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Early stone tools are claimed to be the earliest evidence for the cultural transmission of toolmaking techniques, and with it, cumulative culture. This claim has ostensibly been supported by experimental studies wherein modern humans learned stone tool production (knapping) in conditions that provided opportunities for cultural transmission. However, alternative hypotheses propose that individual learning was sufficient for the expression of early knapping techniques. In order to evaluate this possibility, the capacities of individuals to independently re-innovate early knapping techniques need to be determined. For this, individuals must be tested in cultural isolation, i.e., in a test condition in which knapping techniques cannot be culturally transmitted via demonstrations or reverse engineering. Here, we report on the results of this test condition with human participants (N = 28). Naïve individuals spontaneously re-innovated various early knapping techniques, resulting in products resembling the earliest core and flake technologies. These results contradict previous hypotheses and conclusions of earlier experiments that explicitly implicated cultural transmission in Oldowan stone tool production. They suggest instead that knapping techniques among pre-modern hominins could have been individually derived rather than necessitating cultural transmission.

Plants deploy a variety of chemical and physical defenses to protect themselves against herbivores and pathogens. Organic farming seeks to enhance these responses by improving soil quality, ultimately altering bottom up regulation of plant defenses. While laboratory studies suggest this approach is effective, it remains unclear whether organic agriculture encourages more-active plant defenses under real-world conditions. Working on the farms of cooperating growers, we examined gene expression in the leaves of two potato (Solanum tuberosum) varieties, grown on organic vs. conventional farms. For one variety, Norkotah, we found significantly heightened initiation of genes associated with plant-defense pathways in plants grown in organic vs. conventional fields. Organic Norkotah fields exhibited lower levels of nitrate in soil and of nitrogen in plant foliage, alongside differences in communities of soil bacteria, suggesting possible links between soil management and observed differences in plant defenses. Additionally, numbers of predatory and phloem-feeding insects were higher in organic than conventional fields. A second potato variety, Alturas, which is generally grown using fewer inputs and in poorer-quality soils, exhibited lower overall herbivore and predator numbers, few differences in soil ecology, and no differences in gene-activity in organic and conventional farming systems. Altogether, our results suggest that organic farming has the potential to increase plants' resistance to herbivores, possibly facilitating reduced need for insecticide applications. These benefits appear to be mediated by plant variety and/or farming context.

Datasets associated with EAP19-0828 manuscript

Supplemental figures showing variation in the number of species and resources separately, and their influence on invasion probability and interaction strengths

R code for algorithm to define a vector of carrying capacities to allow coexistence of native species.

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): 35-58, DOI: 10.5281/zenodo.283572

Pustulatirus watermanorum new species (Figures 38���42) Description: Shell small for genus (largest 32.2 x 10.5 mm), fusiform, with rounded whorls, prominent axial ribs and obscure spiral cords, surface generally appearing smooth, waxy. Protoconch of 2 whorls, first whorl broadly globose, nearly flat on top, with rounded sides; second whorl no wider than first, with straight sides and two or three faint axial riblets just before terminus. Teleoconch of 6 to 7 rapidly expanding whorls, each with 8 or 9 broad, evenly rounded axial ribs that extend from suture to suture; about 7 spiral cords on first whorl, including 3 faint ones on sutural ramp; cords increasing in number abapically, penultimate whorl with 4 cords on ramp and 8 more crossing axial ribs, sometimes with single lesser threads between cords crossing ribs; central area of body whorl nearly smooth, but with about 13 very low cords, followed by 2 stronger cords adjacent to junction with base; dorsal surface of siphonal process with 9 or 10 oblique cords of varying strength, sometimes with single faint threads between. Aperture ovate, constricted adapically by parietal node at anal sinus and abapically by tooth-like node opposite entrance fold at junction of columella and siphonal canal; outer lip arcuate, edge crenulated by extensions of interspaces between spiral cords on body whorl, inner surface with about 10 beaded lirae, some incomplete; inner lip adherent, columella with 3 or 4 oblique plicae near base, sometimes with smaller plica adapically; siphonal canal typical for genus, straight, outer edge crenulated, inner edge simple, straight, slightly raised, sometimes forming shallow pseudoumbilicus near tip. Outer surface of shell uniformly orange, interior white. Operculum corneous, brown, drop-shaped, with terminal nucleus at pointed anterior end, outer surface with many concentric, arcuate growth increments; prominent posterior muscle scar occupying about 70 % of inner surface, bounded by thick, raised ridge. Radula unknown. Type Material: Holotype 27.4 x 10.5 mm (Figures 38���40), lv, off Roat��n, Honduras, depth 80 m, in baited shell trap, ANSP 449721. Paratypes: Honduras ��� 1, 27.7 mm, dd, same data as holotype, LC; 2, 26.4 and 23.6 mm, dd, off Roat��n, 30���40 m, in lobster traps, 1996, ANSP 449722; 3, 26.6 and 24.8 mm, lv and 26.7 mm, dd, off Roat��n, 200���300 ft (60.7���91.5 m), ANSP 449723; 20, 29.1, 27.0, 25.1, 20.6, 15.6 and 9.5 mm, lv and 27.6, 27.4, 25.6, 25.5, 24.4, 24.2, 22.1, 19.2, 17.9, 17.6, 15.4, 15.2, 13.5 and 12.1 mm, dd, off Roat��n, 200���300 ft (60.7���91.5 m), shell traps, SC; 2, 25.4 and 25.3 mm, lv, off Roat��n, 80 m, traps, 1999, ANSP 449724; 2, 25.7 mm, lv and 25.0 mm, dd, off Roat��n, 100 m, UF 455469; 1, 25.1 mm, lv, off Roat��n, 480 ft (146.4 m), BMSM 17942; 2, 26.1 (Figures 41���42) and 24.6 mm, lv, off Roat��n, 160 m, traps set on undersea slope, 1995, ANSP 449725; 2, 28.2 and 25.9 mm, dd, Cabo Faisa, Mosquitia, diver, 10-15 m, 1996; ANSP 449726; 1, 29.2 mm, lv, Gorda Bank, 30-35 m, coral reef, on sand, MNHN 25669; 1, 27.7 mm, dd, Gorda Bank, 30-45 m, near coral reef, USNM 1192972; 2, 32.2 and 30.0 mm, lv, Rosalind Bank, 90 ft (27.4 m), from lobster boat, SC. Colombian Islands ��� 4, 27.9, 27.4, 27.1 and 24.5 mm (last with chipped apex), dd, Banco Quitasue��o, 15���18 m, under coral slabs, diver, 2004, ANSP 449727; 1, 27.3 mm, dd, Banco Quitasue��o, 15���18 m, under coral slab, NHMUK 20120250; 3, 25.9, 25.1 and 20.9 mm, dd, Banco Quitasue��o, 15���18 m, LC; 1, 26.2 mm, dd, Banco Quitasue��o, 15-18 mm, LC; 1, 16.6 mm, San Andres Island, rubble, 15 m, diver, 2007, ANSP 422779. Other Material: Honduras ��� 11 specimens, 21.1 to 16.6 mm, all immature with thin lips, dd, off Roat��n, 300 ft (91.2 m), in baited shell traps, LC; 4 specimens, 27.6 to 12.0 mm, all with severe break repairs, etc., dd, off Roat��n, 200���300 ft (60.7���91.2 m), shell traps, SC. Type Locality:Off Roat��n, Honduras, depth 80 m. Etymology: The species name honors the James Waterman family, whose interest and acts of kindness encouraged the Sunderlands in assembling their important collection of western Atlantic gastropods. Distribution:Continental shelf of Honduras, including Gorda Bank and Rosalind Bank, and at Banco Quitasueno and Cayos San Andres, Colombia, western Caribbean Sea; depth range: 10��� 160 m. Remarks: The uniformly bright orange color is helpful in distinguishing P. watermanorum from the partially sympatric P. utilaensis. Pustulatirus watermanorum also differs from P. utilaensis by its proportionally shorter spire and longer body whorl and siphon. Axial ribs of P. watermanorum extend from suture to suture whereas those of P. utilaensis (and P. attenuatus) are separated from the suture adapically by a narrow ramp. Sutures of P. utilaensis are more impressed, whorls are more convex, and spiral cords are stronger. The two largest specimens (32.2 and 30.0 mm sl) constitute a single paratype lot (SC) from Rosalind Bank. Features of the smaller shell are consistent with those of other P. watermanorum, but the larger shell is attenuated, with a relatively taller and more slender spire than its conspecifics. The range of depths we report for this species (10���160 m) is the greatest known for any species of Pustulatirus. We offer no explanation for this but note that all specimens were obtained from commercial dealers, who in turn obtained some of them from fishermen. More reliable locational and bathymetric data may reveal a narrower depth range., Published as part of 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, pp. 35-58 in Zootaxa 3636 (1) on pages 45-46, DOI: 10.5281/zenodo.283572

