Pustulatirus ogum (Petuch, 1979)

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 : 49-53

Pustulatirus ogum (Petuch, 1979)

( Figures 52–62 )

Latirus (Polygona) ogum Petuch, 1979: 519 , 520, figs. 3A, B. Rios, 1985: 107, pl. 36, fig. 470; Costa and Moretzsohn, 1991: 59–60, figs. 4, 5; Rios, 1994: 133, pl. 42, fig. 574; Vermeij and Snyder, 2003: 18.

Latirus ogum —Petuch, 1979: 519, 520. Kaicher, 1986: card 4667; Petuch, 1986a: 9; Petuch, 1987: 140, 144, pl. 27, figs. 1, 2; Petuch, 1988: 163, pl. 39, fig. 13; Costa, 1991: 76; Lyons, 1991: 201; Matthews-Cascon et al., 1991: 1; Rios et al., 1994: 34; Goto and Poppe, 1996: 392; K. and L. Sunderland, 1996: 17, 2 figs; Snyder, 2000: 162; Petuch, 2001: 339; Vermeij and Snyder, 2003: 18, 20; Mallard and Robin, 2005: 18, pl. 47; Robin, 2008: 222, figs. 9; Landau and Vermeij, 2012: 88.

Benimakia ogum (Petuch, 1979) —Vermeij and Snyder, 2003: 15, 18-20, figs. 6a, b. Pisor and Poppe, 2008: 83; Snyder and Vermeij, 2008: 49, 50.

Pustulatirus ogum (Petuch, 1979) —Snyder, Landau and Vermeij in Landau and Vermeij, 2012: 88.

Description: Shell of medium size for genus (largest 53.1 x 17.4 mm), elongate, fusiform, with rounded, rapidly expanding whorls, broad axial ribs, and low spiral cords; spire surface of large shells (> 40 mm sl) nearly smooth except for fine spiral cords on early whorls and stronger cords on body whorl and siphonal process. Protoconch brown, of about 2 equal-sized swollen whorls; first whorl smooth with rounded sides, remaining half whorl with straight sides, smooth except for few faint axial striae and one or two axial riblets near abrupt terminus. Teleoconch of 7 to 8 whorls; spire whorls each with about 7 broad axial ribs, 6 to 9 on body whorl; suture shallow, slightly impressed, somewhat undulant in concert with axial ribs and interspaces; sutural ramp hatched with fine spiral cords crossed by low, thin axial growth increments, hatching nearly obsolete in large shells; whorls 1 and 2 with 2 smooth spiral cords crossing ribs on abapical half of whorl, several finer spiral threads on adapical half (sutural ramp); cords increasing to 3 but diminishing in strength abapically, hardly perceptible on whorls 4 to 7; posterior half of body whorl initially smooth but then showing weak re-emergance of spiral cords, cords becoming increasingly evident on final (abapical) half whorl, about 13 to 15 smooth cords at lip; cords generally increasing in strength toward anterior end of whorl, occasionally with single spiral threads between; siphonal process with 4 or 5 strong, oblique cords, usually with single lesser cords between, sometimes with 7 or 8 small, oblique cords near tip; smaller shells (~ 30–39 mm sl) with prominent large and small cords over entire surface of spire, as many as 14 smilar cords on body whorl. Aperture ovo-elongate, constricted adapically by thick parietal callus and abapically by ridge-like, sometimes bifid node opposite entrance fold at base of columella; outer lip broadly arcuate to semicircular, crenulated by extensions of interspaces between termini of spiral cords of body whorl and siphonal process, conferring serrate or saw-toothed effect, less evident on large shells; serration created by interspace adjacent to large anterior cord near anterior flexure of body whorl larger than neighboring serrata; inner surface of outer lip (of mature shells) with 9 to 13 lirae of various strengths, all (or sometimes only those on abapical side) interrupted as though composed of dashes and dots; lirae separated from lip margin by smooth band-like strip; inner lip adherent, columella arcuate posteriorly, straight anteriorly, with 3 or 4 broad plicae, occasionally with 1 or 2 lesser plicae adapically; siphonal canal typical of genus, rather short, slender, broader adapically, smooth within; outer edge crenulated in accord with termini of interspaces between exterior cords, inner edge raised, smooth, forming narrow pseudoumbilicus near tip. Shell surface uniformly dark reddish brown to light yellowish brown, usually darker on ribs; interior light orange, tan or white. Operculum corneous, dark brown, sometimes nearly black, drop-shaped, longer than wide, with terminal nucleus at pointed, inwardly curved abapical end; outer surface with many fine, arcuate, concentric growth increments. Radula unknown.

