Pustulatirus, Lyons And Martin Avery Snyder, William G., 2013

Lyons And Martin Avery Snyder, William G., 2013, The Genus Pustulatirus Vermeij and Snyder, 2006 (Gastropoda: Fasciolariidae: Peristerniinae) in the Western Atlantic, with Descriptions of Three New Species, Zootaxa 3636 (1), pp. 35-58 : 38-43

Pustulatirus virginensis (Abbott, 1958)

( Figures 8–30 )

[?] Lathyrus annulata [sic] (Bolt. Link)—Mørch, 1852: 99. Non Syrinx annulata Röding, 1798 .

[?] Turbinella annulata (Bolt.) —Krebs, 1864: 16. Poulsen, 1878: 11; Dall, 1885: 314; Clench et al., 1947: 35. Non Syrinx annulata Röding, 1798 .

[?] Peristernia annulata (Bolten) —Melvill, 1891b: 407. Non Syrinx annulata Röding, 1798 .

[?] Peristernia annulata (A. Ad.) —Melvill, 1891b: 411, pl. 2, fig. 1. Non Syrinx annulata Röding, 1798 .

Latirus (Polygona) virginensis Abbott, 1958: 76 , 77, text-figs. 4.7, 4.8, pl. 2, fig. 6. Rios, 1970: 96 (pars); Abbott, 1974: 227; Rios, 1975: 104 (pars); Ortiz-Corps, 1983: 121; Rios, 1985: 107, pl. 36, fig. 471 (pars); Rios, 1994: 133 (pars); Espinosa et al., 1995: 36; Harasewych, 1997: 62; Snyder, 2003: 218; Rios, 2009: 253 (pars). Non Latirus (Polygona) virginensis ‘Abbott’ auctt. northeastern Brazil, = Pustulatirus n. sp.

Latirus virginensis —Abbott, 1958: 77. Warmke and Abbott, 1961: 120, 272, pl. 22, fig. n; Arnow et al., 1963: 169; Wagner and Abbott, 1964: 167; Holeman, 1966: 29, 30; Wagner and Abbott, 1967: 247; Nowell-Usticke, 1971: pl. 6, fig.; Morris and Clench, 1973: 218, pl. 60, figs. 4; Abbott, 1974: color pl. 11, fig. 2493; Ekdale, 1974: 648; Lipka, 1974: 155; Riggs, 1975: 14; Kaicher, 1978: card 1763; Wagner and Abbott, 1978: 80–231; Woodlock, 1980: 189; Pointier et al., 1982: 9; Sarasúa and Espinosa, 1984: 8, 9, 18, fig. 6c; Sutty, 1986: 62; Pointier et al., 1987: 12; Faber, 1988: 82; Lyons, 1991: 177, 178, figs. 36-41; Espinosa et al., 1994: 113; Goto and Poppe, 1996: 393; K. and L. Sunderland, 1996: 17, 2 figs.; Pointier and Lamy, 1998: 131, 2 figs.; Snyder, 2000: 162; Redfern, 2001: 103, pl. 46, fig. 432; Snyder, 2003: 121; Henkel and Kurtz, 2004: 14, 15, fig.; Mallard and Robin, 2005: 19, pl. 52; Rios, 2009: 253 (pars); Landau and Vermeij, 2012: 88. Non Latirus virginensis ‘Abbott’ auctt., northeastern Brazil, = Pustulatirus n. sp.

Latirus [new subgenus] annulatus (Melvill) —Bullock, 1968: 65-67, pl. 4, figs. 2, 14, pl. 5, fig. 10.

Latirus [new subgenus] attenuatus (Reeve) —Bullock, 1968: 67-69, pl. 4, figs. 3–5, 10, pl. 5, figs. 11, 12, 14 (pars). Non Pustulatirus attenuatus (Reeve, 1847) ,?Caribbean region, nec Latirus eppi Melvill, 1891 , treated by Bullock as junior synonym of attenuatus .

Latirus [new subgenus] virginensis Abbott—Bullock, 1968: 71 -73, pl. 4, figs. 1, 19.

Latirus karinae Nowell-Usticke, 1969: 18 , pl. 4, fig. 821. Faber, 1988: 82; Lyons, 1991: 178; Boyko and Cordeiro, 2001: 76; Redfern, 2001: 103; Snyder, 2003: 121; Rios, 2009: 253.

Latirus virgineus [sic] (Abbott)—Santos Galindo, 1977: 222.

Latirus elegans (Gray) —Kaicher, 1986: card no. 4671 (pars; larger [right] shell only). Non Fusus elegans J. E. Gray, 1838 = Fusinus filosus (Schubert and Wagner, 1829) , West Africa, new synonymy.

