The Genus Pustulatirus Vermeij and Snyder, 2006 (Gastropoda: Fasciolariidae: Peristerniinae) in the Western Atlantic, with Descriptions of Three New Species
Author
Lyons And Martin Avery Snyder, William G.
text
Zootaxa
2013
3636
1
35
58
journal article
10.5281/zenodo.283572
61a9ddf5-0a97-4757-8217-e72d41a770a7
1175-5326
283572
C2B24CC9-EE3D-43DC-AB13-22B7346C93DA
Pustulatirus virginensis
(Abbott, 1958)
(
Figures 8–30
)
[?]
Lathyrus annulata
[
sic
] (Bolt. Link)—Mørch, 1852: 99.
Non
Syrinx annulata
Röding, 1798
.
[?]
Turbinella annulata
(Bolt.)
—Krebs, 1864: 16. Poulsen, 1878: 11; Dall, 1885: 314; Clench
et al
., 1947: 35.
Non
Syrinx annulata
Röding, 1798
.
[?]
Peristernia annulata
(Bolten)
—Melvill, 1891b: 407.
Non
Syrinx annulata
Röding, 1798
.
[?]
Peristernia annulata
(A. Ad.)
—Melvill, 1891b: 411, pl. 2, fig. 1.
Non
Syrinx annulata
Röding, 1798
.
Latirus (Polygona) virginensis
Abbott, 1958: 76
, 77, text-figs. 4.7, 4.8, pl. 2, fig. 6. Rios, 1970: 96 (
pars
); Abbott, 1974: 227; Rios, 1975: 104 (
pars
); Ortiz-Corps, 1983: 121; Rios, 1985: 107, pl. 36, fig. 471 (
pars
); Rios, 1994: 133 (
pars
); Espinosa
et al
., 1995: 36; Harasewych, 1997: 62; Snyder, 2003: 218; Rios, 2009: 253 (
pars
).
Non
Latirus (Polygona) virginensis
‘Abbott’
auctt
. northeastern
Brazil
, =
Pustulatirus
n. sp.
Latirus virginensis
—Abbott, 1958: 77. Warmke and Abbott, 1961: 120, 272, pl. 22, fig. n; Arnow
et al
., 1963: 169; Wagner and Abbott, 1964: 167; Holeman, 1966: 29, 30; Wagner and Abbott, 1967: 247; Nowell-Usticke, 1971: pl. 6, fig.; Morris and Clench, 1973: 218, pl. 60, figs. 4; Abbott, 1974: color pl. 11, fig. 2493; Ekdale, 1974: 648; Lipka, 1974: 155; Riggs, 1975: 14; Kaicher, 1978: card 1763; Wagner and Abbott, 1978: 80–231; Woodlock, 1980: 189; Pointier
et al
., 1982: 9; Sarasúa and Espinosa, 1984: 8, 9, 18, fig. 6c; Sutty, 1986: 62; Pointier
et al
., 1987: 12; Faber, 1988: 82; Lyons, 1991: 177, 178, figs. 36-41; Espinosa
et al
., 1994: 113; Goto and Poppe, 1996: 393; K. and L. Sunderland, 1996: 17, 2 figs.; Pointier and Lamy, 1998: 131, 2 figs.; Snyder, 2000: 162; Redfern, 2001: 103, pl. 46, fig. 432; Snyder, 2003: 121; Henkel and Kurtz, 2004: 14, 15, fig.; Mallard and Robin, 2005: 19, pl. 52; Rios, 2009: 253 (
pars
); Landau and Vermeij, 2012: 88.
Non
Latirus virginensis
‘Abbott’
auctt
., northeastern
Brazil
, =
Pustulatirus
n. sp.
Latirus
[new subgenus]
annulatus
(Melvill)
—Bullock, 1968: 65-67, pl. 4, figs. 2, 14, pl. 5, fig. 10.
Latirus
[new subgenus]
attenuatus
(Reeve)
—Bullock, 1968: 67-69, pl. 4, figs. 3–5, 10, pl. 5, figs. 11, 12, 14 (
pars
).
Non
Pustulatirus attenuatus
(Reeve, 1847)
,?Caribbean region,
nec
Latirus eppi
Melvill, 1891
, treated by Bullock as junior synonym of
attenuatus
.
Latirus
[new subgenus]
virginensis
Abbott—Bullock, 1968: 71
-73, pl. 4, figs. 1, 19.
Latirus karinae
Nowell-Usticke, 1969: 18
, pl. 4, fig. 821. Faber, 1988: 82; Lyons, 1991: 178; Boyko and Cordeiro, 2001: 76; Redfern, 2001: 103; Snyder, 2003: 121; Rios, 2009: 253.