Type Material: Holotype USNM 780654 (39.3 x 17.5 mm) ( Figures 52–54 ), dd.

Type Locality: Tide pool at fringing reef around Coroa, Vermelha, Abrolhos Reef Complex, Estado Bahía, Brazil (17°57’S, 39°13’W) (Petuch, 1979: 519).

Other Material: Brazil —1, 21.1 mm, dd, north of Natal, Estado Rio Grande do Norte, depth 10–15 m, 8/ 2004, LC; 2, 33.6 and 33.3 mm, dd, off Barra, Estado Bahía, bryozoan/sand bottom, 10–15 m, diver, 1996, ANSP 449734; 1, 31.7 mm ( Figures 57–58 ), dd, off Barra, Salvador, Bahia, bryozoan/sand bottom, 10–15 m, diver, 1997, ANSP 449733; 1, 23.6 mm, lv, 170 km northeast of Alcobaça, Bahía, depth 15–20 m, diver, 1/2005, LC; 1, 30.9 mm, dd, off Alcobaça, 20–25 m, 1/2003, LC; 1, 47.7 mm, dd, off Alcobaça, 20–25 m, 1/2003, LC; 1, 32.1 mm, lv, off Alcobaça, 20-25 m, 1/2004, LC; 2, 33.8 and 32.8 mm, lv, off Alcobaça, depth 20–25 m, diver, 1/2004, LC; 1, 35.2 mm, lv, 70 km off Alcobaça, 20–25 m, ANSP 449736 ( Figure 62 ; shell figured by Vermeij and Snyder, 2003: figs. 6a, b); 1, 35.7 mm, lv, 70 km off Alcobaça, 20–25 m, diver, 1/2003, LC; 1, 21.6 mm, dd, 70 km off Alcobaça, 20–25 m, diver, 1/2004, LC; 2, 37.5 and 33.6 mm, dd, 70 km off Alcobaça, 20–25 m, diver, 7/2008, LC; 1, 28.2 mm, lv, off Conceição da Barra, Estado Espirito Santo, 60–80 m, net, 8/2006, LC; 3, 31.0, 27.8 and 22.1 mm, dd, off Conceição da Barra, trawled, 60–80 m, 8/2006, LC; 2, 28.8 and 28.1 mm, lv, off Conceição da Barra, lobster nets, 70–90 m, 2008, ANSP 449798; 4, 41.2, 34.0, 30.8 and 26.9 mm, lv, 1, 27.5 mm, dd, off Guarapari, Espirito Santo, under rocks, 15–20 m, diver, 8/1992, ANSP 449728; 3, 23.6 and 18.9 mm, dd, off Guarapari, 17–18 m, dredged, 8/2003, LC; 1, 35.8 mm, lv, off Guarapari, 17–21 m, diver, LC; 1, 42.5 mm, lv, off Guarapari, 17–21 m, diver, LC; 1, 41.0, lv, off Guarapari, under rocks, 17–21 m, diver, LC; 2, 30.9 and 29.1 mm, lv, off Guarapari, bryozoan/sand bottom, 20–25 m, diver, 12/1993; ANSP 449737; 1, 33.3 mm, lv, off Guarapari, 20–25 m, 12/1993, SC; 2, 22.0 and 18.1 mm, lv, off Guarapari, 20–25 m, diver, 12/1993, ANSP 449730; 1, 27.6 mm, dd, off Guarapari, 20–25 m, diver, 11/2003, LC; 1, 23.7 mm, lv, 3, 19.2, 18.3 and 17.2 mm, dd, off Guarapari, 20–25 m, 11/2003, LC; 1, 32.9 mm, dd, off Guarapari, 