Laevistrombus Abbott 1960 (Gastropoda: Strombidae): Indian and Southwest Paciﬁc Species

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Laevistrombus Abbott 1960 (Gastropoda: Strombidae): Indian and southwest Paciﬁc species

Article in Zootaxa · February 2019 DOI: 10.11646/zootaxa.4555.4.2

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The user has requested enhancement of the downloaded file. Zootaxa 4555 (4): 491–506 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2019 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4555.4.2 http://zoobank.org/urn:lsid:zoobank.org:pub:D3B7A64B-3F47-4B0D-9F4D-5F6DB028BEFE Laevistrombus Abbott 1960 (Gastropoda: Strombidae): Indian and southwest Pacific species

STEPHEN J. MAXWELL1, AART M. DEKKERS2, TASMIN L. RYMER1,3 & BRADLEY C. CONGDON1,3 1College of Science and Engineering, James Cook University, Cairns Qld 4870. E-mail: [email protected] 2Oasestraat 79, 1448 NR, Pumerend, The Netherlands. 3Centre for Tropical Environmental and Sustainability Sciences, James Cook University, Cairns Qld 4870.

Abstract

Here we evaluate the taxonomy of the marine gastropod genus Laevistrombus Abbott, 1960 and determine that there are five extant species within this genus, three of which occur in the southwest Pacific. Comparative analyses of this complex have been problematic due to the lack of designated type material. Therefore, we present the type material for L. canarium Linnaeus, 1758; L. taeniatus Quoy & Gaimard, 1834; and L. vanikorensis Quoy & Gaimard, 1834. Current taxonomy has L. vanikorensis absorbed within the L. canarium complex. L. taeniatus is generally held to be a synonym of L. turturella Röding, 1789. We demonstrate that both L. taeniatus and L. vanikorensis are distinct species and reinstate both to species level. Our revision also notes the significant variability in early teleoconch structure within the geographic range of L. vanikorensis, and highlights the need for a greater revision of Laevistrombus, given the diversity in early teleoconch morphology present in southwest Pacific species.

Key words: Laevistrombus, sympatric, taxonomy, revision, species, southwestern Pacific

Introduction

After the first major modern revision of the gastropod family Strombidae (Abbott 1960) the genus Laevistrombus Abbott, 1960 was generally accepted to contain a single extant species. However, the genus has undergone more recent revisions and it is now considered that Abbott (1960) was too conservative in his assessment (Man in’T Veld & de Turck 1998; Kreipl et al. 1999; Liverani 2013). These recent revisions currently recognise three extant taxa, namely L. canarium Linnaeus, 1758, L. guidoi Man in’T Veld & de Turck, 1998 and L. turturella Röding, 1789, although the rank of some taxa is in dispute (Liverani 2013). After further reviewing this complex, we recommend that two more Laevistrombus species, which are currently treated as synonyms post Abbott (1960), should be recognised as distinct species. The first, Laevistrombus vanikorensis Quoy & Gaimard, 1834 is currently considered a synonym of L. canarium. The second, L. taeniatus Quoy & Gaimard, 1834 is currently included under the synonymy of L. turturella (Figure 1). The reinstatement of these two species brings the total number of southwestern Pacific Laevistrombus species to three, and the total number of extant Laevistrombus species recognised worldwide to five. We note that L. guidoi and L. taeniatus are sympatric with L. vanikorensis in the southwestern Pacific region.

Methods and materials

We reviewed the taxonomy of taxa contained currently under the synonymy of L. canarium from first principles. This process involved the identification of all known names given to a targeted set of organisms from the southwest Pacific and India. This search was optimised by using historical reviews from which we could draw a preliminary list of names (Abbott 1960; Man in ‘T Veld & de Turck 1998). Once this list had been devised, it was

Accepted by T. Duda: 3 Oct. 2018; published: 15 Feb. 2019 491 chronologically ordered and the descriptions examined alongside images of types or material referenced in the original descriptions. The information obtained was then chronologically compared, from earliest to latest and the defining characteristics of preceding taxa compared with later. From this comparison a new list of valid taxa was generated based on morphological and spatiotemporal characteristics. The reference lists of all articles were examined for any literature that may have been overlooked originally. This process of type material check, followed by literary internal content and reference search, was repeated for each name on the preliminary list. In the case of taxa that were described based on reference images, such as those species named by Linnaeus (1758) or Röding (1798), these reference images were examined and critiqued to ascertain a general theme or descriptive intent. This critique was presented along with translations of the original descriptions. Finally, all southwest Pacific valid taxa were then presented with original descriptions, type images and information on type locality.

FIGURE 1. The Laevistrombus of Quoy and Gaimard (1834): A) L. canarium (Quoy and Gaimard 1834, pl. 51, fig. 10); B) Lectotype: L. vanikorensis (Quoy and Gaimard 1834, pl. 51, figs. 7 and 8); and C) Lectotype: L. taeniatus (Quoy and Gaimard 1834, pl. 51, figs. 14 and 15).

