The Veliger 51(1):85-103 (March 31, 2010)
THE VELIGER
? CMS, Inc., 2008
Trophonella (Gastropoda: Muricidae), a New Genus from Antarctic
Waters, with the Description of a New Species
M. G. HARASEWYCH
Department of Invertebrate Zoology, MRC-163, National Museum of Natural History, Smithsonian Institution,
P.O. Box 37012, Washington, DC 20013-7012, USA
(e-mail: Harasewych@si.edu)
GUIDO PASTORINO
Museo Argentino de Ciencias Natureles, Av. Angel Gallardo 470, 3? piso, lab. 57,
C1405DJR Buenos Aires, Argentina
(e-mail: gpastorino@macn.gov.ar)
Abstract. The new genus Trophonella is described from the outer shelf and upper continental slope of Antarctica
and islands within the Antarctic Convergence. Four previously known species that had been attributed to the genus
Trophon (Trophon scotianus Powell, 1951; T. echinolamellatus Powell, 1951; T. enderbyensis Powell, 1958; and T.
eversoni Houart, 1997) are included in Trophonella, as is one new species (Trophonella rugosolamellata) described
herein. Trophonella resembles Trophon in gross shell morphology: the members of both genera have large, globose
shells, paucispiral protoconchs, prominent axial lamellae, and short siphonal canals. Trophonella differs from
Trophon in having shells with evenly rounded whorls that lack a well-defined shoulder; rachidian teeth with
distinctive, broadly triangular central cusps, but that lack the marginal cusps of Trophon; characteristic spherical
accessory salivary glands; and a circumpapillar fold on the penis that is absent in Trophon. Relationships of the
genera Trophon and Trophonella, as well as of the subfamily Trophoninae are reexamined by supplementing the data
matrix of Kool (1993b, Table 3) with data for additional taxa. Results support the segregation of Trophonella from
Trophon at the generic level. Based on the relationships of the type species of their respective nominotypical genera,
Trophoninae is either the sister taxon of a narrowly circumscribed Ocenebrinae, or both are part of a larger clade. A
better resolved phylogeny containing a much broader sampling of the more than 50 genus-level taxa that have been
attributed to these two subfamilies will be required in order to delineate more precisely the membership of the clade
and to identify its diagnostic synapomorphies.
INTRODUCTION
Although the family Muricidae is best known for its
temperate and tropical shallow-water representatives, it
also includes a group of taxa inhabiting polar seas and
intervening deeper waters that have generally been
grouped in the subfamily Trophoninae. Cossmann
(1903) proposed the subfamily Trophoninae to contain
the genera Trophon Montfort, 1810, and Aspella
March, 1877. The genus Trophon was subdivided into
the subgenera Trophon and Trophonopsis Bucquoy and
Dautzenberg, 1882. Under Trophon sensu stricto, he
included the sections Trophon, Xanthochorus Fischer,
1884, and Forreria Jouseaume, 1880. Trophonopsis was
further subdivided into Trophonopsis and Boreotrophon
Fischer, 1884. The genus Aspella was not subdivided.
Over the past several decades, there has been general
agreement among researchers that Trophoninae is not
a monophyletic group. As was noted by Radwin and
D'Attilio (1976, p. 175), Trophoninae "has been the
site of assignment of many dissimilar forms that seem,
on the basis of the shell and radular morphology of the
type species of Trophon (T. geversianus), unlikely to
belong there." The genus Aspella is presently assigned
to the subfamily Muricinae on the basis of radular
morphology (Radwin & D'Attilio, 1976, p. 21). These
authors went on to exclude from the subfamily
Trophoninae a number of genera (e.g., Austrotrophon
Dall, 1902; Forreria; Zacatrophon Hertlein & Strong,
1951) because they" ... had radulae characteristic of the
family Thaididae." In a phylogenetic study, Kool
(l993a, fig. 65) showed that, based on the morphology
of its type species, the genus Trophon was closely
related to the genera Nucella and Ocenebra, and he
suggested that future studies would reveal Trophoninae
to be polyphyletic.