Latirus virgineus
[
sic
] (Abbott)—Santos Galindo, 1977: 222.
Latirus elegans
(Gray)
—Kaicher, 1986: card no. 4671 (
pars
; larger [right] shell only).
Non
Fusus elegans
J. E. Gray, 1838
=
Fusinus filosus
(Schubert and Wagner, 1829)
, West Africa, new synonymy.
Latirus
species (
cf.
attenuatus
) (Reeve, 1847)—Sutty, 1986: 62, 64, 65, color fig. 70.
Non
Turbinella attenuata
Reeve, 1847
.
[?] [no genus]
annulatus
(Bolten) Röding—Trew, 1990: 13
.
Latirus eppi
Melvill
y Schapman [
sic
], 1891—Espinosa
et al
., 1994: 113.
Non
Latirus eppi
, =
Pustulatirus eppi
(Melvill, 1891)
,
Curaçao
.
Latirus (Latirus) eppi
Melvill
y Schapman [
sic
], 1891—Espinosa
et al
., 1995: 36.
Non
Latirus eppi
Melvill, 1891
.
Latirus annulata
[
sic
] (Röding, 1798)—Mallard and Robin, 2005: 16.
Non
Syrinx annulata
Röding, 1798
.
Latirus abbotti
Snyder, 2003
—Mallard and Robin, 2005: pl. 39. Robin, 2008: 219, figs. 9.
Non
Latirus abbotti
, =
Polygona abbotti
(Snyder, 2003)
, Caribbean Sea.
Latirus eppi
Melvill, 1891
—Mallard and Robin, 2005: pl. 43 (
pars
; right figs. only, shell from
Puerto Rico
); Robin, 2008: 221, figs.
Non
Latirus eppi
Melvill, 1891
.
Pustulatirus annulata
[
sic
] (Röding, 1798)—Vermeij and Snyder, 2006: 421.
Non
Syrinx annulata
Röding, 1798
.
Pustulatirus attenuata
[
sic
] (Reeve, 1847)—Vermeij and Snyder, 2006: 421, fig. 4B.
Non
Pustulatirus attenuatus
(Reeve, 1847)
,?Caribbean region.
Pustulatirus virginensis
(Abbott, 1958)
—Vermeij and Snyder, 2006: 421, fig. 4H. Rosenberg
et al
., 2009: 654; Zhang, 2011: 129, figs. 445(1–3); Landau and Vermeij, 2012: 88.
Pustulatirus
sp. Zhang, 2011: 129, figs. 446(1–3).
Description
: Shell of medium size for genus (largest 52.7 x
18.7 mm
), elongate, fusiform, slender, with rounded whorls, broad axial ribs, and low spiral cords; outer lip serrate and internal lirae beaded as in generic diagnosis. Variability in shell morphology, size, and color is so extensive that we choose simply to illustrate
types
and several other specimens (
Figures 8–30
) to demonstrate the range of variation. Variation is also exemplified by three shells figured as
Latirus virginensis
by Mallard and Robin (2005) and four shells figured as
Pustulatirus
sp. and
P. virginensis
by Zhang (2011). Further comments on variation appear in remarks for the species.
Type
Material
:
Latirus virginensis
:
holotype
34.2 mm
(
Figures 8–9
), St. Thomas,
Virgin Islands
, ANSP 196459; 2
paratypes
, 32.0 and
29.4 mm
, St. Thomas, ANSP 34975; 2
paratypes
,
36.6 mm
, lv,
29.5 mm
, dd?, St. Thomas, ANSP 34968; 4
paratypes
, 29.8, 29.3, 26.6 (
Figures 10–11
) and
25.4 mm
, dd, “West Indies,” ANSP 34969;
Latirus karinae
:
lectotype
,
32.7 mm
(
Figures 12–13
), “
Puerto Rico
,” AMNH 198490.
Type
Localities
:
Latirus virginensis
: St. Thomas,
Virgin Islands
;
Latirus karinae
: “
Puerto Rico
.”