Pustulatirus ogum (Petuch, 1979) (Figures 52���62) Latirus (Polygona) ogum Petuch, 1979: 519, 520, figs. 3 A, 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, 1986 a: 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. 6 a, 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 Bah��a, 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 Bah��a, 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 Alcoba��a, Bah��a, depth 15���20 m, diver, 1 / 2005, LC; 1, 30.9 mm, dd, off Alcoba��a, 20���25 m, 1 / 2003, LC; 1, 47.7 mm, dd, off Alcoba��a, 20���25 m, 1 / 2003, LC; 1, 32.1 mm, lv, off Alcoba��a, 20-25 m, 1 / 2004, LC; 2, 33.8 and 32.8 mm, lv, off Alcoba��a, depth 20���25 m, diver, 1 / 2004, LC; 1, 35.2 mm, lv, 70 km off Alcoba��a, 20���25 m, ANSP 449736 (Figure 62; shell figured by Vermeij and Snyder, 2003: figs. 6 a, b); 1, 35.7 mm, lv, 70 km off Alcoba��a, 20���25 m, diver, 1 / 2003, LC; 1, 21.6 mm, dd, 70 km off Alcoba��a, 20���25 m, diver, 1 / 2004, LC; 2, 37.5 and 33.6 mm, dd, 70 km off Alcoba��a, 20���25 m, diver, 7 / 2008, LC; 1, 28.2 mm, lv, off Concei����o da Barra, Estado Espirito Santo, 60���80 m, net, 8 / 2006, LC; 3, 31.0, 27.8 and 22.1 mm, dd, off Concei����o da Barra, trawled, 60���80 m, 8 / 2006, LC; 2, 28.8 and 28.1 mm, lv, off Concei����o 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, Itapo�� and Arquip��lago dos Abrolhos, Estado Bah��a; 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 Alcoba��a, Bah��a, 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 (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 Alcoba��a (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 M��rch, 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 (R��ding), 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 S��o 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, S��o 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., Published as part of 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, pp. 35-58 in Zootaxa 3636 (1) on pages 49-53, DOI: 10.5281/zenodo.283572

