Lyonsifusus ansatus ( Gmelin, 1791 )

Lyons, William G. & Snyder, Martin Avery, 2019, Fasciolariidae (Gastropoda: Neogastropoda) of French Guiana and nearby regions, with descriptions of two new species and comments on marine zoogeography of northeastern South America, Zootaxa 4585 (2), pp. 239-268 : 245-247

Lyonsifusus ansatus ( Gmelin, 1791)

( Figures 5–8 )

Murex ansatus Gmelin, 1791: 3556 ( Chemnitz, 1780: pl. 144, fig. 1340 cited for fig.); Hadorn & Rogers, 2000: 10 –12 ( Cernohorsky, 1974: 182, fig. 56 figured only available syntype ZMUC, 131.0 mm; here designated lectotype); Vermeij & Snyder, 2018: 63, 64, fig. 7 (pars; type species of Lyonsifusus ; fig’d specimen 195 mm, from El Pico, Peninsula de Paraguana, Venezuela); non M. ansatus Vermeij & Snyder, 2018: 63 , 64, fig. 6, = Lyonsifusus carvalhoriosi .

Murex versicolor Gmelin, 1791: 3556 ( Chemnitz, 1780: pl. 146, fig. 1348 cited for fig.); Hadorn & Rogers, 2000: 12 (identity uncertain, based on poor fig. by Chemnitz; could be synonym of M. ansatus ).

Syrinx maculata Röding, 1798: 122 ( Chemnitz, 1780: pl. 144, fig. 1340 cited for fig.; same fig. cited by Gmelin for M. ansatus ); Hadorn & Rogers, 2000: 10 (synonym of M. ansatus , based on same Chemnitz fig.).

Fusus torulosus Lamarck, 1816: 6 , pl. 423, fig. 4; Hadorn & Rogers, 2000: 10, 11, 37, pl. 2, figs. 26, 27 (holotype figured; synonym of M. ansatus ); Finet & Snyder, 2012: 6 , 27, fig. 10A (holotype figured).

Fusus distans Lamarck, 1822: 124 ; Horst & Schepman, 1894: 88; Schepman, 1916: 477; Altena, 1969: 12; Hadorn & Rogers, 2000: 10, 11, 37, pl. 2, figs. 28, 29 (holotype figured; junior synonym of Fusinus ansatus ); Finet & Snyder, 2012: 4 , 22, fig. 5A (holotype figured).

Fusus closter Philippi, 1847 in 1847–1850: 115, 116, pl. 5, fig. 1; Hadorn & Rogers, 2000: 10 –12, 37, pl. 2, fig. 25 (type locality Isla de Margarita Venezuela; repository of original figured specimen unknown, designated as holotype by authors, citing ICZN Article 73.1.4; junior synonym of F. ansatus ).

Fusinus closter caboblanquensis Weisbord, 1962: 364 –368, 637, 638, pl. 32, figs. 13, 14, pl. 33, figs. 1, 2 (lower Mare Formation, Venezuela, Pleistocene); Hadorn & Rogers, 2000: 11, 37, figs. 30, 31 (holotype 102.5 mm, figured, PRI 26263, lower Mare Formation; paratype 142.0 mm, Mare Formation; junior synonym of F. ansatus ).

Fusinus caboblanquensis: Weisbord, 1962: 367 , 368 (name used interchangeably as species or subspecies).

? Fusinus spec. ( Fusus distans ?): Altena, 1969: 12 ( Suriname).

Fusinus closter: Rios, 1970: 97 (pars; Venezuela); Macsotay, 1982: 185 ( Venezuela); Okutani in Takeda & Okutani, 1983: 283 (pars; Venezuela); DeJong & Coomans, 1988: 87, pl. 39, fig. 475 ( Aruba); Díaz & Puyana, 1994: 197, pl. 61, fig. 761, pl. 4, left fig. (pars; Caribbean coast of Colombia, on coarse sand bottom, depths 15–60 m; range northern South America including Colombia and Netherlands Antilles); Rios, 1994: 132 (pars; eastern Colombia and Venezuela); Díaz, 1995: 118 (pars; Colombia at Santa Marta zone and Goajira-Paraguaná zone; northern Venezuela and Aruba); Capelo & Buitrago, 1998: 132 (pars; eastern Venezuela at Isla de Cubagua, Isla de Coche, Isla de Margarita, and Peninsula de Araya Puerto La Cruz); Vermeij & Snyder, 2018: 66 (synonym of Lyonsifusus ansatus ); non F. closter , auct. Lange de Morretes, 1949: 100 (Bahia, Brazil).

Fusinus barbarensis Princz, 1973: 188 , 192, 193, 212 (nomen nudum; state of Nuevo Esparta, northern Venezuela); non Fusus barbarensis Trask, 1855 , = Barbarofusus barbarensis, Pleistocene , California, fide Callomon & Snyder (2017: 73).