492 · Zootaxa 4555 (4) © 2019 Magnolia Press MAXWELL ET AL. Once the full species list was derived, those taxa on the list were compared to material in systematic collections to determine if the revised taxa reflected the natural diversity. This was achieved by first examining the original descriptions to determine the intent of the author in naming the organism. In this way, after examination of a larger sample of material, some taxa could be seen to reflect only regional colour forms, or the edges of a highly variable morphological cline and these were subsequently synonymised. Secondary references that did not add further information to the original descriptions, nor comprise major revisions, or contain new generic arrangements, were excluded from the synonymic list. Once the species list for the southwest Pacific was finalised and mapped, the immediate post-nuclear whorls, the early teleoconch, were examined and photographed for comparative studies to confirm species hypotheses.

Abbreviations and Acronyms:

AMD Collection of Aart M. Dekkers, Pumerend, The Netherlands. SM Collection of Stephen Maxwell, Cairns, Queensland, Australia. TM Collection of Trevor and Marguerite Young, Whitsundays, Queensland, Australia. VC Collection of Valda Cantamessa, Proserpine, Queensland, Australia. LLength.

Results

The genus Laevistrombus in the southwestern Pacific contains three species that are currently synonymised or seen as subspecies of L. canarium or L. turturella (Abbott 1960; Man in ‘T Veld & de Turck 1998; Liverani 2013). We suggest that there are distinct differences among some of the synonymized forms that allow the discernment of five species (Table 1; Figure 2). The shell of L. vanikorensis is smaller than that of L. canarium (rarely exceeding 5cm). L. vanikorensis also consistently displays a pattern of dark zigzag lines on a white background across the body of the shell and also on the posterior edge of the outer lip when viewed dorsally. This patterning terminates below the final suture. The dorsal lines are broader, less numerous, and darker than the Indian examples. L. vanikorensis has a less rotund body whorl lacking the high inflated shoulder of the dorsum. While the teleoconch of L. canarium is similar to that of some examples of L. vanikorensis, this is not a consistent feature, with the teleoconch of L. vanikorensis showing great diversity in form (Figure 3). The shell of L. taeniatus differs from L. turturella in having a uniform tan to brown colour, with a grey-blue mid-dorsal band, and an outer lip that is posteriorly more rounded. In contrast, the outer lip of L. turturella is more quadrate. While both species have spiral threads on the teleoconch, L. taeniatus does not have distinctive axial plaits that are present in L. turturella, which has a more angulate teleoconch shoulder (Figure 3). Restricted to Vanuatu, the endemic species L. guidoi differs from L. vanikorensis primarily in the teleoconch, which is more shouldered, with less defined spiral threads (Figure 3). The posterior outer lip of L. guidoi tends to be more calloused and axially recurved than L. vanikorensis. Both species show similarities in pattern when present; however, white shells are uncommon in L. vanikorensis, whereas patterning on the shell is rare in L. guidoi. Recent reviews have relegated the species L. guidoi to sub-specific rank under L. canarium (Liverani 2013). However, Liverani did not distinguish L. vanikorensis from L. canarium. While L. guidoi and L. vanikorensis are sympatric, we suggest that there are demonstrable differences in morphology that enable differentiation of these two forms. Thus L. guidoi should be regarded as full species. Examples of white shells of both L. vanikorensis and L. taeniatus are known, and these are often assigned to L. guidoi based on colour. However, colour should not be the primary determinant for assigning a specimen to a particular taxon.

Systematics

Caenogastropoda Cuvier, 1797

Stromboidea Rafinesque, 1815

Strombidae Rafinesque, 1815

Laevistrombus Abbott, 1960

Synonymy 1753 Gallinula Klein, genus XIII, p. 56, partly. Adams & Adams 1858, p. 259, partly. Chenu 1859, p. 256, partly. Mörch 1852, p. 61. Zittel 1885, p. 258. Issel & Tapparone Canefri 1876, p. 343. Non-binominal pre Linnaeus 1758. 1955 Laevistrombus Kira, p. 31. Kira 1959, p. 36. Bandel 2007, p. 153. nomen nudum. 1960 Laevistrombus Kira ex Abbott, p. 47. Type species: Strombus canarium Linnaeus, 1758.

Diagnosis. The shells of this genus are smooth, rotund and solid. The outer lip is thickened, smooth and anteriorly calloused. The columella is smooth, thickened in the middle.