Pustulatirus biocellatus new species (Figures 46���51) Latirus virginensis Abbott, 1958 ���Bullock, 1968: 72, 73, 99, 101, pl. 4, fig. 1. Matthews, 1968: 248. Non Latirus virginensis Abbott, 1958, Recent, eastern Caribbean. Latirus (Polygona) virginensis Abbott, 1958 ���Rios, 1970: 96 (pars). Rios, 1975: 104 (pars), pl. 29, fig. 440; Rios, 1985: 107 (pars), pl. 36, fig. 471; Rios, 1994: 133 (pars), pl. 42, fig. 575; Rios, 2009: 253 (pars), figs. Non Latirus (Polygona) virginensis Abbott, 1958, Recent, eastern Caribbean. Latirus eppi Melvill, 1891 ���Mallard and Robin, 2005: pl. 43, figs. (pars; 17 -mm shell from Brazil only). Non Latirus eppi Melvill, 1891, Recent, Cura��ao. Pustulatirus eppi (Melvill, 1891) ���Vermeij and Snyder, 2006: 421, fig. 4 C, pars. Non Pustulatirus eppi (Melvill, 1891), Recent, Cura��ao. Description: Shell small for genus (largest 30.0 x 13.8 mm), solid, broadly fusiform. Protoconch of about 2 rounded whorls; first whorl smooth, second with 2 or 3 broad axial riblets on last quarter whorl; riblets increasing in strength toward junction with teleoconch. Teleoconch of as many as 7 whorls bearing prominent broad, wellrounded axial ribs crossed by low, smooth spiral cords; cords diminishing in strength abapically, causing shell surface to appear smooth; whorls 1 and 2 each with 7 or 8 ribs, subsequent whorls each with 6 or 7 ribs; about 3 evenly spaced cords on whorls 1 and 2, joined by 2 or 3 more broad, low cords on sutural ramp of whorl 3, number of cords increasing to about 14 on body whorl of largest shell; most body whorl cords very low and indistinct, presence of some indicated only by color changes atop ribs; 3 to 5 stronger, oblique cords atop siphonal process, sometimes with 1 to 3 faint spiral threads between. Aperture ovo-elongate, constricted adapically by callosity on parietal shield and abapically by prominent node at junction with siphonal canal; outer lip arcuate, serrate on mature shells, particularly on abapical edge in response to termini of extensions between spiral cords of body whorl and outer edge of siphon, inner side with 7 to 11 (usually 8 or 9) well-spaced, strongly beaded lirae, most evident on fully developed lip; inner lip and parietal shield adherent; columella straight, with 4 distinct plicae adapical to entrance fold of short, straight siphonal canal. Shell exterior reddish brown, with creamy white node-like ribs, light-colored ribs of body whorl crossed by narrow brown band, producing two distinctive spots on each rib; some mature shells uncommonly with ribs of body whorl tan, not white, rendering ���biocellate��� effect less evident; interior of shell white except for brown band at edge of outer lip. Operculum corneous, narrow, with anterior terminal nucleus, outer surface covered with many indistinct, concentric, arcuate growth increments. Radula unknown. Type Material: Northeastern Brazil ��� Holotype 30.0 x 13.8 mm (Figures 46���48), dd, off Natal, Estado Rio Grande do Norte, Brazil, depth 10���15 m, MZUSP 108767. Paratypes: 1, 24.0 mm (Figure 51), dd, off Camocim, Estado Cear��, 15���25 m, diver, 2007, ANSP 422778; 1, 26.4 mm, dd, off Camocim, 20-25 m, in octopus pot, LC; 1, 22.1mm, dd, same data, LC; 1, 27.2 mm, dd, off Camocim, 20-25 m, SC; 1, 25.3 mm, dd in octopus pot, off Camocim, 20���35 m, BMSM 17941; 1, 21.2 mm, dd in octopus pot, off Camocim, 20���35 m, NHMUK 20120248; 1, 22.9 mm, dd, off Camocim, 25-30 m, USNM 1192973; 1, 28.3 mm, crabbed in octopus pot, dd, off Camocim, 25- 35 m, LC; 1, 27.7 mm, dd ex pisce, off Fortaleza, Estado Cear��, 16 m, 7 / 1968, ANSP 449751; 1, 25.8 mm, dd ex pisce, off Fortaleza, 12 fm (22 m), AMNH 140147; 1, 23.5 mm, dd, off Rio do Fogo, north of Natal, Rio Grande do Norte, 15���25 m, sand under rock, 2008, ANSP 422777; 2, 16.0 and 20.6 mm, dd, off Rio do Fogo, north of Natal, 15-25 m, UF 455470; 1, 23.7 mm, dd, north of Natal, Rio Grande do Norte, 10���15 m, MNHN 25671; 1, 15.6 mm, dd, north of Natal, 10-15 m, sand under rock, diver, 1999, ANSP 449750; 1, 11.1 mm, dd, north of Natal, 10-15 m, sand under rock, 2000, ANSP 449749; 7, 18.1, 17.1, 17.0, 16.8, 15.7, 12.2 and 10.4 mm, lv, north of Natal, 10���15 m, sand under rock, 2007, ANSP 421133; 1, 22.1 mm, dd, north of Natal, 10���15 m, SC; 1, 18.0 mm, lv, north of Natal, 25 m, 11 / 2003, ANSP 449746; 1, 15.8 mm, lv, Rio Grande do Norte, reef, 10-20 m, UF 455467; 1, 17.0 mm, lv, Rio Grande do Norte, 10-20 m, ANSP 449747; 1, 26.7 mm, dd, off Alcoba��a, Estado Bah��a, 20-25 m, LC; 2, 26.5 (Figures 49���50) and 21.7 mm, dd, off Alcoba��a, Bah��a, Brazil, 40-45 m, ANSP 449745. Other Material: Northeastern Brazil ��� 23, 15.4, 15.3, 14.3, 14.8, 14.3, 13.8, 13.7, 13.6, 13.5, 13.2, 13.2, 13.0, 12.8, 12.6, 12.5, 12.5, 12.3, 12.0, 11.9, 11.7, 11.1, 10.0 and 8.6 mm, dd, north of Natal, 10-15 m, sand under rock, 2004, ANSP 449753; 2, dd, 17.3 and 17.0 mm, north of Natal, Rio Grande do Norte, 10���15 m, LC; 1 lv, 16.8 mm, 13 dd, 10.2-21.1 mm, north of Natal, 10���15 m; 7 lv, 13.3-15.6 mm, Rio Grande do Norte, 10-20 m, LC; 2, 13.6 and 15.5 mm, lv, Rio Grande do Norte, 25 m LC. Venezuela? (records considered spurious)��� 1, 16.1 mm, lv, Los Roques Islands, Venezuela, 6 m, LC; 1, 10.2 mm, dd, off Los Roques Islands, dredged, 200 m, 2006, ANSP 449752. Type Locality: Off Natal, Estado Rio Grande do Norte, Brazil, 10��� 15 m. Etymology: The species name, an adjective, is composed of the prefix bi-, meaning two, the Latin noun ocellus, meaning little eye, and the suffix - atus, - a, - um, meaning ���provided with,��� referring to the pattern of paired spots on the body whorl. Distribution: Off Camocim, Cear�� southward to Alcoba��a, Bah��a, Brazil; depth range 10��� 45 m. Remarks: Pustulatirus biocellatus seems relatively common in moderate depths (10���45 m) on the inner continental shelf off of the states of Cear��, Rio Grande do Norte, and Bah��a in northeastern Brazil. We examined two shells labeled ���off Los Roques, Venezuela,��� depths 6 m and 200 m, but we consider those data suspect and requiring confirmation. This species was reported and figured as P. e p p i (Melvill, 1891) by Vermeij and Snyder (2006), and Brazilian shells have been offered for sale as ��� Latirus ��� eppi, or sometimes as L. cf. L. eppi, for more than a decade. Between 1968 and 1994, records of the species were mistakenly reported as L. virginensis Abbott, 1958 (see synonymy). Recently, Rios (2009) reported and figured Brazilian specimens as L. (Polygona) virginensis, and then cited L. eppi as a junior synonym. However, we have found no valid record of the occurrence of either of those Caribbean species in Brazil, and we reject that synonymy. The pattern of spots on the body whorl immediately distinguishes P. biocellatus from P. eppi and P. virginensis. Although their shells are similar in size, P. biocellatus has more rounded whorls, deeper sutures, and more conspicuous spiral cords than P. e p p i. Shells of P. virginensis are larger and more attenuate than P. biocellatus, with longer, narrower ribs and a relatively longer and more slender siphonal process., Published as part of 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, pp. 35-58 in Zootaxa 3636 (1) on pages 48-49, DOI: 10.5281/zenodo.283572