Other Material
:
Bahama
Islands
—2, 27.1 (
Figure 24
) and
25.5 mm
, lv, reef north of Sandy Cay, West
End
, Grand
Bahama
,
15–17 m
, 1999, ANSP 449716; 2, 20.0 and
10.3 mm
, dd, Sandy Cay, West
End
,
40 ft
[
12.2 m
], coral rubble, LC; 1,
25.5 mm
, lv, Sandy Cay, West
End
, LC; 2 fragments (1 shell?), dd, Gold Rock, Grand
Bahama
,
80 ft
[
24.4 m
], ANSP 369654; 2, 27.9 and
25.8 mm
, lv, Bimini, reef,
35 ft
[
10.7 m
], LC; 1,
24.5 mm
, lv, Bimini, reef,
45 ft
[
13.7 m
], LC; 1,
20.7 mm
(
Figures 27-28
), dd, Bimini, 7/1985, ANSP 499719; 1,
24.2 mm
, lv, Turtle Rocks, South Bimini,
30 ft
[
9.1 m
], LC; 2, 24.6 and
18.9 mm
, lv,
3.2 km
south of Northwest Channel Light,
19.2 km
southwest of Chub Cay,
Bahamas
,
70-90 ft
[
21.4-27.6 m
], LC; 1,
26.4 mm
, dd, Silver Cay Beach, Nassau, New Providence, LC; 2, 33.7 and
24.6 mm
, dd, beach
1.6 km
south of Fresh Creek, Andros, LC; 1, 19.0 mm, dd, off Fresh Creek, Andros, LC; 4, 31.4, 29.7, 20.5 and
20.5 mm
, dd, AUTEC Base, Andros, beached by storm, LC; 1, 28.0 mm, dd, off Windimere, Eleuthera,
3 m
, 6/1988, ANSP 449778.
Turks and Caicos Islands
—1,
23.8 mm
, lv, “
Turks
and
Caicos
,” LC.
Dominican Republic
—1, 30.0 mm (
Figure 25
), lv, 1,
11.7 mm
, dd, northern coast at Castillo, Duarte, rubble,
4 ft
[
1.2 m
], 8/1994, ANSP 449715; 15, 43.3, 42.2, 40.3, 39.2, 39.0, 38.6, 35.2, 33.6, 20.8, 20.0, 19.4, 19.2, 16.3 and 14.0 mm, lv, 14, 40.3, 39.0, 37.2, 32.4, 32.2, 29.4, 27.3, 23.6, 23.1, 23.0, 21.7, 21.0, 19.8 and
19.2 mm
, dd, Las Galeras, Samaná,
3 to 10 ft
[
0.9-3.1 m
],
ex
Glenn Duffy, LC; 1,
41.4 mm
(
Figure 14
), lv, same data, ANSP 449754; 2, 25.1 and
23.6 mm
, lv, Las Galeras, Samaná,
1-2 m
, 1994, ANSP 449791; 1, 20.0 mm, lv, Meces, on dead coral at low water, LC.
Puerto Rico
—1,
21.3 mm
, dd, Bahía Salinas, Cabo Rojo,
15 ft
[
4.6 m
], on
Thalassia
, 3/2000, LC; 1,
16.2 mm
, dd, same locality and date,
18 ft
[
5.5 m
], LC; 1,
34.8 mm
, dd, San Juan Harbor, 1980, ANSP 449771; 1,
24.2 mm
, dd, “
Puerto Rico
,” “
paratype
,” ANSP 219064; 1,
27.4 mm
, dd, “
Puerto Rico
,”
ex
Warmke, UF 162196; 1,
25.7 mm
, dd, “
Puerto Rico
,”
ex
Warmke, UF 162197; 12, 33.1, 32.9, 29.7, 27.9, 27.8, 27.5, 26.7, 26.4, 26.2, 26.0, 21.2 and
20.8 mm
, dd, “
Puerto Rico
,” Usticke Coll. [identified as
Latirus karinae
by Usticke, =
paralectotypes
?], AMNH 191472.
