Siphonaria javanica ( Lamarck, 1819 )

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 : 29-31

Siphonaria javanica ( Lamarck, 1819)

( Figs 12G–H, K, N–O , 15A )

Patella javanica Lamarck 1819: 333 (type locality: Java).— Delessert 1841: unpaginated text, pl. 23, figs 3a–c; Hanley 1858b: 152; Mermod 1950: 696; Trew 1983: 5.

Siphonaria javanica —Blainville 1827: 294.— Anton 1838: 26; H. Adams & A. Adams 1855 (in 1853–58): 271, 1863: 271; Paetel 1873: 117; 1883: 178; 1889: 428; Morrison 1972: 52; Tan & Kastoro 2004: 50; Pinchuck & Hodgson 2009: 371; White & Dayrat 2012: 59, 64, 70.

Siphonaria (Siphonaria) stellata — Hubendick 1945: 29, 31, 72, figs 42, 53; 1946: 48 (in part), pl. 3, fig. 23 (not S. stellata ( Helbling, 1779)) .

Siphonaria (Siphonaria) siquijorensis — Hubendick 1946: 47, pl. 2, fig. 38 (not S. siquijorensis Reeve, 1856 ).

Siphonaria (Siphonaria) laciniosa var. exigua — Hubendick 1946: 47 (in part), pl. 3, fig. 19 (not S. exigua Sowerby I, 1823 ).

Siphonaria laciniosa forma exigua — Hubendick 1947b: 3 (not S. exigua Sowerby I, 1823 ).

Siphonaria laciniosa forma sipho — Hubendick 1947b: 3 (not S. sipho Sowerby I, 1823 ).

Material examined. Type material. Lectotype of Patella javanica Lamarck, 1819 , present designation, from ‘les côtes de Java’ [coasts of Java, Indonesia]) ( MHNG MOLL-50923 , syntype number ‘2’, Fig. 12G ) . Paralectotypes, same data as lectotype ( MHNG MOLL-50923 , syntype number ‘1’, d, Fig. 12H ; syntype number ‘3’, d, Fig. 12I ) .

Other, non-type material. Timor-Leste: N of Dili: Dolokoan Beach , 8°31.424’S, 125°37.091’E, TL01-1 ( AM C.584795 p [M434, SK145]; Fig. 12J ) .

Taxonomic remarks. The type series for Patella javanica Lamarck, 1819 comprises three syntypes labelled 1 to 3. Syntypes #1 and #2 are siphonariids ( Figs 12G–H ); however, syntype #3 is a liotiid ( Fig. 12I ). In the original description Lamarck (1819) stated that the shell had ‘small white sides … fine and longitudinal streaks between its ribs’ while the interior was ‘blackish, bordered with yellow and a white edging’. This description closely matches syntype 2 ( Fig. 12G ), which is herein designated as the lectotype of Patella javanica for stabilisation of the name (Art. 74.1 of the Code). The figure of Patella javanica in Delessert (1841: pl. 23, fig. 3a–c), matches syntype #1 ( Fig 12H ).

Hubendick (1946: 38) treated S. javanica Lamarck, 1819 as variety of S. laciniosa . This treatment is rejected herein. Specimens figured in Hubendick (1946: 47) as S. laciniosa var. sipho (pl. 3, fig. 19, ‘Java’) and S. stellata (pl. 3, fig. 23, ‘Sunda Islands’), respectively, are of S. javanica . The ‘very minute barbs’ on the SPM mentioned by Hubendick (1946: 48) have not been observed. Such barbs on SPM have been observed only in S. japonica , S. camura sp. nov. The RS figured as S. kurracheensis siquiorensis (sic!) in Hubendick (1945: 31, fig 52; from ‘Edam Java’) matches the RS of S. alba as described herein, not S. kurracheensis nor S. javanica ( Fig. 15A ).