Fusinus ansatus: Cernohorsky, 1974: 182 , figs. 56 (syntype 131.3 mm, figured, at University Zoological Museum, Copenhagen; earlier name for Fusus distans Lamarck ); Hadorn & Rogers, 2000: 10 –12, 37–39, pls. 2–4, figs. 25–31, 33, 34, 36–38 (pars; Colombia and Venezuela; Honduras? and Yucatan Peninsula, Mexico?); Rios, 2009: 250, figs. ( Colombia, Venezuela); Mallard & Robin, 2005: 10, pl. 14 (pars; fig. of shell with periostracum); Daccarett & Bossio, 2011: 100, 258, fig. 495 (pars; Colombia at Guajira Peninsula and Tayrona; Venezuela); Mallard & Robin, 2017: 16, 17, figs. 1, 3, 6 (pars; Colombia and Venezuela).

Fusinus barbadensis Princz, 1978: 138 , 168 (nomen nudum; Isla de Margarita, Venezuela); Princz, 1982: 123 (name attributed to Princz, 1978; error for Fusinus closter ).

Fusinus marmoratus: Sutty, 1986: 122 , fig. 136 ( Venezuela); non Fusinus marmoratus ( Philippi, 1846) , Recent, Red Sea.

Fusinus timessus: Carvajal & Capelo, 1995: 165 ( Isla de Coche, northeastern Venezuela); Macsotay & Campos Villarroel, 2001: 94 (Carvajal & Capelo record cited in synonymy for Fusinus martinezi ); non F. timessus ( Dall, 1889) , fide Macsotay & Campos Villarroel (2001: 94), = Heilprinia timessa, Recent , Florida to eastern Mexico.

Fusinus timesus (sic): Capelo & Buitrago, 1998: 132 (eastern Venezuela; Isla de Blanquilla, Isla de Coche, Isla de Margarita and Peninsula de Araya); non F. timessus ( Dall, 1889) .

? Fusinus ansatus: Hadorn & Rogers, 2000: 10 –12, 37–39, pls. 2–4, figs. 32, 35, 39 (pars; Guyana, Suriname, French Guiana and Amapá and Pará, Brazil; figured shells from Guyana and French Guiana); Mallard & Robin, 2017: 16, figs. 2, 3 (pars; figured shells from northern Brazil and Venezuela).

Fasciolaria cf. F. papillosa reevei: Macsotay & Campos Villarroel, 2001: 89 , pl. 11, figs. 1–6 (northeastern Venezuela); non F. papillosa G.B. Sowerby I, 1825 , nomen dubium; nec F. reevei Jonas in Philippi, 1850 , = Triplofusus giganteus (Kiener, 1840) , Recent, Gulf of Mexico and southeastern United States.

Fusinus (Fusinus) frailensis Macsotay & Campos Villarroel, 2001: 92 , 93, pl. 2, figs. 1, 5 (holotype 198 mm, paratype 162 mm; 7 quadrants, Margarita Platform, Venezuela).

Fusinus ( Fusinus ?) martinezi Macsotay & Campos Villarroel, 2001: 94 , pl. 4, figs. 1, 4 (holotype 123.0 mm MBUCV-XIV- 4683 figured, from quadrent E-24 & 2 other quadrants, Margarita Platform, Venezuela).

Fusinus (Fusinus) cf. F. (F.) marmoratus: Macsotay & Campos Villarroel, 2001: 94 , pl. 4, figs. 10, 11; non F. marmoratus ( Philippi, 1846) , Recent, Red Sea.

Fusinus (Fusinus) veatchi: Macsotay & Campos Villarroel, 2001: 94 , 95, pl. 4, figs. 2, 3; non F. veatchi Maury (1917) , Tertiary, Dominican Republic, fide Daccarett & Bossio (2011: 100).

Fusinus ansatus caboblanquensis: Mallard & Robin, 2017: 16 . 17, figs. 4, 5, 7 (specimens from Colombia and Venezuela figured).

Lyonsifusus ansatus: Vermeij & Snyder, 2018: 66 .