U. S.
Virgin Islands
—2, 27.4 and
19.6 mm
, dd, Krause’s Reef, St. Croix,
ex
Usticke via McGinty, UF 135674; 39, 30.2 to 5.0 mm, dd, St. Croix at Long Reef, 2/1964 and Krause’s Reef, 3/1965, Usticke Coll., AMNH 191463; 2, 24.2 and
21.9 mm
, dd, off St. Croix, 1992, ANSP 449788; 20, 29.0, 26.2, 24.6, 22.3, 22.3, 22.3, 22.2, 22.2, 22.1, 21.7, 21.2, 21.0, 21.0, 20.9, 20.8, 20.6, 20.2, 18.6, 17.4 and
16.3 mm
, Cowpet Bay, St. Thomas,
ex
J. E. Holeman, AMNH
170501
; 2, 21.6 and
9.7 mm
, dd, Cowpet Bay, St. Thomas, AMNH 191476; 3, 33.7, 24.8 and
23.7 mm
, dd, Cowpet Bay, St. Thomas, ANSP 449795; 1,
34.3 mm
, dd, Cowpet Bay, St. Thomas, 1990, ANSP 449784; 3, 30.8, 16.9 and
16.4 mm
, dd, Cowpet Bay, St. Thomas,
ex
J. Holeman via McGinty, UF 156255; 1,
38.6 mm
, dd, Crown Bay, St. Thomas,
ex
Usticke via McGinty, UF 135675; 17, 40.9, 37.5, 36.6, 34.9, 33.3, 31.7, 31.1, 29.3, 27.3, 25.1, 22.9, 20.3, 19.0, 16.7, 14.0, 13.8 and 13.0, dd, Yacht Club, Jessup Bay, St. Thomas, 1969, AMNH 191470; 1,
38.4 mm
, dd, Jessup Bay, St. Thomas,
ex
Usticke via McGinty, UF 156254; 1,
36.2 mm
, dd, St. Thomas Harbor,
ex
J. Holeman via McGinty, UF 135673; 2, 28.0 and
26.9 mm
, dd, St. Thomas Harbor, St. Thomas,
ex
McGinty, UF 135677; 4, 30.6, 17.5, 16.9 and
8.9 mm
, dd, Water
Island
, St. Thomas,
ex
J. Holeman via McGinty, UF 363808; 1,
16.9 mm
, dd, West Gregorie Channel between Water Isle and St. Thomas, ANSP 255163; 1,
35.3 mm
, dd, dredged off St. Thomas, 1980, ANSP 449774; 2, 52.7 (
Figure 15
) and
50.3 mm
, dd, off St. Thomas, dredged, 1987, ANSP 449714; 1,
36.9 mm
, dd, off St. Thomas, 1988, ANSP 449779; 1,
45.4 mm
, dd, off St. Thomas, 1990, ANSP 449786; 3, 30.9, 24.4 and
23.2 mm
, dd, off St. Thomas, 1992, ANSP 449790; 9, 40.0, 33.0, 20.8, 20.2, 16.7, 14.5, 10.8, 9.9 and
8.8 mm
, dd, St. Thomas,
ex
Joan Brindley, 1965, AMNH 182520; 3, 47.7 (
Figure 16
), 25.5 and
25.1 mm
, dd, St. Thomas, 1976, ANSP 449713; 3, 18.0, 15.3 and
13.4 mm
, dd, St. Thomas, Usticke Coll., AMNH 191490; 4, 30.1, 29.1, 25.7 and
16.2 mm
, dd, St. Thomas, AMNH 114569; 1,
24.4 mm
, dd, St. Thomas, 1983, ANSP 449776; 2, 23.2 and 19.0 mm, dd, St. Thomas, ANSP 449796; 4, 29.9, 29.7, 28.4 and
24.2 mm
, dd, St. Thomas, LC; 1,
31.9 mm
, lv, St. Thomas, LC; 1,
30.6 mm
(
Figure 26
), dd, Little St. James
Island
,
24 ft
[
7.3 m
], 6/1996, ANSP 449720; 8, 24.2, 22.3, 20.9, 20.6, 20.4, 19.9, 19.9 and
16.8 mm
, lv, Little St. James
Island
,
27 ft
[
8.2 m
],
7/9/2000
, LC; 1,
26.7 mm
(
Figures 29-30
), same data, ANSP 449718; 1,
24.3 mm
, lv, same data, SC; 6, 24.7, 22.6, 22.6, 22.3, 21.7 and
21.5 mm
, lv, Little St. James
Island
,
27 ft
[
8.2 m
], 7/2002, LC; 5, 38.4, 37.7, 31.2, 28.4 and
27.9 mm
, dd,
St. John
, 1978, ANSP 449748; 2, 47.0 and
45.3 mm
, dd,
St. John
, 1980, ANSP 449775; 1, dd,
20.5 mm
, “
Virgin Islands
,” scuba at night in rubble,
9-10 m
, 1997, ANSP 449793; 2, 31.7 and
26.9 mm
, dd, “
Virgin Islands
,” LC.
British Virgin Islands
—1,
29.4 mm
(
Figure 22
), dd,
1.6 km
NE of East Point,
Anegada
Island
, ANSP 249155; 1,
27.7 mm
,
Anegada
, dd, deep-water fish trap, LC; 1,
29.9 mm
, dd,
0.8-1.6 km
SSW, SSE and E of the Bluff, Sir Francis Drake Channel, Beef
Island
, 12 to 14 fm [22.0-
25.6 m
], ANSP 331162; 2, 10.6 and
7.5 mm
, dd, Beef
Island
, LC; 1, 34.0 mm, dd, Beef
Island
, shallow reef, LC; 1,
29.1 mm
, lv, Buck
Island
,
3 ft
[
0.9 m
], eel grass (
Thalassia
), LC; 11, 26.7, 26.6, 24.0, 20.7, 17.7, 17.5, 17.4, 17.0, 15.5, 12.5 and
11.4 mm
, dd, Fat Hog Bay,
Tortola
,
1–5 m
, in fire coral (
Millepora
) and eel grass (
Thalassia
), 2/1985, ANSP 449777; 2, 21.9 and
19.3 mm
, dd, Fat Hog Bay,
Tortola
,
8 ft
[
2.4 m
], LC; 3 rolled fragments, dd,
Virgin Gorda
, Usticke Coll., AMNH 181489.