Our continuing studies on the Muricidae represented
in the collections of the United States Antarctic
Program (USAP) have revealed that four species
previously described in the genus Trophon (i.e.,
Page 86
Trophon scotianus Powell, 1951; T echinolamellatus
Powell, 1951; T enderbyensis Powell, 1958; and T
eversoni Houart, 1997) appear to be closely related to
each other, but they differ substantially from the type
species of the genus Trophon. In this paper, we propose
the new genus Trophonella to include these species, and
we describe an additional new species that we attribute
to this genus. A review of each of the species included
in this new genus is provided, based on more recently
collected specimens.
We also examine the relationships of the genus
Trophon and the subfamily Trophoninae by reanalyz?
ing Kool's (1993b, table 3) data matrix of primarily
ocenebrine and rapanine taxa supplemented by the
addition of data for (1) Trophon geversianus (Pallas,
1769), the type species of the type genus of Trophoni?
nae; (2) Ocenebra erinaceus (Linnaeus, 1758), the type
species of the type genus of Ocenebrinae; (3) Tropho?
nella scotiana (Powell, 1951), the type species of the new
genus Trophonella; (4) Boreotrophon aculeatus Watson,
1882; and (5) Paziella pazi (Crosse, 1869), the type
species of the genus Paziella.
MATERIALS AND METHODS
This study is based primarily on specimens collected for
the United States Antarctic Program (USAP) aboard the
vessels RN Hero, RN Eltanin and RN Professor
Sedlacki and housed in the collections of the National
Museum of Natural History, Smithsonian Institution
(USNM). Additional specimens from several Antarctic
expeditions of the Argentine Republic deposited in the
Museo Argentino de Ciencias Naturales (MACN) were
examined, as were specimens at the Zoological Institute
and Museum, Hamburg (ZMH), and the Senckenberg
Museum, Frankfurt (SMF), collected by the vessels RN
Walther Herwig and RN Polarstern. The notation "win"
following an institutional acronym indicates that no
catalog number was associated with the specimen at the
time it was examined. All specimens were compared with
the holotypes oftheir respective species, which are housed
in The Natural History Museum, London (NHM), the
South Australian Museum (SAM), and at USNM.
Alcohol-preserved animals of most species of Tropho?
nella were dissected, and the gross anatomy of the
anterior part of the alimentary system and pallial
gonoducts were compared. Radulae were prepared
according to the method described by Solem (1972)
and examined using a Scanning Electron Microscope
(SEM). Terminology for rachidian-tooth morphology
follows Kool (1987, fig. 1). Shell ultrastructure data was
obtained from fracture surfaces of shell fragments
removed from the central portion of the lip along the
last shell whorl.
In order to determine the generic and subfamilial
relationships of this distinctive group of Antarctic
The Veliger, Vol. 51, No.1
muricids, we reanalyzed the data matrix in Kool
(1993b, table 3), to which we added data for Tro?
phon geversianus, Ocenebra erinaceus, Trophonella scoti?
ana, Boreotrophon aculeatus, and Paziella pazi. Harase?
wych (1984) was the source ofdata for T geversianus, B.
aculeatus, and P. pazi. Data for T geversianus and o.
erinaceus were obtained from Kool (1993a). Data on
shell ultrastructure of Boreotrophon aculeatus and
Paziella pazi were newly obtained from the voucher
material from Harasewych's (1984) study. Each of the
newly added taxa were scored for the 18 characters and
their character states, as defined in Kool (1993b). The
data were analyzed using maximum parsimony (MP)
and neighbor-joining algorithms ofPAUP* 4.0 Macin?
tosh Beta Version 10 (Swofford, 2002).
SYSTEMATICS
Class GASTROPODA Cuvier, 1797
Order NEOGASTROPODA Wenz, 1938
Family MURICIDAE Rafinesque, 1815
Trophonella new genus
Type species: Trophon scotianus Powell, 1951.