No type specimen of P. javanica was figured in Memod (1950). Morrison (1972: 51) in redefining selected species groups, incorrectly considered S. sipho as a synonym of S. javanica and incorrectly stated this taxon to be the type species of the genus. Further, Morrison (1972: 52) incorrectly treated S. exigua and S. sipho as synonyms of S. javanica based on similarity of the shell and a ‘common reproductive development’. These synonymies are not supported by examination of type specimens and comparative morpho-anatomy.The figured specimen in Morrison (1972: 55, fig. 1, ‘North Borneo’) matches the lectotype ( Fig. 12G ). Reversing taxonomic precedence, Rehder (1980: 97) incorrectly stated ’ Patella javanica Lamarck, 1819 ’ (= S. javanica ) to be a synonym of S. sipho . Records of ‘ S. javanica ’ in Tan & Chou (2000: fig. 116) and Tan & Woo (2010: 61) are misidentifications of S. viridis (refer Comparative remarks, S. viridis ). The record of ‘ S. javanica’ in Tan & Kastro (2004: 50) from Anambas and Natuna Islands, Indonesia is incorrect and attributable to S. sipho .

External morphology ( Figs 12N ).Foot sole centrally pale grey to dark yellow at foot edge, unpigmented; foot wall and cephalic folds dark yellow with irregular black flecks; mantle translucent dark grey to join of foot wall, yellow markings aligning with underside of ribs, black bands between on edge align under rib interstices, edge lobed and thickened to shell lip.

Shell ( Fig. 12G–I, K ; Table S9). Medium sized (max sl mean = 19.3 mm, SD = 9.7 mm, n = 2), ovate; height tall; apex offset central, apical sides convex, posterior straight to concave, protoconch area dark brown; protoconch direction central to weakly heterostrophic (n = 1), shell whorl dextral; growth striae prominent, shell thick; rib count (mean = 54, SD = 1, n = 2), ~ 11 primary ribs pale white, fairly straight, rounded ridges, increasingly raised/broaden and protrude beyond shell lip (~ 1 mm) to unevenly scallop and corrugate the edge; 3 or more interspersed finer secondary ribs pale white with irregular black/brown flecks, rib interstices white, uneven axial brown streaks; paired fused primary ribs form siphonal ridge. Interior shell margin intensely white with fine dark brown rays on edge aligning under rib interstices, siphonal groove distinct; spatula dark chocolate brown; ADM scar indistinct and paler brown with some darker rays to shell margin, CMS convex, paler than shell lip; thickening of shell lip not observed.

Reproductive system ( Fig. 15A; n = 1). Positioned to right side of coelom, against foot wall on foot muscle, under respiratory cavity; epiphallic parts positioned between RAM and BM; GA small, with singular GP through foot wall; AO large, very wide, elongated, bluntly pointed, joined to lower ED and upper GA; ED relatively long, wide, unfolded, joins to outer side of GA; GA, AO, ED all white muscular fibrous tissue; EG medium, joins end of much longer ED, soft whitish tissue, slightly folded; single, coiled end flagellum (F1), appears as an extension of wider ED, lays over back of BM; BD and CD junctions into GA close between ED join and GP, both ducts narrow, very long, smooth, 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 with large distal loop without MA, BC very small, embedded in top of AG, spherical, thin whitish translucent test; HD short, small, folded, links ducts in soft white folded tissues of AG to yellowish granulated HG; AG similar size to HG, sides match curvature of inner foot wall.