Latirus species ( cf. attenuatus ) (Reeve, 1847)—Sutty, 1986: 62, 64, 65, color fig. 70. Non Turbinella attenuata Reeve, 1847 .

[?] [no genus] annulatus (Bolten) Röding—Trew, 1990: 13 .

Latirus eppi Melvill y Schapman [sic], 1891—Espinosa et al., 1994: 113. Non Latirus eppi , = Pustulatirus eppi (Melvill, 1891) , Curaçao.

Latirus (Latirus) eppi Melvill y Schapman [sic], 1891—Espinosa et al., 1995: 36. Non Latirus eppi Melvill, 1891 .

Latirus annulata [sic] (Röding, 1798)—Mallard and Robin, 2005: 16. Non Syrinx annulata Röding, 1798 .

Latirus abbotti Snyder, 2003 —Mallard and Robin, 2005: pl. 39. Robin, 2008: 219, figs. 9. Non Latirus abbotti , = Polygona abbotti (Snyder, 2003) , Caribbean Sea.

Latirus eppi Melvill, 1891 —Mallard and Robin, 2005: pl. 43 (pars; right figs. only, shell from Puerto Rico); Robin, 2008: 221, figs. Non Latirus eppi Melvill, 1891 .

Pustulatirus annulata [sic] (Röding, 1798)—Vermeij and Snyder, 2006: 421. Non Syrinx annulata Röding, 1798 .

Pustulatirus attenuata [sic] (Reeve, 1847)—Vermeij and Snyder, 2006: 421, fig. 4B. Non Pustulatirus attenuatus (Reeve, 1847) ,?Caribbean region.

Pustulatirus virginensis (Abbott, 1958) —Vermeij and Snyder, 2006: 421, fig. 4H. Rosenberg et al., 2009: 654; Zhang, 2011: 129, figs. 445(1–3); Landau and Vermeij, 2012: 88.

Pustulatirus sp. Zhang, 2011: 129, figs. 446(1–3).

Description: Shell of medium size for genus (largest 52.7 x 18.7 mm), elongate, fusiform, slender, with rounded whorls, broad axial ribs, and low spiral cords; outer lip serrate and internal lirae beaded as in generic diagnosis. Variability in shell morphology, size, and color is so extensive that we choose simply to illustrate types and several other specimens ( Figures 8–30 ) to demonstrate the range of variation. Variation is also exemplified by three shells figured as Latirus virginensis by Mallard and Robin (2005) and four shells figured as Pustulatirus sp. and P. virginensis by Zhang (2011). Further comments on variation appear in remarks for the species.

Type Material: Latirus virginensis : holotype 34.2 mm ( Figures 8–9 ), St. Thomas, Virgin Islands, ANSP 196459; 2 paratypes, 32.0 and 29.4 mm, St. Thomas, ANSP 34975; 2 paratypes, 36.6 mm, lv, 29.5 mm, dd?, St. Thomas, ANSP 34968; 4 paratypes, 29.8, 29.3, 26.6 ( Figures 10–11 ) and 25.4 mm, dd, “West Indies,” ANSP 34969; Latirus karinae : lectotype, 32.7 mm ( Figures 12–13 ), “ Puerto Rico,” AMNH 198490.

Type Localities: Latirus virginensis : St. Thomas, Virgin Islands; Latirus karinae : “ Puerto Rico.”