Diagnosis: Shell large, fusiform, inflated. Protoconch
paucispiral (1.5-2 whorls). Teleoconch (6-7 whorls)
globose, without distinct shoulder, with tall, conical
spire. Sculpture of low, rounded, spiral cords and axial
lamellae that may be broadly flaring to short and
rugose. Aperture broadly rounded, with flaring outer
lip and thin inner lip. Siphonal canal short. Operculum
large, oval, with subterminal nucleus. Rachidian teeth
with rectangular basal plate; three major cusps, central
cusp exceptionally broad, triangular, with small denticle
on each side; marginal cusps absent. Accessory salivary
glands small, spherical. Esophageal gland reduced.
Penis with conical papilla surrounded by collar.
Description: Shell large (to 76 mm), fusiform, often
broadly ovate, with strongly convex, evenly rounded
whorls that do not form a distinct shoulder. Proto?
conch paucispiral, occasionally with weak, irregular,
spiral cords. Teleoconch sculpture of low, rounded,
spiral cords with or without scales, and axial lamellae
that range from broadly flaring to short and rugose.
Aperture large, suboval, with thick outer lip, and thin
inner lip. Siphonal canal short, scabrous. Shell
ultrastructure of three layers, the inner two of
orthogonally oriented crossed-lamellar aragonite, the
outermost layer of calcite. Operculum large, filling
aperture, corneous, oval, with subterminal nucleus,
thick rim, and large attachment area.
Animal large, unpigmented. Cephalic tentacles wide,
blunt. Mantle edge smooth, siphon short. Pleuroem-
M. G. Harasewych & G. Pastorino, 2008
bolic proboscis short, broad. Radular ribbon small,
thin, extending beyond rear of buccal mass. Rachidian
teeth with subrectangular, weakly recurved basal plate
and smooth, broad, marginal surfaces that lack
marginal cusps. Central cusp long, exceptionally broad,
triangular; flanked by a single, small to nearly obsolete
lateral denticle on each side; lateral cusps short, robust,
smooth. Lateral teeth of moderate size, basal plate
shorter and narrower than in rachidian teeth, with
single scythelike cusp along outer edge that is slightly
longer than the basal plate. Esophagus short. Salivary
glands very large, ascinous, enveloping esophagus.
Accessory salivary glands small, spherical, and com?
pletely embedded in corresponding salivary gland.
Gland of Leiblein compact, with terminal ampula.
Penis large, wide, with conical papilla rising from a
concavity at its distal end.
Etymology: Trophon (Gr., that which feeds) + ella (L.,
suffix added to form the diminutive).
Included species: Trophonella scotiana (Powell, 1951),
new combination [Type species]; T echinolamellata
(Powell, 1951), new combination; T eversoni (Houart,
1997), new combination; T enderbyensis (Powell,
1958), new combination; T rugosolamellata, new
species.
Remarks: Trophonella is similar to Trophon in gross
shell morphology, as both have globose shells of
comparable size with paucispiral protoconchs, promi?
nent axial lamellae, and short siphonal canals. Species
of Trophonella can be distinguished most readily by
their evenly rounded whorls that lack a well-defined
shoulder. The main differences appear to be anatom?
ical. Trophonella has a circumpapillar fold on the penis
that is absent in Trophon. The accessory salivary glands
of Trohonella are globular, while those of Trophon are
elongate.
The rachidian teeth of Trophonella have distinctively
broadly triangular central cusps and lack the marginal
cusps of Trophon.
Trophonella is restricted in geographic distribution to
Antarctica and islands within the Antarctic Conver?
gence. Records range in depth from 18 m to 474 m
(both for T scotiana), with the majority of specimens
sampled at outer-shelf and upper continental-slope
depths (Figures 20-21). In terms of size and biomass,
species of Trophonella are the largest muricids occur?
ring within the Antarctic Convergence.
Trophonella scotiana (Powell, 1951),
new combination
(Figures 1-13, 20)
Synonymy: Trophon scotianus Powell, 1951, p. 153, pI.
9, figs. 48--49, M88; Carcelles, 1953, p. 189, pI. 2, fig.