20–25 m, 12/2003, LC; 1, 17.2 mm, dd, off Guarapari, 50–60 m, net, 11/2006, LC; 1, 39.0 mm, lv, Ilha Escalvada, off Guarapari, 14–15 m, diver, LC; 1, 30.1 mm, lv, Ilha Escalvada, 15 m, diver, LC; 1, 47.5 mm, lv, Ilha Escalvada, 17 m, diver, LC; 1, 38.8 mm, lv, Ilha Escalvada, 17–21 m, diver, LC; 1, 27.4 mm, dd, Ilha Escalvada, 17–21 m, LC; 2, 37.4 and 32.4 mm, lv, Ilha Escalvada, 17–21 m, diver, LC; 1, 48.9 mm, lv, Ilha Escalvada, 25 m, diver, LC; 2, 34.0 and 30.7 mm, lv, Ilha Escalvada, 25 m, diver, LC; 1, 16.2 mm, dd, “off Ilhas,” Guarapari, 18 m; SC; 3, 13.3, 13.2 and 12.1 mm, dd, Boldro Beach, Fernando de Noronha, under rocks, 20–25 m, diver, 2000, ANSP 449731; 1, 17.3 mm, lv, Boldro Beach, Fernando de Noronha, under rocks, 20–25 m, diver, 7/2000, LC; 2, 40.3 and 38.1 mm, lv, off Arrial do Cabo, Estado Rio de Janeiro, 20–25 m, diver, 2002, ANSP 449732; 1, 50.1 mm, lv, off Arrial do Cabo, 30–35 m, diver, 1/2003, LC; 1, 53.1 mm ( Figures 55–56 ), lv, off Arraial do Cabo, under rocks, 30–35 m, ANSP 449735; 1, 51.5 mm, dd, off Arraial do Cabo, 30-35 m, diver, 1/ 2005, LC; 2, 52.8 and 49.4 mm, dd, off Arraial do Cabo, 30–35 m, diver, 1/2007, LC; 2, 31.3 and 30.4 mm, lv, off Arraial do Cabo, 30–35 m, diver, 3/2007, LC; 1, 27.8 mm, lv, off Arraial do Cabo, under rocks, 30–35 m, diver, 3/ 2007, LC; 1, 49.7 mm, dd, off Arraial do Cabo, 30–35 m, diver, 5/2011, LC; 1, 32.3 mm, dd, off Cabo Frio, Estado Rio de Janeiro, Brazil, 60–90 m, trawled, LC; 1, 22.9 mm ( Figures 59–61 ), lv, off Cabo Frio, Estado Rio de Janeiro, Brazil, depth 80–100 m, “dredged by local fishermen,” ANSP 449799.

Etymology: The species was named for “ Ogum , a Macumba god often associated with the sea” (Petuch, 1979).

Distribution: Brazil, off Natal, Estado Rio Grande do Norte (rare, herein); off Salvador, Itapoã and Arquipélago dos Abrolhos, Estado Bahía; Guarapari, Estado Espirito Santo; and Estado Rio de Janeiro (Petuch, 1979; Rios, 1985; Costa, 1991; Costa and Moretzsohn, 1991; Rios, 1994; Vermeij and Snyder, 2003). Most material we examined was from off Alcobaça, Bahía, Guarapari, Espirito Santo, and Arraial do Cabo, Rio de Janeiro. Depth range: intertidal to 20–25 m (Petuch 1979, Sunderland & Sunderland 1994); 14–80 m (herein).