Other Material: Bahama Islands —2, 27.1 ( Figure 24 ) and 25.5 mm, lv, reef north of Sandy Cay, West End, Grand Bahama, 15–17 m, 1999, ANSP 449716; 2, 20.0 and 10.3 mm, dd, Sandy Cay, West End, 40 ft [12.2 m], coral rubble, LC; 1, 25.5 mm, lv, Sandy Cay, West End, LC; 2 fragments (1 shell?), dd, Gold Rock, Grand Bahama, 80 ft [24.4 m], ANSP 369654; 2, 27.9 and 25.8 mm, lv, Bimini, reef, 35 ft [10.7 m], LC; 1, 24.5 mm, lv, Bimini, reef, 45 ft [13.7 m], LC; 1, 20.7 mm ( Figures 27-28 ), dd, Bimini, 7/1985, ANSP 499719; 1, 24.2 mm, lv, Turtle Rocks, South Bimini, 30 ft [9.1 m], LC; 2, 24.6 and 18.9 mm, lv, 3.2 km south of Northwest Channel Light, 19.2 km southwest of Chub Cay, Bahamas, 70-90 ft [21.4-27.6 m], LC; 1, 26.4 mm, dd, Silver Cay Beach, Nassau, New Providence, LC; 2, 33.7 and 24.6 mm, dd, beach 1.6 km south of Fresh Creek, Andros, LC; 1, 19.0 mm, dd, off Fresh Creek, Andros, LC; 4, 31.4, 29.7, 20.5 and 20.5 mm, dd, AUTEC Base, Andros, beached by storm, LC; 1, 28.0 mm, dd, off Windimere, Eleuthera, 3 m, 6/1988, ANSP 449778. Turks and Caicos Islands —1, 23.8 mm, lv, “ Turks and Caicos,” LC. Dominican Republic —1, 30.0 mm ( Figure 25 ), lv, 1, 11.7 mm, dd, northern coast at Castillo, Duarte, rubble, 4 ft [1.2 m], 8/1994, ANSP 449715; 15, 43.3, 42.2, 40.3, 39.2, 39.0, 38.6, 35.2, 33.6, 20.8, 20.0, 19.4, 19.2, 16.3 and 14.0 mm, lv, 14, 40.3, 39.0, 37.2, 32.4, 32.2, 29.4, 27.3, 23.6, 23.1, 23.0, 21.7, 21.0, 19.8 and 19.2 mm, dd, Las Galeras, Samaná, 3 to 10 ft [0.9-3.1 m], ex Glenn Duffy, LC; 1, 41.4 mm ( Figure 14 ), lv, same data, ANSP 449754; 2, 25.1 and 23.6 mm, lv, Las Galeras, Samaná, 1-2 m, 1994, ANSP 449791; 1, 20.0 mm, lv, Meces, on dead coral at low water, LC. Puerto Rico —1, 21.3 mm, dd, Bahía Salinas, Cabo Rojo, 15 ft [4.6 m], on Thalassia , 3/2000, LC; 1, 16.2 mm, dd, same locality and date, 18 ft [5.5 m], LC; 1, 34.8 mm, dd, San Juan Harbor, 1980, ANSP 449771; 1, 24.2 mm, dd, “ Puerto Rico,” “ paratype,” ANSP 219064; 1, 27.4 mm, dd, “ Puerto Rico,” ex Warmke, UF 162196; 1, 25.7 mm, dd, “ Puerto Rico,” ex Warmke, UF 162197; 12, 33.1, 32.9, 29.7, 27.9, 27.8, 27.5, 26.7, 26.4, 26.2, 26.0, 21.2 and 20.8 mm, dd, “ Puerto Rico,” Usticke Coll. [identified as Latirus karinae by Usticke, = paralectotypes?], AMNH 191472. U. S. Virgin Islands —2, 27.4 and 19.6 mm, dd, Krause’s Reef, St. Croix, ex Usticke via McGinty, UF 135674; 39, 30.2 to 5.0 mm, dd, St. Croix at Long Reef, 2/1964 and Krause’s Reef, 3/1965, Usticke Coll., AMNH 191463; 2, 24.2 and 21.9 mm, dd, off St. Croix, 1992, ANSP 449788; 20, 29.0, 26.2, 24.6, 22.3, 22.3, 22.3, 22.2, 22.2, 22.1, 21.7, 21.2, 21.0, 21.0, 20.9, 20.8, 20.6, 20.2, 18.6, 17.4 and 16.3 mm, Cowpet Bay, St. Thomas, ex J. E. Holeman, AMNH 170501; 2, 21.6 and 9.7 mm, dd, Cowpet Bay, St. Thomas, AMNH 191476; 3, 33.7, 24.8 and 23.7 mm, dd, Cowpet Bay, St. Thomas, ANSP 449795; 1, 34.3 mm, dd, Cowpet Bay, St. Thomas, 1990, ANSP 449784; 