Family Fasciolariidae J. E. Gray, 1847, Published as part of 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, pp. 35-58 in Zootaxa 3636 (1) on page 36, DOI: 10.5281/zenodo.283572

Pustulatirus virginensis (Abbott, 1958) (Figures 8���30) [?] Lathyrus annulata [sic] (Bolt. Link)���M��rch, 1852: 99. Non Syrinx annulata R��ding, 1798. [?] Turbinella annulata (Bolt.) ���Krebs, 1864: 16. Poulsen, 1878: 11; Dall, 1885: 314; Clench et al., 1947: 35. Non Syrinx annulata R��ding, 1798. [?] Peristernia annulata (Bolten) ���Melvill, 1891 b: 407. Non Syrinx annulata R��ding, 1798. [?] Peristernia annulata (A. Ad.)���Melvill, 1891 b: 411, pl. 2, fig. 1. Non Syrinx annulata R��ding, 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; Saras��a and Espinosa, 1984: 8, 9, 18, fig. 6 c; 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) R��ding���Trew, 1990: 13. Latirus eppi Melvill y Schapman [sic], 1891 ���Espinosa et al., 1994: 113. Non Latirus eppi, = Pustulatirus eppi (Melvill, 1891), Cura��ao. Latirus (Latirus) eppi Melvill y Schapman [sic], 1891 ���Espinosa et al., 1995: 36. Non Latirus eppi Melvill, 1891. Latirus annulata [sic] (R��ding, 1798)���Mallard and Robin, 2005: 16. Non Syrinx annulata R��ding, 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] (R��ding, 1798)���Vermeij and Snyder, 2006: 421. Non Syrinx annulata R��ding, 1798. Pustulatirus attenuata [sic] (Reeve, 1847)���Vermeij and Snyder, 2006: 421, fig. 4 B. Non Pustulatirus attenuatus (Reeve, 1847),?Caribbean region. Pustulatirus virginensis (Abbott, 1958) ���Vermeij and Snyder, 2006: 421, fig. 4 H. 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, Saman��, 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, Saman��, 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, Bah��a 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, 1986 a, 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, M��xico, 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 R��ding, 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.] R��ding (1798) reallocated the Chemnitz figures cited by Gmelin, Published as part of 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, pp. 35-58 in Zootaxa 3636 (1) on pages 38-43, DOI: 10.5281/zenodo.283572

