Lyonsifusus carvalhoriosi ( Macsotay & Campos Villarroel, 2001 )

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 : 247-250

Lyonsifusus carvalhoriosi ( Macsotay & Campos Villarroel, 2001)

( Figures 9–10 )

Fusus turris: Durand, 1960: 34 , 37 ( Guyana); Altena, 1969: 43; Altena, 1975: 56; non F. turris Valenciennes, 1832 , Recent, tropical eastern Pacific.

Fusinus sp. Altena, 1969: 43 ( Suriname); Rogers, 1997: 193, fig. ( St. Vincent, Lesser Antilles).

Fusinus closter: Altena, 1969: 12 ( Suriname) ; Rios, 1970: 97, pl. 29, 2 figs. (pars; north and northeast Brazil); Altena, 1975: 56, pl. 4, figs. 3–5 ( Suriname and French Guiana); Rios, 1975: 104, 105, pl. 30, fig. 443; Okutani in Takeda & Okutani, 1983: 282, 283, figs. (pars; Suriname and Brazil); Rios, 1985: 108, pl. 37, fig. 475; Díaz & Puyana, 1994: 197 (pars; range northern South America to northern Brazil); Rios, 1994: 565, pl. 42, figs. 567; Díaz, 1995: 118 (pars; Orinoco Delta to Guiana and Suriname); Capelo & Buitrago, 1998: 132 (pars; Guayana, Venezuela); Le Loeuff and & von Cosel, 2000: 10, 11, 19, 20, 25, 27, 28, 38, 39 ( Guyane); non Fusus closter Philippi, 1850 , = Lyonsifusus . ansatus ( Gmelin, 1791) .

Fusinus nobilis: Sutty, 1986: 64 , 65, fig. 72 ( Barbados); non F. nobilis ( Reeve, 1848) , Recent, western Pacific.

Fusinus ansatus: Hadorn & Rogers, 2000: 10 –12, 37–39, pls. 3, 4, figs. 32, 39 (pars; specimens examined from Martinique (?), St. Vincent, Guyana, Suriname, French Guiana and Amapá and Pará, Brazil figured shells from Guyana and French Guiana); Rios, 2009: 250, figs. (pars; Suriname and Amapá, Brazil); Mallard & Robin, 2005: 10, pl. 14 (pars; 121 mm “Caribbean” specimen and 65.1 mm Guyane specimen figured); Massemin et al., 2009: 154, 155, figs. ( Guyane); Daccarett & Bossio, 2011: 100, 258, fig. 495 (pars; Guyana, Suriname and Brazil); Mallard & Robin, 2017: 16, fig. 2 (pars; figured shell from “N. Brazil); non Lyonsifusus ansatus ( Gmelin, 1791) .

? Fusinus dupetitthouarsi: Massemin et al., 2000: 21 ; non F. dupetitthouarsi (Kiener, 1840) , Recent, tropical eastern Pacific; nec Testud, 1974: 210, 211, 224, Salvador to Florianopolis, Brazil; nec. Rios, 1975: 105, Brazil, after Testud.

Fusinus (Fusinus) carvalhoriosi Macsotay & Campos Villarroel, 2001: 91 –94, pl. 2, figs. 6, 9, pl. 3, figs. 3, 8, pl. 7, fig. 4 (holotype 108.55 mm, paratype 182.0 mm; 17 Quadrants of Margarita Platform,. Venezuela); Snyder, 2003: 63.

Fusinus carvalhoriosi: Rios, 2009: 250 , no. 624 (in synonymy of F. ansatus ); Mallard & Robin, 2017: 27, figs. 1, 2 (type locality Banco de los Frailes, Venezuela; max. size 180 mm sl; depth range 10–120 m; figured shells from “West Indies” and “French Guyana;” range map: Venezuela to Maranhão, Brazil).