Page 87
51; Powell, 1960, p. 154; Okutani, 1986, p. 279, pI. 1,
figs. 3-6; Dell, 1990, p. 207, figs. 346-347; Castellanos
& Landoni, 1993, p. 13, fig. 36; Numanami, 1996,
p. 138, figs. 91 a-c.
Trophon sp. 1 Hain, 1990, p. 63, pI. 6, fig. 9 a,b; pI. 25,
fig. 2.
Description: Shell (Figures 1-6) large (to 70 mm),
fusiform, thin shelled, chalky. Protoconch (Figures 7?
10) tall, increasing in diameter from 450 Ilm to 1.2 mm
in 1112 to P/4 evenly rounded whorls, surface with very
fine, irregular and discontinuous spiral threads. Tran?
sition to teleoconch marked by broadly flaring lip.
Teleoconch of up to 6 slightly globose whorls; spire tall,
conical, less than 50% of shell length. Suture impressed
to abutting. Spire angle 62?-84?. Aperture subpolygo?
nal to evenly rounded, deflected from the coiling axis
by 20?-25?; interior glossy. Siphonal canal moderately
short, narrow, slightly deflected dorsally. Siphonal
fasciole scabrous; pseudoumbilical chink nearly closed
or reduced to a very narrow slit. Outer lip rounded with
reflected edges. Columellar lip narrow, adpressed.
Axial sculpture of regular, thin, strongly developed
lamellae (4-11 on last whorl) that span whorl surface
from suture to siphon, expanded in shoulder region.
Fine, closely spaced growth lines cover whorls and
lamellae. Spiral sculpture, of low, rounded, spiral
cords, broader than interspaces (18-32 on body whorl,
2-6 on siphonal canal) becoming reduced to obsolete
on larger specimens. Shell color white to pale orange?
tan. When color present, lighter bands may be evident,
one below suture, the other above the juncture to the
siphonal canal.
Shell composed of three layers (Figure 11). Inner?
most layer, of crossed lamellar aragonite with crystal
planes oriented perpendicular to the growing edge,
comprises roughly 2% of shell thickness. Middle layer,
also aragonitic, arranged with crystal planes parallel to
growing edge, comprises roughly 36% of shell thick?
ness. Outermost layer is calcitic, comprising the major
part (62%) of the shell's thickness. Operculum (Fig?
ure 43) oval, with terminal nucleus, outer surface with
regular growth lines, inner surface with broad, thickly
glazed rim along posterior edge, and large, round/oval
attachment area, often with 2-3 horseshoe-shaped
scars.
Animal large, compact. Foot short, broad, with
accessory boring organ situated along ventral midline,
opening via ventral pedal gland in females. Tentacles
large, broad, closely situated, with large black eyes.
Mantle edge smooth, osphradium (Figure 48, os)
small, slightly asymmetrical, with 45-55 leaflets per
side, less than 50% the length of the ctenidium
(Figure 48, ct). Proboscis large, muscular, cylindrical,
pleurombolic. Radula (Figures 12-13) short (radular
Page 88 The Veliger, Vol. 51, No.1
Figures I-II. Trophonella scotiana (Powell, 1951). 1. Apertural, 2. lateral, and 3. dorsal views of a medium-sized specimen with
few axial lamellae, USNM 897411, off South Georgia Island, 54?53'S, 35?49'W, in 127-140 m. 4. Apertural view of large specimen
with well-developed lamellae, USNM 901644, Off Borchgrevink Coast, Victoria Land, Antarctica, 74?01 'S, 178?53'E, in 256?
258 m. 5. Apertural and 6. dorsal views of average specimen, USNM 897560, off South Georgia Island, 54?39'S, 37?22'W, in 140?