Genus Pustulatirus Vermeij and Snyder, 2006 Type Species: Latirus mediamericanus Hertlein and Strong, 1951 (Figures 1���4), Recent, tropical eastern Pacific, by original designation (Vermeij and Snyder 2006). Revised Diagnosis: Shells of small to moderate size, largest to about 100 mm sl, generally slender, with tall spire and well-developed siphonal process. Shells ornamented with narrow to broad axial ribs and low spiral cords, latter often most developed near outer lip; whorl surface sometimes nearly smooth, waxy on central sectors and on abapical end of siphonal process; outer lip markedly convex just above basal constriction, edge rendered serrate by extensions of interspaces between spiral cords of body whorl; inner side of outer lip with beaded lirae resembling pustules; 3 or 4 plicae aligned obliquely to axis of columella, sometimes with a smaller plica adapically; adapical and abapical sinuses distinct. Radula formula 1 - 1 - 1, median tooth quadrate, with 3 cusps; lateral teeth broad, somewhat curved; species for which radula known with 6 slender, well-developed, subequal cusps, sometimes with single smaller cusps at inner and/or outer corners. Remarks: Vermeij and Snyder (2006) treated Pustulatirus as feminine. However, Article 30.1. 1 of the International Code of Zoological Nomenclature (ICZN 1999) states, in part, ���a genus-group name that is or ends in a Latin word takes the gender given for that word ���,��� in this case that word being the masculine Latirus. To the original description of Pustulatirus (Vermeij and Snyder, 2006) we add a diagnosis of the radula, figured for a Caribbean species by Abbott (1958) and Bullock (1968), and we note the serrate edge of the outer lip, which we found to occur consistently on all western Atlantic and all but one eastern Pacific species. We caution that pustulose lirae and serrate outer lips are features of mature shells and may be poorly developed or absent on immature specimens. We did not find serrate edges on outer lips of Hemipolygona, Polygona, Leucozonia Gray, 1847, Opeatostoma Berry, 1958, or Bullockus, Lamellilatirus, and Lightbournus Lyons and Snyder, 2008, the other New World peristerniine genera. Outer lips of some Fusolatirus Kuroda and Habe, 1971, an Indo-west Pacific genus, also have serrate edges, but their internal lirae are uninterrupted, not beaded or pustulose. Pustulose lirae are not diagnostic solely for Pustulatirus. Lyons and Snyder (2008) also described pustulose lirae in Lamellilatirus and Bullockus. Pustules varied from few and faint to numerous and distinct among seemingly mature specimens of single species. We did not consistently find pustulose lirae or labral serrata on shells of Latirus hemphilli Hertlein and Strong, 1951, a Panamic species placed in Pustulatirus by Vermeij and Snyder (2006). Both features were uncommon among 44 specimens in 21 lots from M��xico and Panam��. When present, the features tended to occur on immature shells or, in the case of labral serrata, on repaired lips of adult shells. The type species and other eastern Pacific Pustulatirus species have small, tapered protoconchs that suggest a planktotrophic veliger whereas protoconchs of western Atlantic species are lecithotrophic, suggesting demersal development. The eastern Pacific P. mediamericanus and P. praestantior (Melvill, 1891 c) are larger than any Atlantic congener, attaining sizes of about 91 and 100 mm respectively (Pisor & Poppe 2008) whereas the largest Atlantic species, P. ogum and P. virginensis, attain only 53.2 and 52.8 mm (both herein)., Published as part of 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, pp. 35-58 in Zootaxa 3636 (1) on pages 36-37, DOI: 10.5281/zenodo.283572