3, 30.8, 16.9 and 16.4 mm, dd, Cowpet Bay, St. Thomas, ex J. Holeman via McGinty, UF 156255; 1, 38.6 mm, dd, Crown Bay, St. Thomas, ex Usticke via McGinty, UF 135675; 17, 40.9, 37.5, 36.6, 34.9, 33.3, 31.7, 31.1, 29.3, 27.3, 25.1, 22.9, 20.3, 19.0, 16.7, 14.0, 13.8 and 13.0, dd, Yacht Club, Jessup Bay, St. Thomas, 1969, AMNH 191470; 1, 38.4 mm, dd, Jessup Bay, St. Thomas, ex Usticke via McGinty, UF 156254; 1, 36.2 mm, dd, St. Thomas Harbor, ex J. Holeman via McGinty, UF 135673; 2, 28.0 and 26.9 mm, dd, St. Thomas Harbor, St. Thomas, ex McGinty, UF 135677; 4, 30.6, 17.5, 16.9 and 8.9 mm, dd, Water Island, St. Thomas, ex J. Holeman via McGinty, UF 363808; 1, 16.9 mm, dd, West Gregorie Channel between Water Isle and St. Thomas, ANSP 255163; 1, 35.3 mm, dd, dredged off St. Thomas, 1980, ANSP 449774; 2, 52.7 ( Figure 15 ) and 50.3 mm, dd, off St. Thomas, dredged, 1987, ANSP 449714; 1, 36.9 mm, dd, off St. Thomas, 1988, ANSP 449779; 1, 45.4 mm, dd, off St. Thomas, 1990, ANSP 449786; 3, 30.9, 24.4 and 23.2 mm, dd, off St. Thomas, 1992, ANSP 449790; 9, 40.0, 33.0, 20.8, 20.2, 16.7, 14.5, 10.8, 9.9 and 8.8 mm, dd, St. Thomas, ex Joan Brindley, 1965, AMNH 182520; 3, 47.7 ( Figure 16 ), 25.5 and 25.1 mm, dd, St. Thomas, 1976, ANSP 449713; 3, 18.0, 15.3 and 13.4 mm, dd, St. Thomas, Usticke Coll., AMNH 191490; 4, 30.1, 29.1, 25.7 and 16.2 mm, dd, St. Thomas, AMNH 114569; 1, 24.4 mm, dd, St. Thomas, 1983, ANSP 449776; 2, 23.2 and 19.0 mm, dd, St. Thomas, ANSP 449796; 4, 29.9, 29.7, 28.4 and 24.2 mm, dd, St. Thomas, LC; 1, 31.9 mm, lv, St. Thomas, LC; 1, 30.6 mm ( Figure 26 ), dd, Little St. James Island, 24 ft [7.3 m], 6/1996, ANSP 449720; 8, 24.2, 22.3, 20.9, 20.6, 20.4, 19.9, 19.9 and 16.8 mm, lv, Little St. James Island, 27 ft [8.2 m], 7/9/2000, LC; 1, 26.7 mm ( Figures 29-30 ), same data, ANSP 449718; 1, 24.3 mm, lv, same data, SC; 6, 24.7, 22.6, 22.6, 22.3, 21.7 and 21.5 mm, lv, Little St. James Island, 27 ft [8.2 m], 7/2002, LC; 5, 38.4, 37.7, 31.2, 28.4 and 27.9 mm, dd, St. John, 1978, ANSP 449748; 2, 47.0 and 45.3 mm, dd, St. John, 1980, ANSP 449775; 1, dd, 20.5 mm, “ Virgin Islands,” scuba at night in rubble, 9-10 m, 1997, ANSP 449793; 2, 31.7 and 26.9 mm, dd, “ Virgin Islands,” LC. British Virgin Islands —1, 29.4 mm ( Figure 22 ), dd, 1.6 km NE of East Point, Anegada Island, ANSP 249155; 1, 27.7 mm, Anegada, dd, deep-water fish trap, LC; 1, 29.9 mm, dd, 0.8-1.6 km SSW, SSE and E of the Bluff, Sir Francis Drake Channel, Beef Island, 12 to 14 fm [22.0- 25.6 m], ANSP 331162; 2, 10.6 and 7.5 mm, dd, Beef Island, LC; 1, 34.0 mm, dd, Beef Island, shallow reef, LC; 1, 29.1 mm, lv, Buck Island, 3 ft [0.9 m], eel grass ( Thalassia ), LC; 11, 26.7, 26.6, 24.0, 20.7, 17.7, 17.5, 17.4, 17.0, 15.5, 12.5 and 11.4 mm, dd, Fat Hog Bay, Tortola, 1–5 m, in fire coral ( Millepora ) and eel grass ( Thalassia ), 2/1985, ANSP 449777; 2, 21.9 and 19.3 mm, dd, Fat Hog Bay, Tortola, 8 ft [2.4 