150 m. 7-8. Lateral, and 9. apical views of the protoconch. 10. Detail of protoconch sculpture. USNM 901645, W of Graham Land,
M. G. Harasewych & G. Pastorino, 2008
length roughly 30% of aperture length), narrow
(roughly 550 11m wide), extending beyond rear of
retracted proboscis. Rachidian tooth subrectangular,
basal plate moderately concave anteriorly, with short,
smooth marginal area. Central cusp large, broad,
lateral cusps shorter, narrower, with single denticle
between central and lateral cusps (Figure 13, d) often
almost obsolete, but present, even in juvenile speci?
mens. Lateral teeth L-shaped, with single cusp along
outer edge roughly equal in length to width of basal
plate. Juvenile specimens have proportionally larger
denticles, and thinner cusps on rachidian and lateral
teeth. Esophagus very thin. Valve of Leiblein (Fig?
ure 49, vL) ovate. Salivary glands (Figure 49, sg) very
large, envelop the valve of Leiblein and the small,
nearly spherical accessory salivary glands, joining
esophagus via two thick ducts anterior to the valve of
Leiblein. Mid-esophagus very thin, lacks conspicuous
esophageal gland, passes under gland of Leiblein.
Gland of Leiblein large, compact, with blind duct
terminating posteriorly in large ampulla. Stomach
(Figures 50, 51, sto) large, lining the anterior and
dorsal surface of the digestive gland, into which it
opens via ducts (Figure 51, dd). Intestine (Figures 50,
51, i) broad. Penis (Figures 45--47, p) large (> 4 X
tentacle length), broad, dorsoventrally flattened, elon?
gate, with laterally situated sperm duct, dorsal blood
sinus, and conical papilla (Figures 46, 47, 52, pp)
surrounded by a collar. Pallial oviduct broad. Albumin
gland thin, capsule gland (Figure 53, cg) short, globose.
Bursa copulatrix (Figure 53, bc) onion-shaped, with
distal female opening (Figure 53, fo). Rectum (Fig?
ure 53, r) overlays pallial oviduct; rectal gland (Fig?
ure 53, rg) lines anterior dorsal surface of rectum; anus
(Figure 53, a) adjacent to female opening. Egg capsules
of T scotiana lenticular, subcircular in outline,
attached to substrate (Hain, 1990, pI. 6, fig. 9b,
Trophon sp. I). Each capsule is about 19 mm in
diameter, contains 140 eggs that hatch as crawling
juveniles in 24-25 months (Hain & Arnaud, 1992,
p. 307, table 3).
Type locality: Off South Georgia Island, 53?55'S,
38?01'W, in 107 m (Figure 20,0).
Type material: Holotype, NHM 1961542.
Material examined: D indicates empty shells, L
indicates live-collected specimens.
NHM 1961542, Holotype; USNM 887834, 1 L, off
South Georgia Island, 53?50'36"S, 36?18'36"W, 185?
205 m, R/V Islas Orcadas, cr. 575, stn. 30, 19 May
Page 89
1975; USNM 887835, 1 L, off South Georgia Island,
54?01'18"S, 36?50'42"W, 108-119 m, R/V Islas Orca?
das, cr. 575, stn. 24, 17 May 1975; USNM 896099,4 D,
off South Georgia Island, 53?51 'S, 37?38'W, in 97?
101 m, R/V Eltanin cr. 22, stn. 1535, 7 Feb 1966;
USNM 896982, 1 L, off South Georgia Island, 53?57'S,
36?06'W, 151-158 m, R/V Professor Siedlecki, cr. 601,
stn. 97, 13 Dec 1986; USNM 897411, 2 L, off South
Georgia Island, 54?53'S, 35?49'W, in 127-140 m, R/V
Professor Siedlecki, cr. 601, stn. 71, 9 Dec 1986; USNM
897416, 1 L, off South Georgia Island, 55?24'S,
35?22'W, 207-218 m, R/V Professor Siedlecki, cr.