Anguilla—
2, 48.5 and
40.6 mm
, dd?,
Anguilla
, 1988, DLC.
St. Martin
—4, 40.6, 37.7, 29.9 and
22.5 mm
, dd, St. Rose, DLC.
St. Barthelemy
—6, 31.7, 24.4, 24.3, 22.0, 20.3 and
19.1 mm
, off St. Barts, Leeward Islands, 1970,
ex
Usticke, ANSP 421134; 1,
32.8 mm
, dd, off
St. Barthelemy
, ANSP 449794; 3, 35.8, 31.0 and
27.9 mm
, lv, 3, 28.3, 26.3 and
24.5 mm
, dd,
St. Barthelemy
, DLC; 1,
34.2 mm
, lv, “
St. Barthelemy
,
Guadeloupe
,” Gaudiat, 1977, DLC; 1,
31.9 mm
, lv?,
St. Barthelemy
, Pointier, 1986, DLC; 1,
33.6 mm
, lv, off
St. Barthelemy
, trap,
100 m
, ANSP 449797; 1,
32.6 mm
(
Figures 19–21
), lv,
St. Barthelemy
,
Guadeloupe
,
100 m
, ANSP 449717.
Barbuda
—3, 25.2, 23.2 and
22.2 mm
, lv, Cocoa Point,
2-5 m
, LC; 2, 24.8 and
20.9 mm
, lv,
Barbuda
,
5 ft
[
1.5 m
], LC.
Antigua
—34, 40.4, 38.1, 37.2, 34.6, 34.6, 33.3, 32.2, 32.2, 31.5, 31.5, 31.5, 31.2, 30.8, 30.1, 29.9, 29.7, 29.4, 29.2, 28.3, 28.3, 26.5, 25.8, 25.6, 25.1, 24.4, 21.9, 21.1, 20.1, 19.0, 18.4, 16.5, 14.5, 12.0 and
11.9 mm
, dd, Maid
Island
, 6/1961, Usticke Coll., AMNH 191465; 4, 34.0, 27.8, 20.6 and
14.1 mm
, dd, Maid
Island
,
ex
Usticke, McGinty Coll., UF 135671; 16, 50.5 (
Figure 17
), 47.9, 43.8, 38.0, 36.2, 35.6, 33.5, 32.7, 25.8, 25.7, 25.3, 22.4, 22.3, 19.9, 18.9 and
17.3 mm
, dd, St. Johns, Usticke Coll., AMNH 191464; 10, 41.6, 33.3, 29.4, 28.5, 28.2, 25.8, 25.5, 25.3, 21.8 and 20.1, dd, St. Johns, 11/1967, Usticke Coll., AMNH 191466; 6, 51.4, 44.8, 43.9, 38.3, 35.3 and
30.3 mm
, dd, St. Johns, Usticke Coll., AMNH 191467; 1,
26.3 mm
, dd,
Antigua
, Lesser Antilles, “
paratype
[of
virginensis
],” ANSP
210740
; 5, 42.3, 38.3, 35.1, 33.2 and
22.9 mm
, dd,
Antigua
Island
, 1979, ANSP 449768; 1,
21.2 mm
, dd,
Antigua
Island
,
ex
Usticke, Warmke Coll., UF 162195; 1,
27.3 mm
, dd,
Antigua
Island
,
ex
Usticke, UF 162198; 4, 39.2, 39.1, 35.8 and
30.4 mm
, dd, off
Antigua
, 1993, ANSP 449792.
Guadeloupe
—4, 31.8, 27,2, 26.7 and
19.8 mm
, dd, 2, 17.0 and
11.1 mm
, lv, Deshaies, Pointier, 1983, DLC; 2, 33.7 and
24.5 mm
, lv, Deshaies,
18 m
, DLC; 2, 38.3 and
31.8 mm
, Pointe Plate, Pointier, 1982, DLC; 1,
32.8 mm
, lv, Port Louis,
10 m
, DLC; 2, 34.7 (
Figure 18
) and
30.3 mm
, dd,
Guadeloupe
,
30-50 m
, ANSP 449712.