Pustulatirus eppi (Melvill, 1891) (Figures 43���45) Latirus eppi Melvill, 1891 a: 158. Melvill, 1891 b: 394, 411, pl. 2, fig. 11; Horst and Schepman, 1894: 92; Schepman, 1916: 477; van Bentham Jutting, 1927: 6, 32; Coomans, 1958: 49, 93; Warmke and Abbott, 1961: 14; de Jong and Kristensen, 1965: 39; Bullock, 1968: 69, 99, 101, pl. 4, fig. 10; Tello, 1975: 125; Princz, 1982: 123; Trew, 1987: 38; de Jong and Coomans, 1988: 88, pl. 25, fig. 479; Trew, 1990: 2; Lyons, 1991: 177, 178, fig. 42, 43; D��az, 1995: 118; Snyder, 2003: 91; Mallard and Robin, 2005: 17 (pars); Rios, 2009: 253 (pars); Faber, 2010: 8, fig. 1; van der Bijl et al., 2010: 54, 4 figs. Non Latirus eppi Espinosa et al. (1994: 113; 1995: 36; both Cuba), Mallard and Robin (2005: pl. 43, right figs. only, ���Porto Rico���), and Robin (2008: 221, figs. 2, = same figs. of shell from ���Porto Rico���), all = P. v i rg i n e n s i s (Abbott, 1958); nec Latirus eppi auctt. = P. biocellatus Lyons and Snyder, herein. Pustulatirus eppi (Melvill, 1891) ���Vermeij and Snyder, 2006: 421 (pars; non fig. 4 C, = Pustulatirus biocellatus n. sp.). Description: Shell small for genus (largest 24.1 x 10.8 mm), solid, broadly fusiform, axially compressed and compact, with rapidly expanding whorls, broad axial ribs, and low spiral cords. Protoconch too worn for description. Teleoconch with about 6 whorls, suture slightly undulant in accord with ribs and interspaces, sutural ramp lacking; whorls shorter than wide, with 8 or 9 axial ribs extending from suture to suture; spiral sculpture of low cords, 3 or 4 cords on whorls 2 and 3, 4 or 5 cords on whorl 4, cords on later whorls becoming nearly obsolete except for about 3 cords near adapical suture and 1 or 2 on body whorl near junction with siphonal process; about 5 stronger, oblique cords on dorsal surface of broad, abapically tapering siphonal process. Aperture ovo-elongate, constricted adapically by parietal node at posterior sinus and abapically by node opposite entrance fold at base of columella; outer lip lowly arcuate, with flexure near junction with siphonal process; outer lip smooth, worn on specimens examined, but with remnants between termini of cords suggesting extensions of interspaces; inside of outer lip with about 6 pustulose lirae emerging from aperture; inner lip smooth, adherent, with 3 or 4 oblique plicae on abapical half. Shell surface chestnut brown, apparently fading to yellow in older dead shells, interior white. Operculum and radula unknown. Type Material: Holotype 24.1 x 10.8 mm (Figures 43���45), dd, RMNHL. Type Locality: Cura��ao, Netherlands Antilles. Other Material: 1, 20.0 mm, dd, beach at Boca Santa Maria, Cura��ao, ZMA. Etymology: Named for Dr. Epp, its discoverer (Melvill, 1891 a); Carolus T. Epp was a pharmacist with the Netherlands West Indian army on Cura��ao from 1876 until 1886 (van der Bijl et al. 2010). Distribution: The species is known only from Cura��ao, Netherlands Antilles, where it seems to be both endemic and rare. Remarks: The authorship of Latirus eppi has been a subject of debate. The name was attributed solely to Melvill by van Benthem Jutting (1927), Coomans (1958), de Jong and Coomans (1965), Princz (1982), Trew (1987, 1990), Snyder (2003) and Mallard and Robin (2005). Others (Horst & Schepman 1894, Schepman 1916) attributed the name to Schepman and Melvill; and still others (Bullock 1968, de Jong & Coomans 1988, Lyons 1991, D��az 1995, Rios 2009) attributed authorship to Melvill and Schepman, ���Melvill y Schapman [sic]��� (Espinosa et al. 1994, 1995) or, most recently to Melvill and Schepman in Melvill 1891 (van der Bijl et al. 2010, Faber 2010). Melvill (1891), in Notes from the Leyden Museum described but did not figure Latirus eppi, taking credit for the name which he stated that Schepman had asked him to describe. Melvill's note bears the date ���March, 1891,��� but that may be when he wrote it; the paper was published in August 1891 (Dickinson, 2005). In another description (with figure) published the same year in Memoirs and Proceedings of the Manchester Literary and Philosophical Society, Melvill (1891: 394) stated that ���Mr. Schepmann [sic] unites with me in joint authorship��� of Latirus eppi. Melvill cited himself as sole author in the caption for the figure of the type (p. 411, pl. 2, fig. 11), while on the same page Peristernia retiaria Melvill (pl. 2, fig. 13) was attributed to Melvill and Schepmann [sic]. Nothing in the text associates Schepman with Peristernia retiaria, so we surmise that ���Melvill and Schepmann��� was intended for Latirus eppi, and ���Melvill��� was intended for Peristernia retiaria. As indicated on its title page, the manuscript for Melvill���s second paper was received on 24 March 1891, but Rosenberg (2009) cites its publication date as ���post-July��� of that year, so it too could have been published in August. Article 21.3. 1 of the International Code of Zoological Nomenclature, Fourth Edition (ICZN, 1999) specifies that when the year and month but not the day of publication is known with certainty, the date to be adopted is the last day of the month, and Article 21.3. 2 specifies that when neither the day nor the month of publication is known with certainty, the date to be adopted is the last day of the year. In the case at hand, the Code dictates that publication of Notes from the Leyden Museum be assigned a date of 31 August 1891 (i.e., 1891 a), and Memoirs and Proceedings of the Manchester Literary and Philosophical Society be assigned a date of 31 December 1891 (i.e., 1891 b). The International Code of Zoological Nomenclature, Fourth Edition (ICZN 1999), Article 50.1 also states in part: ���The author of a name ��� is the person who first publishes it... in a way that satisfies the criteria of availability.��� As Melvill is the only author identified in the first publication (1891 a), Latirus eppi must be attributed solely to Melvill, and efforts to add Schepman���s name as an author are contrary to provisions of the Code. Bullock (1968: 69, pl. 4, fig. 10) figured the holotype of Latirus eppi, which he treated tentatively as a ���squat specimen, quite unlike the type specimen of [L.] attenuatus.��� Bullock maintained that ���a series of intergrades appear to exist between the two forms,��� but he conceded that further study could result in their separation. Bullock may have been confused by the figure that Melvill (1891 b) provided for L. eppi; the shell in that figure looks more like a specimen of P. virginensis than it does the holotype and other material of P. e p p i as figured by Bullock (1968), Lyons (1991), van der Bijl et al. (2010), Faber (2010) and herein. Those shells certainly are not conspecific with P. v i rg i n e n s i s. Melvill���s figure also may have influenced Mallard and Robin (2005) to misidentify a shell of P. virginensis from Puerto Rico as L. eppi. Shells of P. vi rgi ne n s is are larger, relatively more slender, seldom occur in a single color, and usually have more elevated ribs crossed by more prominent spiral cords than do shells of P. eppi. Dr. Jos�� Espinosa kindly provided a photograph of a specimen that had been reported as Latirus eppi from Cuban waters (Espinosa et al. 1994, Espinosa et al. 1995); that specimen proved to be P. virginensis of a small, dark form with white ribs that is rather common in the Bahama Islands; we illustrate a similar shell (ANSP 449719) in our Figures 27���28. Pustulatirus eppi seems limited to waters around Cura��ao. Most applications of its name (see synonymy) either have been reiterations of a few early records or misidentifications, including a new species described in the next account but also including a previously named species (see remarks for P. virginensis). Of the many Brazilian shells that we have seen offered for sale as Latirus eppi during the past decade, all have proved to be the new species., Published as part of 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, pp. 35-58 in Zootaxa 3636 (1) on pages 46-48, DOI: 10.5281/zenodo.283572

Subfamily Peristerniinae Tryon, 1881., Published as part of 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, pp. 35-58 in Zootaxa 3636 (1) on page 36, DOI: 10.5281/zenodo.283572