m], LC; 3 rolled fragments, dd, Virgin Gorda, Usticke Coll., AMNH 181489. Anguilla— 2, 48.5 and 40.6 mm, dd?, Anguilla, 1988, DLC. St. Martin —4, 40.6, 37.7, 29.9 and 22.5 mm, dd, St. Rose, DLC. St. Barthelemy —6, 31.7, 24.4, 24.3, 22.0, 20.3 and 19.1 mm, off St. Barts, Leeward Islands, 1970, ex Usticke, ANSP 421134; 1, 32.8 mm, dd, off St. Barthelemy, ANSP 449794; 3, 35.8, 31.0 and 27.9 mm, lv, 3, 28.3, 26.3 and 24.5 mm, dd, St. Barthelemy, DLC; 1, 34.2 mm, lv, “ St. Barthelemy, Guadeloupe,” Gaudiat, 1977, DLC; 1, 31.9 mm, lv?, St. Barthelemy, Pointier, 1986, DLC; 1, 33.6 mm, lv, off St. Barthelemy, trap, 100 m, ANSP 449797; 1, 32.6 mm ( Figures 19–21 ), lv, St. Barthelemy, Guadeloupe, 100 m, ANSP 449717. Barbuda —3, 25.2, 23.2 and 22.2 mm, lv, Cocoa Point, 2-5 m, LC; 2, 24.8 and 20.9 mm, lv, Barbuda, 5 ft [1.5 m], LC. Antigua —34, 40.4, 38.1, 37.2, 34.6, 34.6, 33.3, 32.2, 32.2, 31.5, 31.5, 31.5, 31.2, 30.8, 30.1, 29.9, 29.7, 29.4, 29.2, 28.3, 28.3, 26.5, 25.8, 25.6, 25.1, 24.4, 21.9, 21.1, 20.1, 19.0, 18.4, 16.5, 14.5, 12.0 and 11.9 mm, dd, Maid Island, 6/1961, Usticke Coll., AMNH 191465; 4, 34.0, 27.8, 20.6 and 14.1 mm, dd, Maid Island, ex Usticke, McGinty Coll., UF 135671; 16, 50.5 ( Figure 17 ), 47.9, 43.8, 38.0, 36.2, 35.6, 33.5, 32.7, 25.8, 25.7, 25.3, 22.4, 22.3, 19.9, 18.9 and 17.3 mm, dd, St. Johns, Usticke Coll., AMNH 191464; 10, 41.6, 33.3, 29.4, 28.5, 28.2, 25.8, 25.5, 25.3, 21.8 and 20.1, dd, St. Johns, 11/1967, Usticke Coll., AMNH 191466; 6, 51.4, 44.8, 43.9, 38.3, 35.3 and 30.3 mm, dd, St. Johns, Usticke Coll., AMNH 191467; 1, 26.3 mm, dd, Antigua, Lesser Antilles, “ paratype [of virginensis ],” ANSP 210740; 5, 42.3, 38.3, 35.1, 33.2 and 22.9 mm, dd, Antigua Island, 1979, ANSP 449768; 1, 21.2 mm, dd, Antigua Island, ex Usticke, Warmke Coll., UF 162195; 1, 27.3 mm, dd, Antigua Island, ex Usticke, UF 162198; 4, 39.2, 39.1, 35.8 and 30.4 mm, dd, off Antigua, 1993, ANSP 449792. Guadeloupe —4, 31.8, 27,2, 26.7 and 19.8 mm, dd, 2, 17.0 and 11.1 mm, lv, Deshaies, Pointier, 1983, DLC; 2, 33.7 and 24.5 mm, lv, Deshaies, 18 m, DLC; 2, 38.3 and 31.8 mm, Pointe Plate, Pointier, 1982, DLC; 1, 32.8 mm, lv, Port Louis, 10 m, DLC; 2, 34.7 ( Figure 18 ) and 30.3 mm, dd, Guadeloupe, 30-50 m, ANSP 449712. Martinique — 1, 31.8 mm, Anse Mitan, ex J. Holeman via McGinty, UF 135676. St. Lucia —2, 37.2 and 37.1 mm, dd, St. Lucia Island, ex C. W. Sheafer via McGinty, UF 135672. Netherlands Antilles— 1, 23.4 mm, dd, Malmok, Aruba, ex Fr. Fredricus, AMNH 245863; 1, 23.9 mm, dd, Malmok, Aruba, ex M. Koolman, AMNH 245864; 2, 28.2 and 25.3 mm, dd, Malmok, Aruba, “de Jong & Bijur,” ex Jerome M. Bijur Coll., AMNH 245865. No locality – 1, 38.5 mm ( Figure 23 ), dd, ex J. S. Phillips, ANSP 35033 (figured as Latirus elegans (Gray) by Kaicher, 1986a, card 4671, right fig.); 1, 24.8 mm, dd, ex Warmke, incorrectly marked as paratype, UF 162199; 1, 30.6 mm, lv, DLC; 1, 39.1 mm, dd, LC.