601, stn. 60, 8 Dec 1986; USNM 897422, 1 L, off
South Georgia Island, 55?08'S, 35?06'W, 145-153 m,
R/V Professor Siedlecki, cr. 601, stn. 66, 9 Dec 1986;
USNM 897427, 2 L, off South Georgia Island, 55?0I'S,
35?27'W, 117-122 m, R/V Professor Siedlecki, cr. 601,
stn. 67, 9 Dec 1986; USNM 897433, 2 L, off South
Georgia Island, 55?05'S, 35?23'W, 116-121 m, R/V
Professor Siedlecki, cr. 601, stn. 68,9 Dec 1986; USNM
897434, 5 L, off South Georgia Island, 54?31 'S,
38?11'W, 180-190 m, R/V Professor Siedlecki, cr.
601, stn. 36, 4 Dec 1986; USNM 897444, 1 L, off
South Georgia Island, 53?55'S, 37?08'W, 104-122 m,
R/V Professor Siedlecki, cr. 601, stn. 109, 14 Dec 1986;
USNM 897459, 1 L, off South Georgia Island, 53?52'S,
37?33'W, 104-112 m, R/V Professor Siedlecki, cr. 601,
stn. 111, 15 Dec 1986; USNM 897475, 3 L, off South
Georgia Island, 53?52'S, 38?20'W, 113-120 m, R/V
Professor Siedlecki, cr. 601, stn. 120, 16 Dec 1986;
USNM 897490,1 L, off South Georgia Island, 54?11'S,
37?53'W, 113-118 m, R/V Professor Siedlecki, cr. 601,
stn. 40, 5 Dec 1986; USNM 897521, 2 L, off South
Georgia Island, 54?05'S, 38?25'W, 197-207 m, R/V
Professor Siedlecki, cr. 601, stn. 24, 3 Dec 1986; USNM
897560, 1 L, off South Georgia Island, 54?39'S,
37?22'W, 140-150 m, R/V Professor Siedlecki, cr.
601, stn. 46, 6 Dec 1986; USNM 897564, 5 L, off
South Georgia Island, 54?51 'S, 35?38'W, in 84-103 m,
R/V Professor Siedlecki, cr. 601, stn. 72, 9 Dec 1986;
USNM 901642,1 D, Ross Sea, Moubray Pennell Bank,
Victoria Land, Antarctica, 73?22'S, 177?37'E, in 465?
474 m, R/V Eltanin, cr. 27, stn. 1933, 30 Jan 1967;
USNM 901644, 1 L, Ross Sea, Moubray Pennell Bank,
Victoria Land, Antarctica, 74?01'S, 178?53'E, in 256?
258 m, R/V Eltanin, cr. 32, stn. 2018, 14 Jan 1968;
USNM 901645, 1 L, W of Graham Land, Palmer
Peninsula, Antarctica, 66?21.7'S, 66?47'W, in 70?
106 m, R/V Hero, cr. 731, stn. 1861, 1 Mar 1973;
MACN-In 18991, 1 D, Schlieper Bay, South Georgia
Island, in 18 m; MACN-In 18990, 1 L, Cumberland
Palmer Peninsula, Antarctica, 66?21.7'S, 66?47'W, in 70-106 m. Scale bars = 400 ~m. II. Shell ultrastructure. Fracture surface
parallel to growing edge. USNM 887835, off South Georgia Island, 54?01'18"S, 36?50'42"W, 108-119 m.
Page 90 The Veliger, Vol. 51, No.1
Figures 12-19. Radulae of species of Trophonella. 12. Dorsal and 13. right lateral views of the radular ribbon of T scotiana,
specimen in Figure 4.14. Dorsal and IS. right lateral views of the radular ribbon of T echinolamellata, USNM 896041, 2L, NW of
Brabant Island, Palmer Archipelago, Antarctic Peninsula, 63?51 'S, 62?38'W, in 128-165 m. 16. Dorsal and 17. right lateral views of
the radula of T enderbyensis, specimen in Figures 38-40. 18. Dorsal and 19. right lateral views of the radula of the holotype of T
rugosolamellata n. sp. ZMH 2777.