Martinique
— 1,
31.8 mm
, Anse Mitan,
ex
J. Holeman via McGinty, UF 135676.
St. Lucia
—2, 37.2 and
37.1 mm
, dd,
St. Lucia
Island
,
ex
C. W. Sheafer via McGinty, UF 135672.
Netherlands
Antilles—
1,
23.4 mm
, dd, Malmok,
Aruba
,
ex
Fr. Fredricus, AMNH 245863; 1,
23.9 mm
, dd, Malmok,
Aruba
,
ex
M. Koolman, AMNH 245864; 2, 28.2 and
25.3 mm
, dd, Malmok,
Aruba
, “de Jong & Bijur,”
ex
Jerome M. Bijur Coll., AMNH 245865.
No locality
– 1,
38.5 mm
(
Figure 23
), dd,
ex
J. S. Phillips, ANSP 35033 (figured as
Latirus elegans
(Gray)
by Kaicher, 1986a, card 4671, right fig.); 1,
24.8 mm
, dd,
ex
Warmke, incorrectly marked as
paratype
, UF 162199; 1,
30.6 mm
, lv, DLC; 1,
39.1 mm
, dd, LC.
Distribution
: This species, the best known of western Atlantic
Pustulatirus
, ranges throughout the eastern Caribbean region from the northernmost
Bahama
Islands to the Greater Antilles, the
Virgin Islands
, and southward to
Antigua
,
Guadeloupe
,
Martinique
,
St. Lucia
, and
Aruba
in the southern
Netherlands Antilles
; there is also an unverified record from Quintana Roo,
México
, in the northwestern Caribbean (Ekdale 1974). Most records with stated depths are from beaches or the shallow subtidal zone (
1–30 m
) but one lot from off
Guadeloupe
is from
30– 50 m
and two from off
St. Barthelemy
are from
100 m
.
Remarks
: Morphology of individual shells may vary greatly, even among specimens from a single location. Some forms are so different morphologically that it is difficult to believe they represent the same species, but the forms invariably blend into others, rendering it difficult to determine where one ends and another begins. With the large number of specimens we examined from throughout the range, we conclude that the various forms represent a single variable species. We hope this conclusion may be tested in the future using genetic information.
The name
Latirus virginensis
Abbott, 1958
, has been used most often for these shells, but several other names have been associated with the group, including
Syrinx annulata
Röding, 1798
,
Turbinella attenuata
Reeve, 1847
,
Peristernia annulata
“(A. Adams)
” Melvill, 1891, and
Latirus karinae
Nowell-Usticke, 1969
. Most pieces of this taxonomic puzzle were examined by Bullock (1968), who recognized a species-group that included
Latirus attenuatus
(Reeve, 1847)
,
L. annulatus
(Melvill, 1891)
, and
L. virginensis
Abbott, 1958
, all occurring sympatrically in the West Indies. Bullock (1968) noted a “tremendous amount” of morphological variation among shells in these groups and also noted many intergrades that left relationships among the species unclear. Bullock proposed (in thesis) but did not formally introduce a new subgeneric name for the group, with
Turbinella attenuata
, a species we already discussed, as its
type
species. Bullock cited the range of what he called
L. attenuatus
as
Cuba
to the Lesser Antilles, reported records from
Cuba
,
the Dominican
Republic,
Puerto Rico
, and St. Thomas,
U. S.
Virgin Islands
, and figured specimens from
the Dominican
Republic,
Puerto Rico
, and St. Thomas, but we have seen no material from any of those locations that conforms to the
type
of
P. attenuatus
. Bullock (1968: 69) also cited
Latirus eppi
“Melvill and Schepman
” as a junior synonym of
L. attenuatus
, characterizing the
holotype
of
L. eppi
as a squat specimen quite unlike the
holotype
of
L. attenuatus
but noting that “a series of intergrades appear to exist between the two forms.” He concluded that further study may distinguish them as separate species. We provide an account for
Pustulatirus eppi
as a separate species-level taxon, but we consider shells assigned by Bullock to
L. attenuatus
(except its
holotype
and the
holotype
of
L. eppi
) to be more appropriately assigned to
Pustulatirus virginensis
.
The name
Latirus annulatus
(Melvill)
traces to a shell figured by Chemnitz (1780: pl. 141, fig. 1316). Chemnitz (1780) grouped the figure with several others (pl. 140, figs.