Comparative remarks. In the molecular phylogeny ( Figs 1 , 3 ), S. javanica ( laciniosa group, unit 65) is represented by a single individual only. The species is genetically well-differentiated from other species by COI distances of ≥ 27.6% (Table S6). Throughout the range of S. javanica , we found eight congeners occurring in partial sympatry. Siphonaria viridis (sympatric in northern PNG and Timor-Leste) has a smaller shell with less raised ribs and less scalloped edge, a shorter and broader ED and slightly larger BC. Siphonaria atra (sympatric in northern PNG) has a lower, darker, unpatterned shell with less raised darker and narrower ribs and a shorter ED. Siphonaria normalis has a smaller shell with less raised ribs, a less to unscalloped edge, and a shorter ED and larger BC. Siphonaria opposita (sympatric on Timor-Leste) has a lower, paler, unpatterned shell with narrower ribs, less scalloped edge, flared siphonal ridge, a shorter ED, a more pointed AO, and a bursal loop on BD. Siphonaria alba has a lower darker, unpatterned shell, a twisted ED, and a larger BC. Siphonaria forticosta sp. nov. has a lower, darker shell with less raised ribs, weaker edge scalloping, a smaller AO and ED, a larger BC, and a longer F1. Siphonaria planucosta sp. nov. has a smaller, lower, darker shell with less raised and even ribbing, a less scalloped edge, a shorter ED and AO, and a larger BC. Siphonaria campestra sp. nov. has a lower shell with less raised ribs, less scalloped edge, darker interior, and shorter ED, F1, and AO. Siphonaria javanica is conchologically resembles several species of the laciniosa group but differs anatomically (i.e., RS structure mainly size of BC; epiphallus parts of ED, F1). Hubendick (1945: fig. 53) figured the distal parts of the reproductive system of what he considered to be S. kurracheensis var. siquiorjensis from Java. However, this figure matches closely the anatomy of S. javanica ( Fig. 15A ) and therefore we consider this treatment as a misidentification. Shells from Indonesia depicted as S. javanica by Dharma (1992: pl. 17, fig. 1) closely resemble S. sipho , while those depicted by Dharma (2005: pl. 79, figs 19a–b) from East Java and Bali are correctly attributed to S. javanica . Specimens from Palawan identified as ‘ S. javanica ’ by Springsteen & Leobrera (1986: pl. 81, figs 21–22) are misidentified specimens of S. sipho . A specimen from Singapore figured by Tan & Chou (2000: fig. 116) as ‘ S. javanica ’ closely resembles S. viridis with respect to shell colouration, number and prominence of primary ribs, interior margin, size rather than S. javanica . There are currently no confirmed records of S. javanica from Singapore. A record of ‘ Siphonaria javanica ’ from Hainan ( Hasegawa et al. 2001: 28) is likely a misidentification from outside of the known distribution of this species ( Fig. 11 ). The correct identity of these specimens is unknown.

Distribution and habitat. Recorded from throughout Indonesia and Timor-Leste ( Fig. 11 ). In this study collected on exposed to moderately exposed rocky boulder shores at upper littoral levels.

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 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 11. Occurrence records of S. sipho, S. javanica and S. japonica.

FIGURE 12. Shells of S. sipho and S. javanica. A–F. S. sipho. A. Honshu, Boso Peninsula, AM C.584934 [M488]. B–C. Okinawa, AM C.584910 [M508]. D. Philippines, Cebu AM C.585120 [SK094]. E. Holotype of S. cornuta USNM5850. F. Okinawa, Tancha Bay, AM C.584925. G–H, J–K. S. javanica. G. Lectotype of P. javanica MHNG-MOLL-50923 (specimen #2). H. Paralectotype MHNG-MOLL-50923 (specimen #1). I. Paralectotype MHNG-MOLL-50917 (specimen #3, not a siphonariid). J. Timor-Leste, Dili, AM C.584795 [M434]. K. Dili, AM C.585948 [SK563]. L–O. Animals in situ. L. S. sipho, Okinawa. M. S. sipho, Okinawa [M488]. N. S. javanica, Dili, [M434]. O. S. javanica, Dili. Scale bars =10 mm.

FIGURE 15. Reproductive anatomy of S. javanica, S. obliquata, and S. australis. A. S. javanica. Timor-Leste, Dili, AM C.584795 [M434]. B–E. S. japonica. B–C. Hong Kong, ZRC.MOL.24905 [M476, SK283]. D–E. Honshu, Boso Peninsula, Neotype AM C.584938 [M489, SK308]. F–H. S. obliquata. F–G. NZ, Dunedin, Neotype NMNZ M.331450 [M515, SK421]. H. South Island, West Coast, TS, NMNZ M.331115 [M516, SK422]. I–J. S. australis I–J. NZ, Stewart Is, AM C.585247 [M480, SK284]. Unlabelled scale bars 1 mm.