M. G. Harasewych & G. Pastorino, 2008 Page 91
2010o
Om
50m
200m
150m
50 m t----t------t-----j
Om ....,...-.,......,
o 2 3
500 m _ ................._"--....1
T scotianus (.)
type locality (0)
400 m r---+--+---+-+--I
250 m
100m
350 m til"'-+--+---+-+--I
300m
150 m t----t------t-----j
200m
250 m
400m ................. -
T rugosolamellata (ffi)
T eversoni (.)
type locality & depth
unknown
450 m Ir-+--+---+-+--I
350 m t--+---+--I
300 m t--t---t-----j
100 m t--t---t-----j
Figure 20. Geographic and bathymetric distributions of Trophonella scotiana, T rugosolamellata n. sp., and the only locality
reported for T eversoni.
Bay, South Georgia Island, in 36 m; ZMH win, 1 D,
off South Georgia Island, 54?13.2'S, 37?49.6'W, in
122 m, RN Walther Herwig, cr. 68, stn. 15; ZMH win,
I L, off South Georgia Island, 53?55.9'S, 38?29.6'W, in
145 m, RN Walther Herwig, cr. 68, stn. 19; ZMH win,
3 L, I D, off South Georgia Island, 53?39.5'S,
37?8.9'W, in 155 m, RN Walther Herwig, cr. 68, stn.
30; ZMH win, 1 D, off South Georgia Island,
53?47.3'S, 3r12.2'W, in 141 m, RN Walther Herwig,
cr. 68, stn. 31; ZMH win, 3 L, off South Georgia
Island, 53?59.8'S, 36?56.2'W, in 150 m, RN Walther
Herwig, cr. 68, stn. 35; ZMH win, I L, off South
Georgia Island, 53?43.3'S, 36?26'W, in 230 m, RN
Walther Herwig, cr. 68, stn. 40; ZMH win, 1 L, off
South Georgia Island, 53?59.1'S, 36?20.6'W, in 198 m,
RN Walther Herwig, cr. 68, stn. 44; ZMH win, 1 L, off
South Georgia Island, 53?52.9'S, 36?4.2'W, in 235 m,
RN Walther Herwig, cr. 68, stn. 55; ZMH win, 4 L, off
South Georgia Island, 54?32.6'S, 35?58.1 'W, in 155 m,
RN Walther Herwig, cr. 68, stn. 64; ZMH win, I L, off
South Georgia Island, 54?48'S, 35?23.6'W, in 215 m,
RN Walther Herwig, cr. 68, stn. 70; ZMH win, 4 L, off
South Georgia Island, 55?15.2'S, 34?46.8'W, in 240 m,
RN Walther Herwig, cr. 68, stn. 78; ZMH win, 2 L, off
South Georgia Island, 55?0.37'S, 35?03.8'W, in 124 m,
RN Walther Herwig, cr. 68, stn. 79; SMF win, I D, off
the Princess Martha Coast, Queen Maud Land,
Antarctica, 700 29'S, 8?07'W, in 270-303 m.
Literature records: PS ANT V/3 St. 593, 73?5S'S,
23?38'W, in 330 m (Hain 1990); JAR?-25, St. A,
70? 14'S, 24?23.9'?, Breid Bay, in 310 m (Okutani 1986;
Numanami 1996).
Distribution: Off South Georgia Island, the Antarctic
Peninsula, and the Ross Sea, the Weddell Sea, and
Breid Bay, Antarctica, at depths of 18-474 m. See
Figure 20.
Page 92
Remarks: This distinctive taxon, the largest species of
Trophonella in terms of biomass, had, until recently,
been known from less than a dozen specimens (Dell,
1990:208; Numanami, 1996:138). A survey of the
collections of Antarctic mollusks at USNM, SMF
and ZMH, as well as a search of the literature, has
uncovered more than 60 specimens and/or records,
most from the vicinity of South Georgia Island and the
Weddell Sea. Dell (1990:208) and Numanami
(1996:140) both noted variation in shell morphology
of this species, with the degree of whorl inflation and
the number of axial lamellae per whorl both increased
in large specimens.
Trophonella echinolamellata (Powell, 1951)
new combination
(Figures 14, 15, and 21-28)
Synonyms: Trophon echinolamellatus Powell, 1951: 152,
pI. 9, figs. 44, 45. Trophon equinolamellatus [sic]
Pastorino, 2002:fig. 22.