Murex ansatus: Vermeij & Snyder, 2018: 63 , 64, fig. 6 (pars; figured specimen 170 mm, Gulf of Venezuela, trawled, 50 m); non Lyonsifusus ansatus ( Gmelin, 1791) .

Lyonsifusus carvalhoriosi: Vermeij & Snyder, 2018: 66 (species included in new genus).

Types. Fusinus carvalhoriosi Macsotay & Campos Villarroel, 2001 : Holotype: 108.55 mm, Quadrant C-20, Banco de Los Frailes, Margarita Platform, Venezuela, MBUCV-XIV-4029. Paratype: 182.0 mm, Margarita Platform, presumably at MBUCV ( Macsotay & Campos Villarroel, 2001: 92, pl. 3, figs. 3, 8).

Material examined. French Guiana— 1-dd, 132.7 mm, G0 sta. 3, 05°00.9’N, 51°18.1’W, 60– 58 m, 4/16/ 1999; 1-dd, 227.2 mm, G0 sta. 10, 05°19.1’N, 51°12.7’W, 80– 79 m, 4/18/1999; 1-lv, 177.8 mm, G0 sta. 11, 05°12.8’N, 51°17.4’W, 66–70 m, 4/18/1999; 2-dd, 80.6 and 124.1 mm, G0 sta. 12, 05°01.3’N, 51°22.0’W, 60 m, 4/ 19/1999; 27-dd, 45.8–171.1 mm, G0 sta. 17, 05°06.3’N, 51°43.7’W, 70– 68 m, 4/19/1999; 2-lv, 52.8 and 54.0 mm, G0 sta. D17, 05°03.3’N, 51°10.7’W, 70 m, 4/19/1999; 1-lv, 109.1 mm, G0 sta. 18, 05°01.2’N, 51°21.7’W, 60–61 m, 4/19/1999; 1-lv, 82.6 mm, 1-dd, 102.2 mm, off Cayenne, dredged, 50 m, 1970, ANSP 465696; 2-lv, 62.1 and 142.6 mm, off Cayenne, dredged, 64 m, LC; 1-lv, 89.7 mm, off Cayenne, 55 m, LC; 1-dd, 71.3 mm, “French Guiana,” 91 m, RC. Barbados —1-lv, 108.3 mm, 1-dd, 112.0 mm, “off Barbados,” 18–27 m, LC. St. Vincent— 1- dd, 70.5 mm, dived, 15 m, RC. Colombia —1-dd, 59.5 mm, off Guajira Peninsula, RC. Venezuela —1-dd, 170.0 mm, Gulf of Venezuela, trawled, 50 m, ANSP 476305. Guyana —1-dd, 169.6 mm, off Georgetown, trawled, 1987, ANSP 465691; 1-dd, 97.1 mm, off Georgetown, RC; 3-dd, 101.9, 109.0 and 118.2 mm, off Berbice County, trawled, muddy sand, 60–80 m, 8/1987, ANSP 465692; 4-dd, 140.0, 150.0, 160.0 and 160.0 mm, off Berbice County, trawled, muddy sand, 8/1987; ANSP 465693; 3-lv, 97.7, 99.8 and 117.7 mm, off Berbice County, muddy sand, 50 m, 1988, ANSP 465695. Suriname —1-dd, 59.0 mm, “off Suriname/French Guiana,” trawl, ~ 100 m, 8/ 1960, LC. Brazil —2-dd, 84.3 and 120.2 mm, off Cabo Norte, Amapá, R/V Oregon, 110–120 m, ANSP 465694; 1- lv, 101.0 mm, 1-dd, 106.4 mm, off Oiapoque, Amapá, 100–120 m, R/V Oregon, 11/1957, LC; 2-dd, 91.4 and 103.6 mm, same data, RC; 1-dd, 80.5 mm, off Oiapoque, Amapá, R/V Oregon, 11/1957, LC; 1-dd, 61.1 mm, off Amapá, trawled, 128–146 m, R/V Oregon, LC; 1-dd, 183.1 mm, São Luis do Ribsuar, Maranhão, 50 m, RC.