FIGURE 2. The five extant members of the Laevistrombus: A) L. turturella Cebu Island, Philippines, 2015, L 70.5 mm (SM 10.009); B) L. taeniatus Boli Passage, Solomon Islands, Muddy sand 6 m, ex. R. Moylan, 1997, L 51.5 mm (SM 11.001); C) L. guidoi Espiritu Santo, Vanuatu, 50–60 ft. in Rubble, 1970, ex. B. and A. Deynzer, 54.5 mm (SM 9.001); D) L. canarium Keelamkarai, India, 5 m, 2015, L 55.0 mm (SM 8.002); and E) L. vanikorensis Alexandra Reef, Qld., On reef sand, ♀, 01/07/ 2013 (SM 7.040), colour variations.

High Absent Vanuatu Rounded Fusiform L. guidoi adapicaly Triangulate rare examples Weakly crenulatedWeakly Angularly roundedAngularly calloused which extends The length of theaperture length of The Thickened and posteriorly andposteriorly Thickened with some central some with thickening White, patterningWhite, of but slight zigzag lines may be present in be lines may zigzag Completely Completely Rounded Rounded Crenulated Triangulate L. vanikorensis Moderately high high Moderately central thickening Triangularly ovate Triangularly white shells are known known are shells white May or may not be present present be not or may May East Indian ocean to northwestern northwestern to ocean Indian East sporadically sporadically in the northern Pacific Thickened and callousedThickened posteriorly Pacific, Indonesia, Philippines, from Philippines,Pacific, Indonesia, from The length of the aperture with some length The theof aperture some with Solid zigzag dark purpleSolid tan, zigzag or to light Queensland to Wallis and Futuna Islands Futuna and Wallis Queensland to orange lines shell. on a white Low ovate known known Present Deltoid Rounded India and India and Sri Lanka Crenulated Triangulate L. canarium L. canarium below centre Inflated and Inflated round Thickened calloused Thickened colour forms known.colour forms Fine zigzag axial axial lines, Fine zigzag orange on shell. a white Completely white shells white Completely The length of theaperture length of The Patterning varies with solid with Patterningvaries ranging from dark purple dark ranging from to with some central some with thickening

High centre Present Present calloused calloused in the north Triangulate L. taeniatus sutural ramp sutural ramp Ovately fusiform fusiform Ovately central violet band Angularly roundedAngularly East Indonesia, southern southern Indonesia, East Papua New Papuato Guinea New the the length theUniform of Thickened and moderately andmoderately Thickened Brown toBrown a strong with tan some thickening the below some Axial andAxial plicate crenulated Solomon Islands and Rabaul Rabaul and Islands Solomon Rounded predistinctive with aperture, some examples with aperture, examples some

species. High Present Present Rounded calloused calloused Indonesia crenulated crenulated Triangulate L. turturella L. turturella Strongly angulateStrongly light to adapically adapically light to Laevistrombus Triangularly fusiform Triangularly Obsoletely plicate andObsoletely Eastern Indian toocean Dark brown, tanDark to white northwestern Pacific into Pacific northwestern Thickened mid to basally basally to mid Thickened Thickened and moderately andmoderately Thickened

Comparative analysis analysis of Comparative

TABLE 1. Character Colour Dorsal range Geographical CallosityColumella Shape view) (dorsal Teleoconch shape Teleoconch teleoconch Later axial whorl shape shape axial Teleoconch Early sculpture Teleoconch Early Teleoconch Early Sub-sutural cord Outer lip

INDIAN AND SOUTHWEST PACIFIC LAEVISTROMBUS Zootaxa 4555 (4) © 2019 Magnolia Press · 495 FIGURE 3. Early teleoconch studies (post-nuclear): A) Laevistrombus taeniatus Boli Passage, Solomon Islands, Muddy sand 6 m, R. Moylan, 1997 (SM 11.001); B) L. taeniatus Solomon Islands, ex. B. Swan Coll., 1985 (SM 11.002); C) L. taeniatus Hula Village, Papua New Guinea, ex. A. Hinton, 1966 (SM 11.003); D) L. turturella Cebu Island, 2015 (SM 10.009); E) L. guidoi Espiritu Santo, Vanuatu, 50–60 ft. in rubble, ex. B. and A. Deynzer, 1970 (SM 9.001); F) L. canarium Keelamlarai, India, 5 m, 2015 (SM 8.002); G) L. vanikorensis Rokui, Papua New Guinea, low tide in sand, ex G. Buick (SM 7.055); H) L. vanikorensis Rokui, Papua New Guinea, low tide in sand, ex G. Buick (SM 7.055); I) L. vanikorensis Boli Passage, Solomon Islands, muddy sand 6 m, R. Moylan, 1997 (SM 11.001); J) L. vanikorensis Rabaul, Papua New Guinea, ex. D. Martin (SM 7.053); K) L. vanikorensis Bootless Bay, Papua New Guinea, ex. A. Hinton (SM 7.054); and L) L. vanikorensis Alexandra Reef, Qld., on reef sand, ♀, 01/07/2013 (SM 7.040) (Not to scale).