Distribution: This species, the best known of western Atlantic Pustulatirus , ranges throughout the eastern Caribbean region from the northernmost Bahama Islands to the Greater Antilles, the Virgin Islands, and southward to Antigua, Guadeloupe, Martinique, St. Lucia, and Aruba in the southern Netherlands Antilles; there is also an unverified record from Quintana Roo, México, in the northwestern Caribbean (Ekdale 1974). Most records with stated depths are from beaches or the shallow subtidal zone (1–30 m) but one lot from off Guadeloupe is from 30– 50 m and two from off St. Barthelemy are from 100 m.

Remarks: Morphology of individual shells may vary greatly, even among specimens from a single location. Some forms are so different morphologically that it is difficult to believe they represent the same species, but the forms invariably blend into others, rendering it difficult to determine where one ends and another begins. With the large number of specimens we examined from throughout the range, we conclude that the various forms represent a single variable species. We hope this conclusion may be tested in the future using genetic information.

The name Latirus virginensis Abbott, 1958 , has been used most often for these shells, but several other names have been associated with the group, including Syrinx annulata Röding, 1798 , Turbinella attenuata Reeve, 1847 , Peristernia annulata “(A. Adams) ” Melvill, 1891, and Latirus karinae Nowell-Usticke, 1969 . Most pieces of this taxonomic puzzle were examined by Bullock (1968), who recognized a species-group that included Latirus attenuatus (Reeve, 1847) , L. annulatus (Melvill, 1891) , and L. virginensis Abbott, 1958 , all occurring sympatrically in the West Indies. Bullock (1968) noted a “tremendous amount” of morphological variation among shells in these groups and also noted many intergrades that left relationships among the species unclear. Bullock proposed (in thesis) but did not formally introduce a new subgeneric name for the group, with Turbinella attenuata , a species we already discussed, as its type species. Bullock cited the range of what he called L. attenuatus as Cuba to the Lesser Antilles, reported records from Cuba, the Dominican Republic, Puerto Rico, and St. Thomas, U. S. Virgin Islands, and figured specimens from the Dominican Republic, Puerto Rico, and St. Thomas, but we have seen no material from any of those locations that conforms to the type of P. attenuatus . Bullock (1968: 69) also cited Latirus eppi “Melvill and Schepman ” as a junior synonym of L. attenuatus , characterizing the holotype of L. eppi as a squat specimen quite unlike the holotype of L. attenuatus but noting that “a series of intergrades appear to exist between the two forms.” He concluded that further study may distinguish them as separate species. We provide an account for Pustulatirus eppi as a separate species-level taxon, but we consider shells assigned by Bullock to L. attenuatus (except its holotype and the holotype of L. eppi ) to be more appropriately assigned to Pustulatirus virginensis .

The name Latirus annulatus (Melvill) traces to a shell figured by Chemnitz (1780: pl. 141, fig. 1316). Chemnitz (1780) grouped the figure with several others (pl. 140, figs. 1306–1309; pl. 141, 1314–1316), all of which he associated with a (non-binominal) species-group. Gmelin (1791) later assigned all of the Chemnitz figures (and some by other authors) to his new species Murex polygonus . [For several decades thereafter, some authors (e.g., Bosc 1802, Dillwyn 1817, Wood 1818, Lamarck 1822, Anton 1838) indiscriminately repeated all of the Chemnitz figures that Gmelin had cited for polygonus , but those citations are not germane here.] Röding (1798) reallocated the Chemnitz figures cited by Gmelin as follows: (1) M. polygonus Gmelin was restricted to figures 1306 and 1307 and reclassified as Fusus polygonus —now known as Latirus polygonus of the Indo-West Pacific; (2) figures 1308 and 1309 were reassigned to Fusus tapetepersicum Röding, 1798 —now a junior synonym of Latirus gibbulus (Gmelin, 1791) , another Indo-West Pacific species and the type species of Latirus ; (3) figures 1314 and 1315 were reassigned to Fusus angulatus Röding, 1798 —now Polygona angulata (Röding, 1798) of the southern Caribbean region; and (4) Chemnitz figure 1316 was reassigned both to Syrinx annulata Röding, 1798 , and to Syrinx clathrata Röding, 1798 . Link (1807) reclassified S. annulata as Cymatium annulatum but did not mention S. clathrata . Lacking other criteria by which to differentiate them, the two Röding names are primary subjective synonyms. Regardless of which name is preferred (one may argue that Link, 1807 preferred annulata ), we believe that Chemnitz figure 1316 is too poor to allow confident identification to species level, and both names should be considered nomina dubia.

Mørch (1852) first listed Lathyrus annulatus “Bolt. ” (i.e., Röding) among the West Indian fauna; H. and A.

Adams (1853) listed the name as Latirus annulatus (Bolten) ; and Krebs (1864) listed Turbinella annulata (Bolten) among West Indian shells. Finally, Peristernia annulata (Bolten) was listed by Melvill (1891b: 407) but refigured as Peristernia annulata (A. Ad.) by Melvill (1891b: 411, fig. 1). Bullock (1968: 65) contended that Chemnitz pl. 141, fig. 1316, the only figure cited by Röding for Syrinx annulata , “could certainly be a Latirus , [but] it is impossible to determine the species as the illustration is poor and the aperture is not shown. Therefore, Röding's name should be considered a nomen dubium and the [name] annulatus credited to Melvill.”