Remarks:We suspected initially that shells identified here as Pustulatirus ogum (Petuch, 1979) comprised more than one species. Larger shells (> 40 mm sl) agree with the original description and illustration of Latirus ogum ; they are consistently brown, most have relatively smooth spires and body whorls, and their outer lips are only slightly serrate. Intermediate-sized shells (~ 30-39 mm sl, most with thin lips indicating immaturity) range from dark reddish brown to yellowish brown; spiral cords are well developed on all whorls and their interspaces extend prominently over the edge of the outer lip, conferring to the edge a distinctly serrate appearance. The cord that marks the abapical edge of the central area of the body whorl is larger than those around it, and an interspace adjacent to that cord usually is larger and more prominent than its neighbors. Small shells (<30 mm sl) are yellow or light brown, in contrast to the generally darker color of intermediate-sized shells, but spiral cords and serrata are similar to those of the intermediate group. However, all features of individual groups tend to blend with those of other groups, and all groups show similar geographic and bathymetric distributions. Consequently, we found no objective criterion by which to distinguish groups, and we conclude that all shells represent growth stages within a single species.

Vermeij and Snyder (2003: 19, 20) reclassified Latirus ogum in Benimakia Habe, 1958 , because “adult specimens with an intact outer lip usually have a distinct, low labral tooth at the end of an enlarged spiral cord separating the central sector of the last whorl from the concave base,” and they figured as B. ogum a 35.2 mm immature specimen from 70 km off Alcobaça (ANSP 449736, above). Landau and Vermeij (2012: 88) reconsidered the classification of Latirus ogum in Benimakia , stating: “M. A. Snyder and we now assign L. ogum to the genus Pustulatirus ” but they provided little explanation for that action. Our rationale for reclassification involves the form of the “distinct, low labral tooth” that prompted Vermeij and Snyder (2003: 20) to place the species in Benimakia ; that “tooth” is the large serration formed by the interspace between spiral cords (described above; Figure 62 ). Similar serrata occur on shells of other Pustulatirus , but this tooth-like structure is unlike that of Benimakia ( Figure 63 ), whose tooth, as in Leucozonia ( Figure 64 ), is formed by direct extension of a prominent cord, not by an interspace between cords.

Of ten species classified in Benimakia by Vermeij and Snyder (2003) and Snyder and Vermeij (2008), all except ‘ Benimakia’ ogum are Indo-West Pacific taxa. Habe (1958) described and figured the radula of B. rhodostoma (Dunker, 1860) , the type species of Benimakia , and noted its morphology to resemble that of Peristernia Mørch, 1852 , in having large and small cusps scattered across the width of lateral teeth whereas lateral cusps of Latirus Montfort, 1810 , are more equal in size or taper gradually from the inner to the outer cusp edge. Abbott (1958) described and figured a radula of L. virginensis , and Bullock (1968) figured radulae that he assigned to L. virginensis and L. attenuatus (i.e., the P. v i rg i n e n s i s species complex); those radulae more resemble those of Latirus than they do the radulae of Peristernia or Benimakia . Radulae of the Peristernia and Benimakia type have not been reported for any western Atlantic or eastern Pacific species, and it seems unlikely that such radulae will be found among species we assign to Pustulatirus .