Laevistrombus canarium Linnaeus, 1758 Figures 4 & 5 A–D

Synonymy 1758 Strombus (Ampliati) canariu Linnaeus, 10th edn., p. 745, no. 438. = Strombus (Ampliati) canarium Linnaeus, 1767, p. 1211, no. 507, partly. Gmelin 1791, p. 3517, no. 24, partly. = Lambis canarium Gmelin—Röding 1798, p. 65, nos. 828 and 829. = Strombus canarium Born, 1778, p. 279. Dillwyn 1817, p. 670. Gould 1833, p. 85. Quoy & Gaimard 1834, p. 73, no. 9, pl. 51, fig. 10. Anton 1839, p. 85, no. 2807. Kiener 1843, p. 33, pl. 29, fig. 1. Hanley 1856, p. 124, pl. 25, fig. 27. Martens 1889, p. 189, in part. Mohanraj et al. 2010, p. 205, pl. 1. = Gallinula canarium Linnaeus—Mörch 1852, p. 61. = Strombus (Gallinula) canarium Linnaeus—Adams & Adams 1858, p. 259.

496 · Zootaxa 4555 (4) © 2019 Magnolia Press MAXWELL ET AL. = Harpago canarium Linnaeus—Hanley 1860, p. 74. = Strombus (Strombus) canarium Linnaeus—Tryon 1885, pp. 110 and 136, pl. 2, fig. 19. = Strombus (Laevistrombus) canarium Linnaeus—Abbott 1960, p. 48. Wagner & Abbott 1978, p. 09–653. Walls 1980, pp. 75 and 76. 1777 Alata gallinula gibba Martini, p. 107, pl. 79, fig. 818. = Strombus (Gallinula) gibbus Martini—Adams & Adams 1858, p. 259. = Strombus gibbus Martini—Abbott 1960, p. 49. = Epidromis gibba Seba—Adam & Leloup 1938, p. 109. = Strombus gibbus Tapparone-Canefri—Dodge 1956, p. 275. = Strombus gibbus Issel & Tapparone-Canefri—Wagner and Abbott 1978, p. 09–654. Walls 1980, p. 189. Man in ‘T Veld & de Turck 1998, p. 95. 1777 Alata tenuis Martini, p. 168, pl. 89, fig. 876. 1791 Strombus (Ampliati) latissimus Gmelin, 13th edn., p. 3516, no. 21β, partly.

Linnaeus (1758) provided six literary illustrations, some of which support the hypothesis that the species intended was the Indian and Sri Lankan L. canarium (Figure 4). The reference to “Argenv. Conch t. 17, f. Q” is interpreted as coming from d'Argenville (1742) L’ Histoire Naturelle Eclaircie dans deux de ses Parties Principales La Lithologie et la Conchyliologie dont l’une Traite des Pierres et l’autre des Conquillages shows an illustration of L. canarium (Figure 4A). The final reference provided by Linnaeus is to “Klein. Ostr t. 4, f. 73” refers to Klein (1753) Tentamen Methodi Ostracologicae. This illustration differs from L. vanikorensis in having a blunter spire and a greatly inflated body whorl, with rounded lip and no pattern, which is more indicative of L. turturella (Figure 4B). The Petiver (1709–1711) Catalogus Classicus and Topicus, t. 98, f. 11 reference by Linnaeus can be attributed to L. canarium, determined by the overall shape, the shift in pattern on the presutural ramp, and the shortened spire (Figure 4C). The reference to “Gualt. Test. t. 32, f. N” points to an image from Gualtieri (1742) Index Testarum Conchyliorum, which shows both dorsal and ventral sides of the shell (Figure 4D). The outer lip of this specimen is not fully calloused, but the short spire and broad and inflated body whorl indicates the Gualtieri (1742) images are illustrations of L. canarium. Two images from Rumphius (1705, 1741) D’Amboinsche Rariteitkamer cited as “Rumph. Mus. t. 36, f. X” and “t. 36, f. N” deal with two different species. The plate 36 figure N is probably L. vanikorensis, given the locality of Amboina (Ambon) (Figure 4E). While plate 36 figure X illustrates Dolomena septimus Duclos, 1844 which is the only non-Laevistrombus cited by Linnaeus (1758) (Figure 4F). Type material. To stabilise the nomenclature we hereby select a syntype from the Linnaean Collection (London) {A–F 0020064 specimen 1} lot LSL. 435, as lectotype of L. canarium Linnaeus, 1758. Type locality. Abbott (1960) designated for L. canarium Ambon as the type location based on “O. Asiae” in the Linnaeus (1758) text. However, images from the Linnaean Society Collection clearly conform to the Indian examples of L. canarium (http://linnean-online.org/17229/ accessed 12/02/2017). Furthermore, the Ambon specimens at hand are Strombus taeniatus Quoy & Gaimard, 1834. Therefore, we redesignate the type locality of L. canarium to Keelamlarai, India, and base our comparisons on specimens from there. Description. “488. S. testæ labro rotundato brevi retuso, spiraque lævi” (Linnaeus 1758, p. 745). The Strombus shell lip is round, short and blunted; the spire is smooth (Translation SM). Distribution. Laevistrombus canarium is restricted to India and Sri Lanka. Records: India—Keelarkarai (AMD x 4; SM x 2; TM x2); Kottaipattinam (AMD x 2); Rameswaren (AMD x 5; SM x 4). Sri Lanka— Trimcomalee (AMD x 1).