Melvill (1891b) referred authorship for the name with his figured shell not to Bolten (Röding) but to Arthur Adams. According to Trew (1992), the only species that Adams named annulatus , - a, - um were in Fissurellidae and Pyramidellidae , families unlikely to be confused with Fasciolariidae . However, if Melvill’s reference to Adams as author was in error, publication by Melvill of the unused combination Peristernia annulata and the accompanying new figure constituted valid introduction of a name. Thus, the binomen Peristernia annulata “(A. Ad.)” Melvill (1891b) seems to be available for taxonomic purposes; it only remains to prove what that name represents. The type specimen must be the shell that Melvill figured, but the location of that specimen is unknown and its identity could not be confirmed. Trew (1990) reported a specimen of annulatus in the Melvill-Tomlin Collection at the National Museum of Wales. At our request, museum staff photographed that specimen, which proves to be a young Hemipolygona carinifera (Lamarck, 1816) ( Figure 31 ) quite unlike Melvill’s figure. We hesitate to replace the well-known Latirus virginensis Abbott with the Melvill name. Absence of an original description and lack of access to a type render the status of Peristernia annulata uncertain. Until the figured type is found and its identity confirmed, we recommend that Melvill’s name be considered a species inquirenda.

Bullock (1968: 65-67) reported as Latirus annulatus (Melvill) specimens from Cuba, Jamaica, St. Thomas and Antigua, figured by that name shells from Jamaica, St. Thomas and Antigua, and cited the species’ range as Cuba to the Virgin Islands and southward in the Lesser Antilles. Bullock characterized L. annulatus as “the most commonly found and least variable of the [species group].” He distinguished L. annulatus from L. attenuatus by its “more angular shoulder on the body whorl and by having white axial ribs on every whorl,” stating that L. virginensis differs from L. annulatus “by having a more rounded body whorl lacking the white color of the axial ribs.” Shells that Bullock figured as L. annulatus generally resemble Melvill’s figure, which in turn resembles shells that Nowell-Usticke (1969) named Latirus karinae from Puerto Rico, the Virgin Islands, and Antigua. Nowell-Usticke (1971) tacitly acknowledged synonymy of L. karinae with L. virginensis by captioning as virginensis in 1971 the figure he had used for karinae in 1969; his figured shell is now the lectotype of karinae (see Boyko and Cordiero, 2001). Latirus karinae was formally relegated to synonymy with L. virginensis by Faber (1988).

Bullock (1968: 71, 72) characterized Latirus virginensis as a very variable species whose status “will not be known until large series of specimens have been studied. The relationship between this species, attenuatus , and annulatus is very unclear. The type of virginensis is a mature specimen and seems different from the largest annulatus seen.” Bullock defined the range of L. virginensis as Puerto Rico and the Virgin Islands to Brazil, citing specimens examined from Puerto Rico (four lots) and St. Thomas (one lot). Bullock also cited his pl. 4, fig. 19 for L. virginensis , but its caption identifies that shell as L. attenuatus from St. Thomas.

Abbott (1958) defined the type series of Latirus virginensis as: holotype ANSP 196459; two paratypes ANSP 34975; two paratypes ANSP 34968; and four paratypes ANSP 34969, and we examined all of these specimens at ANSP. We also examined three other specimens (ANSP 210740; ANSP 219064; UF 162199) labeled as paratypes of Latirus virginensis , but Abbott did not mention those specimens and they cannot be type material. Abbott (1958) only figured the holotype of L. virginensis , and Bullock’s thesis does not indicate that he examined the type series at ANSP, but Lyons (1991) illustrated the holotype and three paratypes from the series; those figures suggest that, had he seen them, Bullock would have classified the holotype and two paratypes (Lyons figs. 36–40) as L. attenuatus and one paratype (Lyons fig. 41) as L. annulatus , further exemplifying the confusing variability of shells in this species-group.

Simplistically, there are two dissimilar morphotypes in the group. The first type, a “ virginensis ” form, is typified by a mostly smooth shell with broad axial ribs, low to absent spiral cords, and a rather waxy shell surface. The shell color is usually olivaceous brown with lighter-colored ribs, but completely orange or yellow specimens are known, and some shells are mostly orange or yellow but with brown patches between ribs on posterior whorls. A population from Little St. James Island in the U. S. Virgin Islands is uniformly olivaceous.

The other morphotype, a “ karinae ” form, is typified by robust, swollen ribs and well-defined spiral cords. The shell color is usually orange-brown but may be yellow, dark brown, or even black, with white axial ribs in stark contrast to the background color. Axial ribs are usually more developed on the karinae form than on the virginensis form and may occur on all whorls, but more often become subdued on the penultimate and body whorls of large specimens.