1306–1309
; pl. 141,
1314–1316
), all of which he associated with a (non-binominal) species-group. Gmelin (1791) later assigned all of the Chemnitz figures (and some by other authors) to his new species
Murex polygonus
. [For several decades thereafter, some authors (e.g., Bosc 1802, Dillwyn 1817, Wood 1818, Lamarck 1822, Anton 1838) indiscriminately repeated all of the Chemnitz figures that Gmelin had cited for
polygonus
, but those citations are not germane here.] Röding (1798) reallocated the Chemnitz figures cited by Gmelin as follows: (1)
M. polygonus
Gmelin
was restricted to figures
1306 and 1307
and reclassified as
Fusus polygonus
—now known as
Latirus polygonus
of the Indo-West Pacific; (2) figures
1308 and 1309
were reassigned to
Fusus tapetepersicum
Röding, 1798
—now a junior synonym of
Latirus gibbulus
(Gmelin, 1791)
, another Indo-West Pacific species and the
type
species of
Latirus
; (3) figures
1314 and 1315
were reassigned to
Fusus angulatus
Röding, 1798
—now
Polygona angulata
(Röding, 1798)
of the southern Caribbean region; and (4) Chemnitz figure 1316 was reassigned both to
Syrinx annulata
Röding, 1798
, and to
Syrinx clathrata
Röding, 1798
. Link (1807) reclassified
S. annulata
as
Cymatium annulatum
but did not mention
S. clathrata
. Lacking other criteria by which to differentiate them, the two Röding names are primary subjective synonyms. Regardless of which name is preferred (one may argue that Link, 1807 preferred
annulata
), we believe that Chemnitz figure 1316 is too poor to allow confident identification to species level, and both names should be considered
nomina dubia
.
Mørch (1852) first listed
Lathyrus annulatus
“Bolt.
” (i.e., Röding) among the West Indian fauna; H. and A.
Adams (1853) listed the name as
Latirus annulatus
(Bolten)
; and Krebs (1864) listed
Turbinella annulata
(Bolten)
among West Indian shells. Finally,
Peristernia annulata
(Bolten)
was listed by Melvill (1891b: 407) but refigured as
Peristernia annulata
(A. Ad.)
by Melvill (1891b: 411, fig. 1). Bullock (1968: 65) contended that Chemnitz pl. 141, fig. 1316, the only figure cited by Röding for
Syrinx annulata
, “could certainly be a
Latirus
, [but] it is impossible to determine the species as the illustration is poor and the aperture is not shown. Therefore, Röding's name should be considered a
nomen dubium
and the [name]
annulatus
credited to Melvill.”
Melvill (1891b) referred authorship for the name with his figured shell not to Bolten (Röding) but to Arthur Adams. According to Trew (1992), the only species that Adams named
annulatus
, -
a
, -
um
were in
Fissurellidae
and
Pyramidellidae
, families unlikely to be confused with
Fasciolariidae
. However, if Melvill’s reference to Adams as author was in error, publication by Melvill of the unused combination
Peristernia annulata
and the accompanying new figure constituted valid introduction of a name. Thus, the binomen
Peristernia annulata
“(A. Ad.)” Melvill (1891b) seems to be available for taxonomic purposes; it only remains to prove what that name represents. The
type
specimen must be the shell that Melvill figured, but the location of that specimen is unknown and its identity could not be confirmed. Trew (1990) reported a specimen of
annulatus
in the Melvill-Tomlin Collection at the National Museum of
Wales
. At our request, museum staff photographed that specimen, which proves to be a young
Hemipolygona carinifera
(Lamarck, 1816)
(
Figure 31
) quite unlike Melvill’s figure. We hesitate to replace the well-known
Latirus virginensis
Abbott
with the Melvill name. Absence of an original description and lack of access to a
type
render the status of
Peristernia annulata
uncertain. Until the figured
type
is found and its identity confirmed, we recommend that Melvill’s name be considered a
species inquirenda
.
Bullock (1968: 65-67) reported as
Latirus annulatus
(Melvill)
specimens from
Cuba
,
Jamaica
, St. Thomas and
Antigua
, figured by that name shells from
Jamaica
, St. Thomas and
Antigua
, and cited the species’ range as
Cuba
to the
Virgin Islands
and southward in the Lesser Antilles. Bullock characterized
L. annulatus
as “the most commonly found and least variable of the [species group].” He distinguished
L. annulatus
from
L. attenuatus
by its “more angular shoulder on the body whorl and by having white axial ribs on every whorl,” stating that
L. virginensis
differs from
L. annulatus
“by having a more rounded body whorl lacking the white color of the axial ribs.” Shells that Bullock figured as
L. annulatus
generally resemble Melvill’s figure, which in turn resembles shells that Nowell-Usticke (1969) named
Latirus karinae
from
Puerto Rico
, the
Virgin Islands
, and
Antigua
. Nowell-Usticke (1971) tacitly acknowledged synonymy of
L. karinae
with
L. virginensis
by captioning as
virginensis
in 1971 the figure he had used for
karinae
in 1969; his figured shell is now the
lectotype
of
karinae
(see Boyko and Cordiero, 2001).
Latirus karinae
was formally relegated to synonymy with
L. virginensis
by Faber (1988).