Laevistrombus guidoi Man in’T Veld & de Turck, 1998 Figures 5 E–F

Synonymy 1998 Strombus (Laevistrombus) guidoi Man in ‘T Veld & de Turck, p. 95. Kreipl et al. 1999, pp. 11 and 26, pl. 41, figs. 1 and 2. = Laevistrombus canarium guidoi Man in ‘T Veld & de Turck—Liverani 2014, p. 24, pl. 143, figs. 1 and 3. Strombus (Laevistrombus) canarium Linnaeus—Cernohorsky 1972, p. 72, pl. 18, fig. 8a.

Type material. The Holotype is held in the Natuurhistorisch Museum Rotterdam, lot 15975.

INDIAN AND SOUTHWEST PACIFIC LAEVISTROMBUS Zootaxa 4555 (4) © 2019 Magnolia Press · 497 FIGURE 4. The Linnaeus (1758) reference images of Laevistrombus canarium: A) d'Argenville (1742) L’ Histoire Naturelle Eclaircie dans deux de ses Parties Principales la Lithology et la Conchyliologie dont l’une Traite des Pierres et l’autre des Conquillages t. 17 f. Q; B) Klein (1753) Tentamen Methodi Ostracologicae t. 4, f. 73; C) Petiver (1709–1711) Catalogus Classicus and Topicus t. 98, f. 1; D) Gualtieri (1742) Index Testarum Conchyliorum t. 32, f. N; E) Rumphius (1705, 1741) D’Amboinsche Rariteitkamer t. 36, f. N; F) Rumphius (1705,1741) D’Amboinsche Rariteitkamer. t. 36, f. X.

Type locality. The type locality for L. guidoi is Port Vila, Vanuatu (Man in’T Veld & de Turck 1998). Original description. “Shell shape: A smooth, solid shell, heavy in comparison to its size, with up to nine whorls. A slender, relatively low to high spire. The first whorls with a radial structure. Aperture wide posteriorly, narrower anteriorly. Outer lip pointed posteriorly and much thickened. Columella much thickened from its centre to its anterior; with a thin glaze on its posterior, which covers the ventral part of the body whorl. A coloured line marks the outline of this glazed area. The columella is thickened at its extreme posterior end, forming a lip that protrudes above the suture. From this lip, the margin curves concavely upward and bends back in some specimens, into the direction of the spire. Also due to the thickened outer lip, a narrow and deep posterior canal arises. The siphonal canal is fairly long, the stromboid notch weak. Sculpture: The first seven whorls with a radial structure, subsequently becoming smoother. In a few specimens with some narrow varices on the whorl. With up to 18 radial lines on the anterior portion of the body whorl. Columella and aperture smooth. Colour: Solid white, in some specimens with a pattern of obscure yellow lines. All adult specimens, without exception with a golden-yellow, metal like gloss on the columella as well as the outer lip. The inside of columella and aperture is white.

498 · Zootaxa 4555 (4) © 2019 Magnolia Press MAXWELL ET AL. Periostracum: Unknown but probably light brown judging from faint traces” (Man in’T Veld & de Turck 1998, p. 103). Distribution. Laevistrombus guidoi is restricted to Vanuatu (Figure 6). Records: Vanuatu—Vila Bay (Cernohorsky 1972); Port Vila (Man in ‘T Veld & de Turck 1998). Material examined from private collections: Vanuatu—No data (AMD x 6); Esprito Santo (SM x1); Efate (VC x 3); Malekula (TM x1). Discussion. Laevistrombus guidoi is consistently white in colour, with only rare examples showing a pattern similar to the typical L. vanikorensis. The extension of the outer lip callosity and level of “metal like gloss” is highly variable and dependent on the age of the animal’s shell: older animals have greater callosity and more gloss.

FIGURE 5. Laevistrombus canarium: A) syntype, designated from lot LSL. 435, Linnaean collection (http://linnean- online.org/17229/ {A–F 0020064 specimen 1} accessed 12/02/2017); B) Keelamkarai, India, 5 m, 2015, L 58.7 mm (TM); C) Keelamkarai, India, L 59.0 mm (AMD STR2569); and D) Rameswaran, India, L 60.9 mm (AMD STR0259); Laevistrombus guidoi: E) Efate Island, Vanuatu (VC) 59 mm; and F) Vanuatu, 50–60 ft. in Rubble, 58.0 mm (AMD STR0031).