Vermeij and Snyder (2006) reassigned four living western Atlantic species and several fossils to Pustulatirus . The living species were P. annulatus (Röding) , P. attenuatus and P. virginensis [collectively the P. virginensis species-complex] and P. eppi (‘Melvill’; = Latirus eppi auctt ., non Melvill, 1891, = P. biocellatus , herein). The larger size and often smooth surface of adult shells suggest P. ogum to be more closely related to P. virginensis than to other western Atlantic species, but P. ogum also much resembles the Panamic P. mediamericanus , the type species of Pustulatirus (compare Figures 1–3 and 55–56 ). These two species are among the largest congeners in their respective regions; P. mediamericanus attains a size of 91.0 mm sl (Pisor & Poppe 2008) and is exceeded in the tropical eastern Pacific only by P. praestantior at 99.9 mm sl (Pisor & Poppe 2008). In the tropical western Atlantic P. ogum , at 53.1 mm sl (ANSP 449735) seems to be the largest species of its genus, the next largest being P. v i rg i n e n s i s at 52.7 mm sl (ANSP 449714). As with shells of P. mediamericanus , the broad axial ribs of P. og u m assume a relatively lower profile as specimens approach maturity; the smooth surfaces of intermediate spire whorls and the initial portion of the body whorl are supplanted by spiral cords that become prominent near the terminal edge of the shell in each species.

Petuch (1979) speculated that “ Latirus sp.” of Rios (1975) from “Couves Is.,” Estado São Paulo, depth 50 m, might be a juvenile shell of Latirus ogum , but Rios et al. (1994) assigned that specimen to their new species Latirus devyanae . The immature shell that Rios (1975) figured as Latirus sp. is the same shell that Rios et al. (1994) figured as the holotype of L. (Polygona) devyanae in their fig. 3, but that specimen is not the same shell they identified as the holotype in figs. 1 and 2. We believe the latter shell, larger and mature, is the true 36.3-mm holotype of L. devyanae . The shell in fig. 3 seems to be their paratype 3, the only shell that Rios et al. (1994) cited as from Isole di Couves, São Paulo, depth 50 m. However, Rios (1975) reported dimensions of his Couves Is. shell as 33 x 15 mm whereas Rios et al. (1994) cited paratype 3 as 26.8 x 12.0 mm. Given the evident immaturity of the shell, it seems probable that the smaller reported value is correct, but the specimen should be remeasured to confirm its size.

Figures

FIGURES 38 – 68. Western Atlantic Pustulatirus and Pseudolatirus species. 38 – 42. Pustulatirus watermanorum new species. 38 – 40. Holotype, ANSP 449721, 27.4 mm, off Roatán, Honduras. 41 – 42. Paratype, ANSP 449725, 26.1 mm, off Roatán. 43 – 45. Pustulatirus eppi (Melvill, 1891) Holotype, RMNHL, 24.1 mm, Curaçao. 46 – 51. Pustulatirus biocellatus new species, Brazil. 46 – 48. Holotype, MZUSP 108767, 30.0 mm, north of Natal. 49 – 50. Paratype, ANSP 449745, 26.5 mm, off Alcobaça. 51. Paratype, ANSP 422778, 24.0 mm, off Camocim. 52 – 62. Pustulatirus ogum (Petuch, 1979), Brazil. 52 – 54. Holotype, USNM 780654, 39.3 mm, Abrolhos reef complex. 55 – 56. ANSP 449735, 53.1 mm, off Arraial do Cabo. 57 – 58. ANSP 449733, 31.8 mm, off Barra, Salvador. 59 – 61. ANSP 449799, 22.9 mm, off Cabo Frio. 62 – 64. Labral “ dentition. ” 62. Pustulatirus ogum, ANSP 449736, enlarged serratum of 35.2 mm shell, off Alcobaça, Brazil. 63. Benimakia species, ANSP 449800, labral tooth of 23.7 mm shell, Zanzibar. 64. Leucozonia nassa (Gmelin, 1791), ANSP 195297, labral tooth of 41.2 mm shell, New Providence, Bahamas. 65 – 68. Pseudolatirus tumbeziensis Olsson, 1932. 65 – 66. ANSP-IP 81622, 61.8 mm, Paraguana Peninsula Venezuela, Cantaure Formation, Miocene. 67. Same specimen, showing exterior cords and threads and edge of lip. 68. Same specimen, showing plicae in siphonal canal.

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.