Pustulatirus attenuatus (Reeve, 1847) (Figures 5���7) Turbinella attenuata Reeve, 1847: pl. 13, fig. 69. Reeve, 1860: 121; Krebs, 1864: 16; Kobelt in K��ster and Kobelt, 1876: 101, 102, pl. 24, fig. 5; Dall, 1885: 314; Clench et al., 1947: 35; Bullock, 1968: 67; Snyder, 2003: 45. Lathyrus attenuata [sic] (Reeve)���M��rch, 1852: 99. Latirus attenuatus (Reeve) ���H. and A. Adams, 1853: 152. Kobelt, 1877: 58; Tryon, 1881: 90, 234, 299, pl. 67, fig. 122, pl. 68, fig. 144; Paetel, 1887: 162; Abbott, 1958: 77; Bullock, 1968: 26; Santos Galindo, 1977: 221; Sutty, 1986: 62. Turbinella (Plicatella) attenuata Reeve���Kobelt, 1876: 20. Turbinella attenuata ? Reeve���Arango y Molina, 1880: 221. Plicatella attenuata (Reeve) ���Dall, 1885: 240. Latirus alternatus [sic] (Reeve)���Melvill, 1891 b: 403. Latirus [unnamed subgenus] attenuatus (Reeve) ���Bullock, 1968: 67 (pars). Non Latirus attenuatus Bullock (1968: 67���69, pl. 4, figs. 3 ���5, 10, pl. 5, figs. 11, 12, 14), = Pustulatirus virginensis (Abbott, 1958), Recent, eastern Caribbean. Turbinella attenuale [sic] Coomans, 1974: 185. [?] Latirus attenuata [sic] (Reeve, 1847)���Mallard and Robin, 2005: 16, pl. 40. Pustulatirus attenuata [sic] (Reeve)���Vermeij and Snyder, 2006: 421 (pars). Non Pustulatirus attenuata [sic] Vermeij and Snyder (2006: fig. 4 B), = Pustulatirus virginensis (Abbott, 1958), Recent, eastern Caribbean. Description: Shell small for genus (holotype 31.1 mm sl), narrowly fusiform, with rounded whorls, broad axial ribs, well-developed spiral cords, shallow suture, and narrow post-sutural ramp bearing closely-packed axial lamellae. Teleoconch of about 7 regularly expanding convex whorls separated by shallow suture; suture undulant in accord with adjacent whorls and interspaces; each whorl with about 8 broad axial ribs; 4 subequal spiral cords on first whorl, enlarging in size but not increasing in number on succeeding whorls of spire, 10���11 cords on body whorl; cords on sutural ramp low, parallel to and undulating in concert with suture, crossed by numerous welldeveloped subsutural lamellae; cords crossing ribs larger than those of ramp, sometimes with single spiral threads between; 6 or more oblique cords of unequal size on siphonal process, occasionally with single smaller thread between. Aperture ovate, constricted adapically by thick parietal node and abapically by small tooth-like node opposite fold at base of columella; outer lip broadly arcuate, rendered serrate by extensions of interspaces between spiral cords, inner surface with about 8 beaded lirae; columella with 4 oblique plicae, another smaller plica adapically; siphonal canal typical of genus, outer lip thin, crenulated by termini of interspaces between larger dorsal cords, inner lip simple, straight. Shell outer surface yellow with white axial ribs on first 3 or 4 teleoconch whorls, interior white. Operculum and radula unknown. Type Material: Holotype (Figures 5���7), 31.1 mm, dd, locality unknown, NHMUK 196735. Type Locality:Unknown; probably tropical western Atlantic. Remarks: The holotype of Turbinella attenuata Reeve, 1847 resembles shells in the Pustulatirus virginensis species-complex of the eastern Caribbean and also resembles shells of a new species from Honduras and Panam�� that is described herein, yet it differs from both. Described without locality, T. attenuata was soon recognized as a western Atlantic species by nineteenth century authors (M��rch 1852, Krebs 1864, Kobelt in K��ster & Kobelt 1876, Kobelt 1876, 1877, Arango y Molina 1880). However, Kobelt in K��ster and Kobelt (1876) suggested that the name might represent a variety of Turbinella infundibulum (Gmelin, 1791), now the type species of Polygona Schumacher, 1817, prompting Tryon (1881) to relegate the name to synonymy with that species. Tryon���s action effectively shelved the name until Abbott (1958) mentioned specimens at ANSP, previously labeled Latirus attenuatus, among those he was naming L. virginensis. Bullock (1968, in thesis) also addressed the name but distinguished it as a species separate from L. virginensis and others in a related species-complex (see below). The name (as Turbinella attenuale [sic]) also appeared on an early list of shells from St. Martin prepared by H. E. Rijgersma, but Coomans (1974) dismissed it as a supposed synonym of Latirus [= Polygona] brevicaudatus (Reeve, 1847). Then Sutty (1986: 62, 64, 65, fig. 70) reported and figured as ��� Latirus species (cf. attenuatus (Reeve, 1847)) ��� a shell from Guadeloupe that in her opinion ���bears a distinct resemblance to Latirus virginensis Abbott, 1958 and, better still, [to] L. attenuatus (Reeve, 1847).��� Thereafter, Snyder (2003) cited L. attenuatus as a valid Caribbean species that ranged from Cuba to the Lesser Antilles, apparently sympatrically with L. virginensis. Mallard and Robin (2005) cited L. attenuata [sic], range Cuba to Lesser Antilles, and L. virginensis, range ���Caribbean,��� again suggesting sympatry. Those authors speculated that attenuata may be the species they figured as L. bernadensis Bullock, 1974, on their plate 41, but that shell was correctly identified and is a species of Polygona. Vermeij and Snyder (2006) reclassified both attenuatus and virginensis in Pustulatirus and treated them as distinct. In this we concur. However, we include other shells figured as attenuatus by Bullock (1968), Sutty (1986) and Vermeij and Snyder (2006) in Pustulatirus virginensis., Published as part of 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, pp. 35-58 in Zootaxa 3636 (1) on pages 37-38, DOI: 10.5281/zenodo.283572

Subfamily Peristerniinae Tryon, 1881.