Laevistrombus taeniatus Quoy & Gaimard, 1834 Figure 7

Synonymy 1834 Strombus taeniatus Quoy & Gaimard, p. 75, no. 10, pl. 51, figs. 14 and 15. = Strombus taeniata Quoy—Reeve 1851, spc. 51. = Strombus canarium var. taeniatus Duclos, 1844, in Chenu, pl. 7, fig. 6. = Strombus (Strombus) isabella taeniatus Horst & Schepman, 1908, p. 213. Strombus (Strombus) canarium var isabella Lamarck—Tryon 1885, pp. 110 and 136, pl. 3, fig. 21. Strombus (Gallinula) isabella Lamarck—Melvill & Standen 1899, p. 165.

INDIAN AND SOUTHWEST PACIFIC LAEVISTROMBUS Zootaxa 4555 (4) © 2019 Magnolia Press · 499 Strombus canarium Linnaeus—Hinton 1972, p. 6, pl. 3, fig. 18. Hinton 1977a, p. 13, no. 5a. Hinton 1977b, p. 10, nos. 6b and c. Strombus (Laevistrombus) turturella Röding—Man in ‘T Veld & de Turck 1998, p. 98.

Type material. We hereby select the only surviving syntype in the Muséum National d’Histoire Naturelle, Paris: Collection: Molluscs: MNHN-IM-2000-32460 as lectotype of L. taeniatus Quoy & Gaimard, 1834. Type locality. The type locality for L. taeniatus is Ambon, Indonesia. Original description. “Strombus, testa obovata, laeve, basi striata, tribus vittis violaceo-fuscis cincta; spira brevi, mucronata; labro crasso, dilatato, antice marginato” (Quoy & Gaimard 1834, p. 75). Translation: Strombus, shell ovate, smooth, with striate base, surrounded by three violet bands, spire short, mucronate, lip callused, dilated, anteriorly margined (Translation SM).

FIGURE 6. The apparent distribution (solid lines) of the three species of Laevistrombus in the southwestern Pacific based on known locations (dots) and geographical features which may affect distributions such as island chains. The type location for Laevistrombus taeniatus in the Indonesian region is also illustrated.

“This small stromb has the form and the size of Canarium. Only its back is a little more abruptly bulging, and the anterior part is deeply indented. It differs from the known species by the three transverse bands of violet brown that run over the body whorl and that are divided by a background of a pretty white. The band in the middle is largest, and the posterior one is smallest. It is the one that traverses all turns of the spire by decreasing width, so as to become not more than a reddish line at the top. These three kinds of ribbons do not reach right up to the right edge, which is thick, rounded and white, of the same (colour) as the rest of the aperture and the columella. The anterior part of the canal is striated at the bottom” (Quoy & Gaimard 1834, p. 75, Translated AMD). Distribution. Laevistrombus taeniatus has a restricted range within the southwest Pacific, ranging from southern Papua to the Solomon Islands (Figure 6). There is evidence of a distributional gap between the Indonesian type population and distribution of the rarer members of the southwest Pacific populations. At the time of the Quoy and Gaimard expedition, Ambon was a significant regional trading centre (Quoy & Gaimard 1833). Records: Australia—Murray Island (Melvill & Standen 1899). Papua New Guinea—Porebada (Hinton 1972); Hula (Hinton 1972); Manus Island, Admiralty group (Hinton 1972); Loyalty Islands (Man in‘T Veld & De Turck 1998). Material examined from private collections: Papua New Guinea—No data (VC x2); Hula Village (SM x 1). Solomon Islands—No data (SM x 1); Boli Passage (SM x1). Images Published online from dealer specimen traders: Ambon, Indonesia. Discussion. Laevistrombus taeniatus possesses a high degree of variability in colouration and is constantly paler in the western side of its range. The Indonesian examples range in colour from tan to white, with a highly variable number and colouration of banding on the body whorl. The southwest Pacific representatives of this species are consistently coloured, with a large central band on the main tanned body whorl, while the two bands alluded to in the original description vary in intensity from dark to absent.

500 · Zootaxa 4555 (4) © 2019 Magnolia Press MAXWELL ET AL. FIGURE 7. Laevistrombus taeniatus: A) syntype 1: Muséum National d’Histoire Naturelle, Paris: Collection: Molluscs: MNHN-IM-2000-32460 (https://science.mnhn.fr/institution/mnhn/collection/im/item/2000-32460?lang=fr_FR accessed 12/02/ 2017), L 40.8 mm; B) Solomon Islands, ex. A. Hinton, 1966, ex SM No. 11.003, L 52 mm; C) Papua New Guinea (VC) 58.5 mm; D) Solomon Islands, ex. B. Swan, 1985, L 60.5 mm (SM 11.002); and E) an image of shells for sale online showing the extensive colour variation in the western Ambon area of the species range.