Types. Murex ansatus Gmelin, 1791 : lectotype at ZMUC, 131.0 mm, designated by Hadorn and Rogers (2000: 10– 12), previously figured as syntype by Cernohorsky (1974: 182, fig. 56); Murex versicolor Gmelin, 1791 : location of type unknown, based on Chemnitz (1780: pl. 146, fig. 1348); Fusus torulosus Lamarck, 1816 : holotype MHNG INVE 51710, 145 mm, locality on label: Mer Rouge ( Finet & Snyder 2012: 6, 27, fig. 10A); Fusus distans Lamarck, 1822 : holotype MHNG INVE 51709, 108 mm, locality on label: Océan Indien ( Finet & Snyder 2012: 3, 22, fig. 5A); Fusus closter Philippi, 1847 : locality not stated, figured specimen recognized as holotype by Hadorn and Rogers (2000: 10), repository unknown; Fusus closter caboblanquensis Weisbord, 1962 : holotype PRI 26263, 102.5 mm, Lower Mare Formation, paratype PRI 26264, 142.0 mm, Upper Mare Formation, Pleistocene, Venezuela ( Weisbord 1962: 366, pl. 32, figs. 13, 14, pl. 33, figs. 1, 2); Fusinus frailensis Macsotay & Campos Villarroel, 2001 : holotype MBUCV-XIV-4142, 196.0 mm, Quadrant A-21, paratype MBUCV-4038, Quadrant C- 20, Margarita Platform, Venezuela ( Macsotay & Campos Villarroel, 2001: 93, pl. 2, figs. 1, 5); Fusinus martinezi Macsotay & Campos Villarroel, 2001 : holotype 123.0 mm, MBUCV-XIV-4683, 4631 or 4852, Quadrants E-24, C- 19 or D-21, Margarita Platform ( Macsotay & Campos Villarroel, 2001: 94, pl. 4, figs. 1, 4).

Material examined. Northern coast of Colombia and Venezuela —80-lv/dd, 16.1–225.3 mm, 0–120 m, LC. Suriname— 1-lv, 152.4 mm, “off Suriname,” trawled, 80 m, LC. French Guiana —3-lv, 121.5, 126.0 and 164.4 mm, off French Guiana , trawled, 50–80 m, 1984, ANSP 465682 ; 1-dd, 58.7 mm, French Guiana , 91 m, RC . Brazil —1-dd, 95.5 mm, off Amapá, R/ V Oregon, 1957, ANSP 465681 ; 1-dd, 163.8 mm, off Amapá, 100–120 m, R/ V Oregon, RC .

Remarks. Variations in shell morphology and color have contributed to an extensive synonymy for L. ansatus . Early authors divided the complex into two species: one with dark-colored, strongly carinate, rough shells and one with light-colored, relatively smoother shells with only a single large cord providing peripheral carination. The pairs Murex ansatus Gmelin, 1791 – M. versicolor Gmelin, 1791 and Fusus torulosus Lamarck, 1816 – F. distans Lamarck, 1822 each exemplify that dichotomy. Murex ansatus and Fusus torulosus soon disappeared from use, perhaps because the rough form was seldom encountered. Lack of accurate locality data also complicated identification. It was long believed that F. distans came from the Philippines, so when Fusus closter Philippi, 1850 was described from Venezuela, that name was adopted for the Caribbean species. Fusinus closter caboblanquensis Weisbord, 1962 , introduced for a Venezuelan Pleistocene fossil, was soon adopted for the rugose living form as well (e.g. Petuch 1987; Mallard & Robin 2017). Princz (1973) reported a species as F. barbarensis (non Fusus barbarensis Trask, 1855 ) from Isla de Margarita and nearby islands off the Caribbean coast of Venezuela but later ( Princz 1978) listed the Venezuela species as F. barbadensis , a nomen nudum that he still later (1982) cited as a junior synonym of Fusinus closter .

Type specimens of Murex ansatus , Fusus torulosus , F. distans , F. closter , F. closter caboblanquensis , F. frailensis and F. martinezi are well illustrated and, except for F. closter , are available in various repositories. A type specimen for Murex versicolor has not been located; Hadorn and Rogers (2000: 12) declared that name to be “definitely not recognizable after the stylized drawings [of Chemnitz],” but noted that it could be conspecific with the non-carinate form of Fusinus ansatus ; those authors followed Reeve (1847), Tryon (1881) and others in treating the name as a nomen dubium. We concur that the Chemnitz figure closely resembles the non-carinate L. ansatus , so we retain the name provisionally as a synonym of L. ansatus . The location of a type specimen for Fusus closter is also unknown (see Coan & Kabat 2017 for discussion of problems locating Philippi’s types), but the original figure clearly depicts the non-carinate form of L. ansatus and its locality, “ Insula Margarita in India Occidentali,” confirms its identity.

