriola . Hubendick (1946: 18) treated Mouretus as a synonym of Siphonaria . The type species of Mouretus is S. adansonii Blainville, 1824 by original designation of Blainville (1824b: 162), not ‘ M. mouretus Blainville’ ( Morrison, 1972: 53; a nomen nudum). Hubendickula was ranked as section in Siphonaria s. str. by Hubendick (1955: 5–6). The name Planesiphon is unavailable from Iredale (1940: 437, 441) as no type species was designated (Art. 13.1 of the Code). However, Zilch (1959: 86), subsequently made the name available referring to Iredale’s (1940) description while designating Planesiphon elegans as the type species. He treated the taxon as a subgenus of Siphonaria . The prior type species designation of S. elegans Iredale, 1940 by McAlpine (1952: 42) is not valid because Planesiphon was not treated as an accepted name (Art. 69.1 of the Code).

Hubendick (1945, 1946, 1955) and Morrison (1963) treated Heterosiphonaria as a valid section of Siphonaria s. str. Benhamina has been ranked as a valid genus by Powell (1939: 217; 1946: 91; 1955: 120; 1979: 293, pl. 54, figs 12–13), Borland (1950: 385–393), Morton and Miller (1968: 302, 336, 338, 354, 378), and Morley (2004: 130), but as a section in Liriola by Hubendick (1945: 55; 1946: 9, 11, 18, 24–26, 64). Our results indicate that the monotypic genus Benhamina is unwarranted because its general morphological and molecular characteristics fall within the range of Siphonaria . Moreover, we synonymize Talisiphon Iredale, 1940 with Siphonaria , and its type species, S. virgulata , with S. funiculata . The type species of Simplisiphonaria Hubendick, 1945 is Siphonaria cookiana Suter, 1909 by original designation. This species is a junior objective synonym of Siphonaria australis Quoy & Gaimard, 1833 because of Boreham’s (1959) subsequent lectotype designation (a juvenile shell of S. australis from a mixed syntype lot; Jenkins, 1983: 1, 21, figs 3e–g). The type species of the following genus-group taxa are herein considered as junior synonyms of Siphonaria species: Mestosiphon Iredale, 1940 (type species M. eumelas = S. atra ); Ellsiphon Iredale, 1940 (type species E. marza = S. denticulata ); Triellsiphon Iredale, 1940 (type species Triellsiphon acervus = S. atra ); Legosiphon Iredale, 1940 (type species L. optivus = S. viridis ); Ductosiphonaria Hubendick, 1945 (type species ‘ S. bifurcata’ [non Reeve, 1856] = S. zelandica ); Planesiphon Zilch, 1959 (type species P. elegans Reeve, 1856 = S. zelandica ).

Siphonaria as delineated herein reveals considerable phylogenetic structure ( Fig. 1 ). However, none of the principal clades retrieved herein is found to exhibit a consistent morphological feature that allows us to consistently distinguish its members from those of any other clade. Therefore, we treat all previously introduced genus-group names listed above as junior synonyms of Siphonaria . Given the lack of clearly identifiable diagnostic characteristics, we do not find it useful or even justifiable to maintain subgenera within Siphonaria for any of the species examined herein. The sole exception is Siphonaria thersites (type species of Liriola ). This species is the sister group of all remaining Siphonaria species ( Figs 1–4 ) and is distinguished by the combination of a very low-level occurrence intertidally, the animal not being fully enclosed by a reduced and often flattened shell, the BD in the RS totally enclosed within the CD and a relatively small and short SPM. Should these characters be revealed as synapomorphies of a distinct taxonomic lineage that incorporates additional extralimital species, then the name Liriola was available for such a natural group. However, none of the species from the IWP examined herein is member of this potential lineage.

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 3. Maximum Likelihood phylogram (partial, species not collapsed). Clades J–L (laciniosa and plicata groups) 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 (see Tables 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 4. Maximum Likelihood phylogram (partial, species not collapsed). Clades C–F (normalis, lateralis and pectinata groups) 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 (see Tables 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.