Remarks. Identifying type material of Lyonsifusus carvalhoriosi is problematic. Macsotay and Campos Villarroel (2001) reported specimens from 17 quadrants of the Margarita Platform. Although they illustrated three specimens, they provided dimensions only for the holotype and a paratype and referred only the holotype to an illustration (pl. 3, figs. 3, 8), without reference to a locality or catalogue number. The caption for pl. 3, figs. 3 and 8 does not state that the specimen is the holotype but provides a locality, Quadrant C-20, Banco de Los Frailes. Information on their p. 92 indicates material from C-20 is contained in lot MBUCV-XIV-4029, where we expect that the holotype may be found. The figured holotype is identical morphologically to specimens from the Guianas and elsewhere that we report as L. carvalhoriosi .

Lyonsifusus carvalhoriosi has a moderately heavy shell with broadly swollen axial ribs crossed by relatively few raised spiral cords on all whorls; most shells lack a prominent peripheral cord bearing interrupted dashes of brown, although faint indicators of such a cord are evident on a few shells in the present material. Spiral cords are equally large on some specimens and variously large and small on others. Shell background colors range from yellowish-tan to white; areas between ribs of many shells are golden or dark brown, whereas intercostal areas of others are simply white. Collectively these features distinguish L. carvalhoriosi from the several overlapping varieties of L. ansatus . Of those varieties, L. carvalhoriosi most resembles the one named F. closter , but the F. closter variety has a lighter-weight shell with axial ribs confined to the upper spire and not continuing onto the penultimate whorl. Lack or diminishment of a peripheral cord causes lateral profiles of the spire and body whorl of L. carvalhoriosi to be rounded, whereas the lateral profiles of L. ansatus whorls tend to be peripherally carinate.

Macsotay and Campos Villarroel (2001) introduced F. carvalhoriosi for specimens from the Margarita Platform of northeastern Caribbean Venezuela. The species was reported as Fusinus closter from Venezuela and Amapá, Brazil by Rios (1970; 1975), then by that name from Venezuela, Suriname and Amapá by Rios (1985; 1994) and later as F. ansatus (pars) from Guyana and French Guiana by Hadorn and Rogers (2000) and from Venezuela, Suriname and Amapá by Rios (2009). Sutty (1986) also figured two specimens from Barbados as Fusinus nobilis ( Reeve, 1848) , a western Pacific species. We regard Antillean records of F. ansatus (and F. closter ) from Barbados and St. Vincent to represent L. carvalhoriosi . The shell from St. Vincent reported as Fusinus sp. by Rogers (1997) is L. carvalhoriosi and is included among our material examined. We consider suspect an Antillean record from Martinique ( Hadorn & Rogers 2000); no representative of the L. ansatus complex is reported at any island of the French West Indies in the comprehensive treatment of Fasciolariidae by Lamy and Pointier (2017).

A 59.5 mm specimen labeled “off Goajira Peninsula, Colombia ” in the Rogers collection and a 170 nn specimen from the Gulf of Venezuela (ANSP 476305) are the only L. carvalhoriosi we saw among several hundred L. ansatus from the northern coasts and continental shelf of Venezuela and Colombia. Based on records and specimens examined, L. carvalhoriosi ranges from Barbados southward along eastern Venezuela and the Guianas to northeastern Brazil. Excellent photos of a 92 mm specimen from off Amapá are available at the Conquiliologistas do Brasil website. That website reports that the species in Brazil occurs only off Amapá, depth range 30–150 m, size range 90–180 mm, but we report here a 227.2 mm specimen from French Guiana and a 183.1 mm specimen from off Maranhão, Brazil.