Laevistrombus vanikorensis Quoy & Gaimard, 1834 Figure 8

Synonymy 1834, Strombus vanikorensis Quoy & Gaimard, p. 73, no. 8, pl. 51, figs. 7 to 9. = Strombus canarium var. vanikorensis Duclos, 1844, in Chenu, pl. 7, figs. 4 and 5. = Strombus vanikorensis Quoy—Reeve 1851, spc. 46. = Strombus (Strombus) canarium vanikorensis Horst & Schepman—Adam & Leloup 1938, p. 110. = Strombus vanicorensis Quoy—Tryon 1885, p. 146. = Strombus vanicorensis Tryon—Wagner & Abbott 1978, p. 09–658. Strombus canarium Linnaeus—Kiener 1843, p. 33, pl. 29, fig. 1a. Allan 1950, p. 99, pl. 17, fig. 2. Rippingale & McMichael 1961, p. 56, pl. 5, fig. 15. Wilson & Gillett 1971, p. 40, pl. 18, fig. 1. Deas 1971, unnumbered page/plate, fig. 5. Jansen 1996, p. 17, figs. 53b and c. = Strombus (Strombus) canarium Linnaeus—Tryon 1885, pp. 110 and 136, pl. 2, fig. 18. = Strombus (Gallinula) canarium Linnaeus—Chenu 1859, p. 256, figs. 1590 and 1591. Melvill & Standen 1899, p. 165.

INDIAN AND SOUTHWEST PACIFIC LAEVISTROMBUS Zootaxa 4555 (4) © 2019 Magnolia Press · 501 = Canarium (Labiostrombus) canarium Linnaeus—Solem 1958, p. 220. = Strombus (Laevistrombus) canarium Linnaeus—Abbott 1960, p. 36, pl. 17, fig. 15. Wilson & Gillett 1979, p. 73, pl. 14, fig. 1. = Laevistrombus canarium Linnaeus—Maxwell et al. 2017, p. 133. Laevistrombus canarium guidoi Man in ‘T Veld & de Turck—Liverani 2013, p. 24, pl. 143, fig. 2.