Hadorn and Rogers (2000) recognized considerable intraspecific variability within the group and settled on one name, Fusinus ansatus , for the complex. Macsotay and Campos Villarroel (2001: 91–95, pls. 2–5, 7, 11), apparently unaware of work by Hadorn and Rogers, retained F. closter at species level (with F. caboblanquensis as a junior synonym), introduced three new species-level names ( F. carvalhoriosi , F. frailensis and F. martinezi ) and additionally applied the names F. marmoratus ( Philippi, 1846) , F. veatchi ( Maury, 1917) and Fasciolaria cf. papillosa reevei (Jonas in Philippi, 1850) , all for specimens of the L. ansatus species-complex living essentially sympatrically on the Margarita Platform off northeastern Venezuela. Of these names, Fasciolaria papillosa reevei was introduced for a smooth form of Triplofusus giganteus (Kiener, 1840) (Fasciolariinae) of the southeastern United States and eastern Mexico ( Snyder et al. 2012); Fusinus marmoratus is a poorly understood name sometimes applied to a species that occurs off central and southern Brazil but which also has been associated with F. verrucosus ( Gmelin, 1791) of the northern Red Sea. Fusinus veatchi is a fossil species from the Pliocene Gurabo Formation of the Dominican Republic, and we concur with Daccarett and Bossio (2011: 100) that the name is inappropriate for Recent Venezuelan specimens. Features of all of the Venezuelan “taxa” seem to overlap morphologically, leading us to conclude that most taxa distinguished by Macsotay and Campos Villarroel should be subsumed as synonyms of Lyonsifusus ansatus , and we combine them for this discussion. The only name by Macsotay and Campos Villarroel that we retain as probably valid for a species-level taxon is F. carvalhoriosi ; see the species account of Lyonsifusus carvalhoriosi for rationale. Perhaps future genetics studies will provide better understanding of morphological variability within this group.

Records of L. ansatus and L. carvalhoriosi published without illustrations or diagnoses are difficult to assign and, because the taxa may be incorrectly identified, some locality and depth records remain uncertain. Lyonsifusus ansatus has been reported from Martinique, Barbados, Aruba, Colombia, and Venezuela to northern Brazil ( Hadorn & Rogers 2000; Daccarett & Bossio 2011). The species seems abundant in northeastern Venezuela ( Capelo & Buitrago 1998; Macsotay & Campos Villarroel 2001), where it occurs from the intertidal zone seaward to about 200 m; and it is common off Colombia ( Daccarett & Bossio 2011), and figures of F. closter from Aruba ( De Jong & Coomans 1988: pl. 39, fig. 475) are of L. ansatus , but we can only report with confidence its occurrence in those nations. Most figures in reports from the Lesser Antilles, the Guianas and Brazil, as well as most specimens we examined, represent L. carvalhoriosi (see that species account for those records), but this does not preclude the possibility of L. ansatus in some of those areas. A shell that Hadorn and Rogers (2000: 38, fig. 35) cited as from Barbados seems to be L. ansatus . Díaz (1995) and Capelo and Buitrago (1998) mentioned F. closter from off the Orinoco delta in Guayana; we have not seen specimens from that area, but it is relatively near the confirmed eastern range of L. ansatus . Other reported localities are patently incorrect: Hadorn and Rogers (2000: 11) rightfully questioned specimens labeled “ Honduras ” and “ Yucatán Peninsula, Mexico ”; there is no credible evidence that L. ansatus occurs north of continental Colombia in the western Caribbean region.

We examined a few specimens trawled off Suriname, French Guiana and Brazil that may be variants of L. ansatus . The shells are white and covered with raised spiral cords, including a larger peripheral cord ( Figs. 5–6 ), sometimes with faint brown dots ( Figs. 7–8 ), but the axial ribs are low, faint, and confined to early whorls of the spire, unlike the prominent, broad axial ribs that characterize typical L. carvalhoriosi . Similar shells are illustrated by Mallard and Robin (2017: 16, figs. 2, 3), the first from northern Brazil and the other from Venezuela, and another similar shell from Brazil is figured as Fusinus frailensis on the Conquiliologistas do Brasil website. Such specimens are uncommon among material from the Guianas.

Figures

FIGURES 5–16. Guianas Fusininae. 5–8. Lyonsifusus ansatus (Gmelin, 1791). 5–6. 163.8 mm, off Amapá, Brazil, R/V Oregon, depth 100–120 m, LC. 7–8. 162.4 mm, Suriname, depth 80 m, LC. 9–10. Lyonsifusus carvalhoriosi Macsotay & Campos Villarroel, 2001, 142.6 mm, off Cayenne, French Guiana, depth 64 m, LC. 11–12. Aristofusus benjamini Hadorn, 1997, 171.6 mm, CP4379, 06°33’N, 52°26’W, depth 130–131 m, MNHN IM-2013-56484. 13–14. Fusinus chocolatus (Okutani, 1983), 45.9 mm, CP4377, 06°36’N, 52°29’W, depth 157–162 m, MNHN IM-2014-6076. 15–16. Fusinus dilectus (A. Adams, 1856), 77.0 mm, off Paramaraibo, Suriname, R/V Oregon, RC.