Most reports of Lyonsifusus ansatus (and F. closter ) from the Guianas and Brazil involve L. carvalhoriosi . Specimens from the Guianas figured as F. closter by Altena (1975), Okutani in Takeda and Okutani (1983) and as F. ansatus by Massemin et al. (2009) all seem to be L. carvalhoriosi , and the GREEN 0 specimens that Le Loeuff and von Cosel (1999) reported as F. closter have proved upon re-examination to be L. carvalhoriosi . Specimens of L. carvalhoriosi from the Guianas were figured by Altena (1975) and Okutani in Takeda and Okutani (1983) as F. closter and by Massemin et al. (2009) as F. ansatus . Altena (1969; 1975) also proposed that Durand’s (1960) report of Fusinus turris ( Valenciennes, 1832) from French Guiana was probably F. closter [auct., = L. carvalhoriosi ]. See Poorman (1981: figs. 1, 3) for figures of a syntype of the eastern Pacific species F. turris . Brazilian reports of F. closter by Rios (1970; 1975; 1985; 1994) are accompanied by figures of L. carvalhoriosi . Published records of F. ansatus from Brazil mention only Amapá, Pará and Maranhão, but the Conquiliologistas do Brasil website depicts the species (as F. ansatus and F. frailensis ) across northern Brazil, with records collectively from Amapá, Maranhão, Ceará, Rio Grande do Norte, Paraíba, Pernambuco and Alagoas, in depths of 10– 150 m. We believe that reported occurrences southward from Ceará require confirmation.

Lyonsifusus carvalhoriosi is not present in GUYANE collections but is well represented in G0 material, suggesting that the species may have a restricted bathymetric range off the Guianas. G0 specimens were taken only at stations in depths of 60–80 m and, as F. closter , were among species identified by Le Loeuff and von Cosel (2000: 11) as showing strong affinity for those depths. Two GUYANE stations were made in 60 m depths but the next deeper station was in 93–95 m, so preferred habitat for L. carvalhoriosi may have fallen between depths of GUYANE stations. We examined several other L. carvalhoriosi specimens from Guyana and French Guiana with recorded depths between 60 and 80 m, but some others were from as shallow as 50 m or as deep as 100 m. Altena (1975) reported several collections (as F. closter ) off Suriname in depths of 17.5–27 fm (32.0– 49.4 m), and Okutani in Takeda and Okutani (1983) reported specimens trawled off Suriname in 35– 83 m. Le Loeuff and von Cosel (2000) included F. ansatus among the principal taxa of offshore (60–80 m) sand bottom communities, whereas Massemin et al. (2009) reported that F. ansatus is trawled off Guyane occasionally in depths of 40– 200 m. All of these records are likely to have involved L. carvalhoriosi . Most Brazilian records are from greater depths (100–146 m).

After early reports (as Fusus distans ) from Suriname ( Horst & Schepman 1894; Schepman 1916), the next report from the Guianas was by Altena (1969), who reported F. closter dredged offshore of Suriname by the Coquette in 1957. Altena proposed that the Horst and Schepman shells differed by “their brownish-purple color and details of their form” from those obtained by the Coquette, surmising that the latter “probably belong to one variable species which may be F. closter .” Díaz (1995) included “Orinoco delta to Suriname ” in the range of F. closter . Hadorn and Rogers (2000) did not distinguish F. ansatus from the later-described F. carvalhoriosi , so their records from the Guianas could represent either taxon; specimens of both species from French Guiana are contained in the Rogers collection. Massemin et al. (2009) correctly figured a specimen from Guyane as F. ansatus , whereas the specimen that Mallard and Robin (2005: pl. 14) figured as F. ansatus from Guyane is L. carvalhoriosi , as is a “Caribbean” specimen figured on the same plate. The sole record of L. ansatus we have seen from Guyana is by Hadorn and Rogers (2000), who reported a 140.0 mm specimen from off Georgetown, depth 45 m, but the only specimen in the Rogers collection, labeled as Fusinus ansatus from off Georgetown, is a 97.1 mm shell that proves to be L. carvalhoriosi .

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.