Type material. We hereby select the first syntype in the Muséum National d’Histoire Naturelle, Paris: Collection: Molluscs: MNHN-IM-2000-32459 (two syntypes), size 55.3 mm as lectotype of L. vanikorensis Quoy & Gaimard, 1834. The other syntype, with the writing “Vanikoro” in the mouth, becomes the paralectotype. Type locality. The type locality for L. vanikorensis is Vanikoro Island, Solomon Islands. Original description. “Strombus, testa obovata, laevi, alba, lineis castaneis confertissimis longitudinalibus maxime flexuosis picta; spira conic-elongata, mucronata; apertura alba; labro crasso dilatato” (Quoy & Gaimard 1834, p. 73) Strombus, shell ovate with a narrow basal end, smooth, white, with many longitudinal winding chestnut lines, spire elongated and conical, mucronate, aperture white, lip thickened and calloused (Translation SM). “On the first impression, one mistakes this species for Strombus canarium; but that species differs by its consistently larger measurements of all examples, by the longer spire that is more conical and pointed, and by the longer canal. Furthermore, its aperture is nicely white, in places yellow as with the Str. canarium, and the axial lines of which it is covered with are violet brown, which forms very sharp zigzags, and in some places are simply wavy. The part of the body whorl, close to the columella, is a little yellow. Its straight edge is dilated and thick to the perfect state, slim and membranous at a young stage, with brown crosslinks, sometimes colourless; the kind that makes the shell look dark. The periostracum is thin and yellow. The body of the animal is generally grey- greenish, stitched with brown. The filaments of the eye stalks are very long. The operculum is long, reddish and regularly toothed on the sides. The eyes are pale yellow, with a black circle exteriorly, and a brown one near the pupil” (Quoy & Gaimard 1834, p. 73, translated AMD). Distribution. Laevistrombus vanikorensis is commonly collected in Queensland from the Cape York Peninsula down to Dingo Beach (Figure 6). To the east, it ranges from Rabaul, Papua New Guinea along the island chains to the Wallis and Futuna Islands. In the northwest, it may range from Myanmar (Burma) to south Vietnam, Philippines and Indonesia, although these populations are not linked to the southwest Pacific populations. There are three historical institutional records for the Keppel Bay region. However, recent examples of white specimens sold by dealers purporting to come from Keppel Bay (VC, TM) are considered unverified. One further institutional record indicating a more southerly distribution is also considered dubious: New South Wales, Long Bay, Sydney, with the single specimen taxonomically problematic as it resembles specimens more typical of L. turturella from the northwestern Pacific. Records: Australia—Bedford Beach, Cooktown (Abbott 1960); Hayman Island (Abbott 1960); Mossman (Rippingale & McMichael 1961; Cernohorsky 1972); Murray Island, Torres Straits (Melvill & Standen 1899); Port Douglas (Abbott 1960). Solomon Islands—Florida Island (Solem 1958). New Caledonia— Touho Bay (Abbott 1960); Baie de l’Orphelinat (Abbott 1960). Material examined from private collections: Australia Alexandra Reef (SM x 134); Archer Point (SM x 1); Bowen Sand Flats (SM x 1); Cape York Peninsula (SM x 7); Cockle Bay, Magnetic Island (SM x 1); Cooya Beach (SM x 7); Dingo Beach (AMD x 1, SM x 16); Double Island Cairns (SM x 4); Four Mile Beach (AMD x1,SM x 114); Garners Beach (SM x 1); Gloucester Passage (AMD x 1); Kurrimine Beach (SM x 7); Machans Beach (SM x 2); Oak Beach (SM x 3); Pallarenda (AMD x1, SM x 6); Saunders Beach (SM x 1); Shelly Beach, Townsville (SM x 1); Wonga Beach (SM x 11); Yorkeys Knob (SM x 2); Yule Point (SM x 187). Papua New Guinea—Bootless Bay (SM x 1); Porebalda (VC x1); Rabaul (SM x1); Rokui (SM x 2). Solomon Islands—Boli Passage (SM x 1; TM x2; VC x 1); Kolambangara Island (TM x1); Marau Sound (AMD x 1). New Caledonia—No data (AMD x1); Thio (AMD x 2); Ouvéa Island (AMD x 1). Wallis and Futuna—No data (AMD x 1). Discussion. Laevistrombus vanikorensis is only associated with near-shore habitats, and is not known from the outer reef system. The benthic substrate that this species prefers is highly variable, and it has been observed living in muddy (Dingo Beach), rubbly (Wonga Beach), fine clean (Archer Point) and silty (Alexandra Reef) sand types. This species is highly gregarious, and adults appear to migrate only within a small home range. Within an adult colony, there are consistent, and significant, sexual dimorphism and sex imbalance, with females being both larger and more numerous than males (Maxwell et al. 2017). Laevistrombus vanikorensis possesses a high degree of consistency in the patterning of dark zigzag lines on a

502 · Zootaxa 4555 (4) © 2019 Magnolia Press MAXWELL ET AL. white to cream background. However, the frequency and distribution of these lines can be quite variable. In the eastern parts of the range, pale and white shells are known. Specimens from the western part of the range, particularly Queensland, have a distinctive bluish band below the suture, which is often missing in eastern specimens. A rare golden form is also known from Queensland. Limitations. The determination of the range of L. vanikorensis from literary sources without illustration is problematic; particularly with the locality records of the three sympatric species encapsulated within those of L. canarium (see Abbott 1960). Therefore, a greater emphasis on actual distribution has been given to collection material over literary records.

FIGURE 8. Laevistrombus vanikorensis: A) syntype 1: Muséum National d’Histoire Naturelle, Paris: Collection: Molluscs: MNHN-IM-2000-32459 (https://science.mnhn.fr/institution/mnhn/collection/im/item/2000-32459 accessed 12/02/2017) size 55.3 mm.; B) syntype 2: Muséum National d’Histoire Naturelle, Paris: Collection: Molluscs: MNHN-IM-2000-32459 (https:// science.mnhn.fr/institution/mnhn/collection/im/item/2000-32459 accessed 12/02/2017); C) Archer Point, Qld., Low tide on sand, ♂, 01/06/2013, L 40 mm (SM 7.017); D) Wonga Beach, Qld., Low tide on sand and rubble, ♀, 11/06/2014, L 30 mm (SM 7.032); E) Marau Sound, Diver, Solomon Islands, L 63.7 mm (AMD STR0290); F) Wallis and Fortuna Islands, L 42.0 mm (AMD STR0886); G) Kolambangara Island, Solomon Islands, 1988 (TM); and H) Boli Passage, Solomon Islands, Low tide in muddy sand, ex. R.Moylan (TM).

Acknowledgements

The authors would like to thank Valda Cantamessa, and Trevor and Marguerite Young for access to their well curated and extensive collections. We also thank those who offered invaluable information on current ranges, particularly Doug Thorn. The authors would also like to thank Valeriia Goriunova for her assistance in photographing some of the images used in the plates. We would like to thank Anne Butler for her valuable proof reading skills. Furthermore, the authors are grateful to Thomas Duda from the University of Michigan, and the two anonymous reviewers for their valuable comments that strengthened the arguments contained within the manuscript.

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