Bullock (1968: 71, 72) characterized
Latirus virginensis
as a very variable species whose status “will not be known until large series of specimens have been studied. The relationship between this species,
attenuatus
, and
annulatus
is very unclear. The
type
of
virginensis
is a mature specimen and seems different from the largest
annulatus
seen.” Bullock defined the range of
L. virginensis
as
Puerto Rico
and the
Virgin Islands
to
Brazil
, citing specimens examined from
Puerto Rico
(four lots) and St. Thomas (one lot). Bullock also cited his pl. 4, fig. 19 for
L. virginensis
, but its caption identifies that shell as
L. attenuatus
from St. Thomas.
Abbott (1958) defined the
type
series of
Latirus virginensis
as:
holotype
ANSP 196459; two
paratypes
ANSP 34975; two
paratypes
ANSP 34968; and four
paratypes
ANSP 34969, and we examined all of these specimens at ANSP. We also examined three other specimens (ANSP
210740
; ANSP 219064; UF 162199) labeled as
paratypes
of
Latirus virginensis
, but Abbott did not mention those specimens and they cannot be
type
material. Abbott (1958) only figured the
holotype
of
L. virginensis
, and Bullock’s thesis does not indicate that he examined the
type
series at ANSP, but Lyons (1991) illustrated the
holotype
and three
paratypes
from the series; those figures suggest that, had he seen them, Bullock would have classified the
holotype
and two
paratypes
(Lyons figs. 36–40) as
L. attenuatus
and one
paratype
(Lyons fig. 41) as
L. annulatus
, further exemplifying the confusing variability of shells in this species-group.
Simplistically, there are two dissimilar morphotypes in the group. The first
type
, a “
virginensis
” form, is typified by a mostly smooth shell with broad axial ribs, low to absent spiral cords, and a rather waxy shell surface. The shell color is usually olivaceous brown with lighter-colored ribs, but completely orange or yellow specimens are known, and some shells are mostly orange or yellow but with brown patches between ribs on posterior whorls. A population from Little St. James
Island
in the
U. S.
Virgin Islands
is uniformly olivaceous.
The other morphotype, a “
karinae
” form, is typified by robust, swollen ribs and well-defined spiral cords. The shell color is usually orange-brown but may be yellow, dark brown, or even black, with white axial ribs in stark contrast to the background color. Axial ribs are usually more developed on the
karinae
form than on the
virginensis
form and may occur on all whorls, but more often become subdued on the penultimate and body whorls of large specimens.
Shell sizes vary considerably among and between populations. Large shells (>
35 mm
sl) were examined from
the Dominican
Republic, St. Thomas and
St. John
in the
U. S.
Virgin Islands
,
Anguilla
,
St. Martin
,
St. Barthelemy
,
Antigua
,
Guadeloupe
, and
St. Lucia
. We saw no shells larger than
28 mm
from
Aruba
, and only two shells larger than
30 mm
from the
Bahama
Islands.
Finally, having examined nearly
400 specimens
, we could not distinguish one phenotype that could not be linked by intergrades to other quite different phenotypes. We conclude that all of the specimens represent a single, morphologically variable species for which
Pustulatirus virginensis
(Abbott, 1958)
is the first available name.
The shell that Kaicher (1986: card 4671) figured as
Latirus elegans
(Gray)
is in the ANSP collection (ANSP 35033;
Figure 23
), where it was catalogued as “
L. elegans
A. Ad. (?)” without stated locality. That specimen, actually
P. virginensis
, is much darker brown than indicated by Kaicher’s figure but is identifiable by a chip in the abapical portion of the outer lip that is visible on the specimen and in the figure. It differs from the shell Kaicher figured on the same card as the
holotype
of
Fusus elegans
Gray, 1838
, which has a shorter, more canted siphonal process, relatively shorter and broader axial ribs, and distinct threads between spiral cords. We doubt that the shell Kaicher figured as
holotype
of
Fusus elegans
Gray
(NHMUK 1968453) is the shell that Gray described from
Sierra Leone
; his description includes the phrase “with … rather distant, acute, raised, narrow, brown topped spiral ridges,” a condition that fits perfectly shells of
Fusinus filosus
(Schubert and Wagner, 1829)
from West Africa but not evident on the NHMUK shell, and we are inclined to treat
F. elegans
Gray
as a junior subjective synonym of
Fusus filosus
Schubert and Wagner.
A shell from off Fortaleza, Ceará,
Brazil
that Bullock (1968) reported and figured as
Latirus virginensis
is instead a new Brazilian endemic which we describe as
Pustulatirus biocellatus
n. sp.
in an account that follows. The shell figured as
Latirus virginensis
from the Bay Islands of
Honduras
by K. and L. Sunderland (1996) is
Pustulatirus utilaensis
n. sp.
, described next.