Shell sizes vary considerably among and between populations. Large shells (> 35 mm sl) were examined from the Dominican Republic, St. Thomas and St. John in the U. S. Virgin Islands, Anguilla, St. Martin, St. Barthelemy, Antigua, Guadeloupe, and St. Lucia. We saw no shells larger than 28 mm from Aruba, and only two shells larger than 30 mm from the Bahama Islands.

Finally, having examined nearly 400 specimens, we could not distinguish one phenotype that could not be linked by intergrades to other quite different phenotypes. We conclude that all of the specimens represent a single, morphologically variable species for which Pustulatirus virginensis (Abbott, 1958) is the first available name.

The shell that Kaicher (1986: card 4671) figured as Latirus elegans (Gray) is in the ANSP collection (ANSP 35033; Figure 23 ), where it was catalogued as “ L. elegans A. Ad. (?)” without stated locality. That specimen, actually P. virginensis , is much darker brown than indicated by Kaicher’s figure but is identifiable by a chip in the abapical portion of the outer lip that is visible on the specimen and in the figure. It differs from the shell Kaicher figured on the same card as the holotype of Fusus elegans Gray, 1838 , which has a shorter, more canted siphonal process, relatively shorter and broader axial ribs, and distinct threads between spiral cords. We doubt that the shell Kaicher figured as holotype of Fusus elegans Gray (NHMUK 1968453) is the shell that Gray described from Sierra Leone; his description includes the phrase “with … rather distant, acute, raised, narrow, brown topped spiral ridges,” a condition that fits perfectly shells of Fusinus filosus (Schubert and Wagner, 1829) from West Africa but not evident on the NHMUK shell, and we are inclined to treat F. elegans Gray as a junior subjective synonym of Fusus filosus Schubert and Wagner.

A shell from off Fortaleza, Ceará, Brazil that Bullock (1968) reported and figured as Latirus virginensis is instead a new Brazilian endemic which we describe as Pustulatirus biocellatus n. sp. in an account that follows. The shell figured as Latirus virginensis from the Bay Islands of Honduras by K. and L. Sunderland (1996) is Pustulatirus utilaensis n. sp., described next.

Figures

FIGURES 1 – 37. New World Pustulatirus species. 1 – 4. Pustulatirus mediamericanus (Hertlein and Strong, 1951). 1 – 3. ANSP 449744, 50.0 mm, off Cebaco Island, Panamá, 36.6 m. 4. Same specimen, protoconch. 5 – 7. Turbinella attenuata Reeve, 1847. Holotype, NHMUK 196735, 31.1 mm, locality unknown. 8 – 11. Latirus virginensis Abbott, 1958. 8 - 9. Holotype, ANSP 196459, 34.2 mm, St. Thomas, U. S. Virgin Islands. 10 – 11. Paratype, ANSP 34969, 26.6 mm West Indies. 12 – 13. Latirus karinae Nowell- Usticke, 1969. Lectotype, AMNH 198490, 32.7 mm, Puerto Rico. 14 – 30. Pustulatirus virginensis. 14. ANSP 449754, 41.4 mm, Las Galeras, Samana, Dominican Republic. 15. ANSP 449714, 52.7 mm, off St. Thomas, U. S. Virgin Islands. 16. ANSP 449713, 47.7 mm, St. Thomas. 17. AMNH 191464, 50.5 mm, St. Johns, Antigua. 18. ANSP 449712, 34.7 mm, off Guadeloupe, 30 - 50 m. 19 – 21. ANSP 449717, 32.6 mm, off St. Barthelemy, 100 m. 22. ANSP 249155, 29.4 mm, off East Point, Anegada Island. 23. ANSP 35033, 38.5 mm, shell figured as Latirus elegans (Gray) by Kaicher (1986), locality unknown. 24. ANSP 449716, 27.1 mm, north of Sandy Cay, West End, Grand Bahama Island, 15 - 17 m. 25. ANSP 449715, 30.0 mm, Castillo, Duarte, Dominican Republic, 1.2 m. 26. ANSP 449720, 30.6 mm, Little St. James Island, U. S. Virgin Islands, 7.3 m. 27 – 28. ANSP 449719, 20.7 mm, Bimini, Bahamas. 29 – 30. ANSP 449718, 26.7 mm, Little St. James Island, 8.3 m. 31. Hemipolygona carinifera (Lamarck, 1816). Shell erroneously labeled “ Latirus annulatus ” in Melvill-Tomlin Collection; NMWC photo. 32 – 37. Pustulatirus utilaensis new species. 32 – 34. Holotype, ANSP 449739, 29.2 mm, off Sandy Key Reef, Utila, Honduras. 35. Paratype, ANSP 449664, 26.6 mm, southwest coast of Utila. 36 – 37. Paratype, ANSP 449659, 31.6 mm, Roatán.