Pustulatirus utilaensis new species (Figures 32���37) Latirus virginensis Abbott, 1958 ���K. and L. Sunderland, 1996: 17, 2 figs. Non Latirus virginensis Abbott, 1958, Recent, eastern Caribbean. Description: Shell small for genus (largest 31.6 x 11.5 mm), moderately fusiform, with rounded whorls, broad, somewhat adapically shouldered axial ribs, low spiral cords, shallow suture, and narrow sutural ramp bearing closely-packed axial lamellae. Protoconch of about 2 whorls; first whorl smooth, broadly globose, with rounded sides; second whorl no wider than first, with straight sides, about 4 broad axial riblets near terminus. Teleoconch of 6 to 7 rapidly expanding convex whorls, each with 8 or 9 broad axial ribs, whorls separated by well-developed suture; suture undulant in accord with adjacent ribs and interspaces; 5 subequal spiral cords on first whorl, becoming more numerous on each succeeding whorl, 10-11 cords on penultimate whorl and 14���16 on body whorl; cords on sutural ramp low, parallel to and undulant in concert with suture, and crossed by numerous welldeveloped subsutural lamellae; cords crossing ribs larger than those of ramp, sometimes with single spiral threads between; 6 or more oblique cords of unequal size on siphonal process, occasionally with single smaller threads between, larger oblique cords raised at intersections with subaxial growth lines, forming lamella-like scales, similar scales sometimes on anterior-most cord of body whorl. Aperture ovate, constricted adapically by thick parietal node and abapically by tooth-like node opposite fold at base of columella; outer lip broadly arcuate, crenulated by extensions of interspaces between spiral cords, inner surface with 10���12 beaded lirae; columella with 2 or 3 oblique plicae, sometimes with smaller plicae adapically; siphonal canal typical of genus, outer lip thin, crenulated by termini of interspaces between larger dorsal cords, inner lip simple, straight, slightly raised, sometimes forming shallow pseudoumbilicus near tip. Outer shell surface uniformly reddish brown or dark brown, interior white. Operculum thin, corneous, yellow or light brown, ovate, with anterior terminal nucleus, outer surface with many arcuate growth increments. Radula unknown. Type Material: Holotype 29.2 x 11.2 mm (Figures 32���34), lv, off Sandy Key Reef, Utila, Bay Islands, Honduras, depth 70 ft [21.4 m], ANSP 449739. Paratypes: Honduras ��� 5, 28.6, 25.8 and 24.0 mm, lv, 2, 13.4 and 13.2 mm, dd, with same data as holotype, SC; 4, 26.6 (Figure 35), 22.6, 18.9 and 18.9 mm, lv, from live reef off southwestern coast of Utila, 40���60 ft [12.2���18.3m], ANSP 449664; 2, 27.1 and 16.7 mm, lv, high-profile reef at Utila, 65 ft [19.8 m], LC; 1, 26.5 mm, lv, same data, USNM 1192971; 1, 30.4 mm, lv, southern coast of Utila, outcrop of rock wall, 40 ft [12.2 m], MNHN 25670; 5, 27.2, 25.9, 25.4, 20.9 and 18.4 mm, lv, same data, LC; 1, 28.4 mm, lv, same data, BMSM 17940; 1, 22.3 mm, lv, off Oyster Bed Lagoon, Utila, coral reef at night, 10-15 m, ANSP 449741; 1, 14.0 mm, lv, Ragged Cay, Utila, reef at night, 12 m, ANSP 449740; 1, 15.5 mm, dd, Ragged Cay, Utila, reef at night, 13 m, ANSP 449742; 2, 21.7 and 16.7 mm, dd, Utila, high profile reef at night, 65 ft (19.8 m), LC; 2, 28.3 and 26.0 mm, lv, Utila, UF 455468; 1, 26.5 mm, lv, Utila, NHMUK 20120249; 1, 22.1 mm, lv, Utila, 20���30 ft [6.1���9.1 m], diver, 5 / 2008, ANSP 449743; 1, 17.5 mm, lv, Utila, 20 ft [6.1 m], night dive, LC; 2, 16.0 and 14.1 mm, dd, Utila, 60 ft [18.3 m], ANSP 449662; 1, 17.0 mm, dd, Utila, reef, ANSP 449660; 1, 25.4 mm, lv, pillar coral reef, Roat��n, Honduras, 30 ft [9.1 m], UF 455471; 1, 25.4 mm, lv, reef at Roat��n, 40 ft [12.2 m], LC; 1, 31.6 mm (Figures 36���37), lv, Roat��n, living reef, 10-12 m, ANSP 449659. Panam�� ��� 1, 21.6 mm, dd, Isla Escudo de Veraguas, eastern Panam��, depth 12-15 m, under rocks, ANSP 449661; 1, 22.3 mm, dd, same data, LC; 1, 22.1 mm, dd, Isla Escudo de Veraguas, Panam��, 20-25 m, ANSP 449663. Other Material: Honduras ��� 1 worn shell, 27.2 mm, dd, Utila, live reef off southwestern coast, 12.2���18.3 m, LC; 1 shell with broken apex, 21.6 mm, dd, Utila, reef, LC; 3 broken or encrusted shells, 25.5 and 23.3 mm, dd and 21.1 mm, lv, same data as holotype, SC. Type Locality: Off Sandy Cay Reef, Utila, Bay Islands, Honduras, depth 21.4 m. Etymology: The species is named for the island of Utila, Bay Islands, Honduras, Caribbean Sea, where it seems to be most abundant. Distribution: Known principally from vicinity of Utila, Bay Islands, Honduras, depth range: 6.1���21.4 m; three specimen examined were from nearby Roat��n, Honduras, and three more were from Isla Escudo de Veraguas, eastsoutheast of Bocas del Toro, Panam��. Remarks:All Honduran shells of P. utilaensis are identical morphologically, differing only in color related to their localities of capture. Specimens from Utila are consistently reddish brown, but all six shells from Roat��n, Honduras and Isla Escudo de Veraguas, Panam�� are dark brown. The Panam�� shells are more worn and smooth, but their patterns of ribs and cords as well as their size and general outlines are consistent with those of Honduran specimens and we believe them to be conspecific. Morphological features of P. utilaensis are consistent within parameters defined in the description, and unworn shells never have smooth surfaces or lighter colored axial ribs like those that often occur on P. virginensis, whose shells are often larger than the maximum length known for P. utilaensis. Shells of P. utilaensis also resemble the type of P. attenuatus, but the latter is a relatively more slender shell with axial ribs that are less shouldered adapically; P. utilaensis shells are consistently reddish brown or dark brown whereas the type of P. attenuatus is yellow with white ribs on early adapical whorls., Published as part of 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, pp. 35-58 in Zootaxa 3636 (1) on pages 43-45, DOI: 10.5281/zenodo.283572

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