Description: Shell (Figures 22-24) large (to 69.8 mm),
fusiform, inflated, thin shelled, with scabrous surface.
Protoconch (Figures 25-27) tall, increasing in diameter
from 300 to 920 J..lm in 13/4 to 2 evenly rounded whorls,
surface smooth, frequently pitted. Transition to tele?
oconch distinguished by weak flare in lip. Teleoconch
of up to 7 evenly rounded whorls; spire tall, conical,
slightly less than half the shell length. Suture distinctly
impressed. Spire angle 66-72?. Aperture broadly ovate,
deflected from the coiling axis by about 25?, interior
with glazed, lustrous white layer that does not extend to
the end of the flared, rounded, outer lip, nor to the edge
of thick, adpressed, columellar lip. Demarcation
between apertural glaze and apertural lip very pro?
nounced, especially in pigmented shells. Siphonal canal
short, narrow, slightly deflected dorsally. Siphonal
fasciole scabrous; pseudoumbilical chink pronounced,
constricted to varying degrees by thick columellar lip.
Outer lip rounded, weakly reflected. Columellar lip
thick, overlaying sculpture of previous whorl. Axial
sculpture of numerous (14-16 on first teleoconch
whorl, 42-50 on last whorl), extremely short lamellae
that intersect with closely spaced spiral cords that are
as broad as interspaces (16-20 on body whorl, 2-4 on
siphonal canal) to produce a scaly surface. On the final
whorl of very large specimens, several lamellae may be
close together and thickened, giving the appearance of
varices that may be spaced 1/4 to 111 whorl apart. Shell
white, or, more commonly pale orange-cinnamon.
When color is present, lighter bands may be evident,
one below the suture, the other above the juncture to
the siphonal canal.
The shell is composed of three layers (Figure 28).
Innermost layer very thin (approximately 2% of shell
The Veliger, Vol. 51, No.1
thickness), consisting of crossed lamellar aragonite with
crystal planes oriented perpendicular to growing edge
of the shell. Middle layer of crossed lamellar aragonite,
with crystal planes colabrally oriented, accounts for
approximately 30% of shell thickness. Outermost layer
thickest (- 68% of shell thickness), calcitic. Operculum
(Figure 42) D-shaped, with straight adaxial margin,
terminal nucleus, fine growth lines along external
surface, and broad, glazed rim along inner surface.
Attachment area without scars.
The animal is similar to that of T. scotiana in most
respects. The radula (Figures 14, 15) is slightly longer
(radular length ~ 0.45 aperture length) and has
proportionally broader rachidian teeth with more
pronounced denticles.
Type locality: Off Cape Bowles, Clarence Island,
Antarctica, 61 ?25'S, 53?46'W, in 342 m. (Figure 21, "7).
Type material: Holotype and one paratype, NHM
1961541.
Material examined: D indicates empty shells, L
indicates live collected specimens.
NHM 1961541, Holotype and one paratype; USNM
638874, 1 L, off Victor Hugo Island, west coast of
Palmer Peninsula, Antarctica, 65?08'S, 66?04'W, in
135 m, stn. ED28, 22 Mar 1959; USNM 678397, 3 L,
off Palmer Peninsula, Antarctica, 600 48/S, 44?13.5W,
in 188 m, RN Eastwind, stn. EW66-028, 11 Feb 1966;
USNM 846180, 1 D, east of Victor Hugo Island, west
coast of Palmer Peninsula, Antarctica, 65?04/S,
65?53'W, in 150 m, RN Polar Duke, 8 Sept 1985;
USNM 870316, 1 L, NW of Brabant Island, Palmer
Archipelago, Antarctic Peninsula, 63?51 'S, 62?38'W,
128-165 m, RN Eltanin, cr. 6, stn. 439, 9 Jan 1963;
USNM 870593, 2 L, NE of Joinville Island, Antarctic
Peninsula, 62?41/S, 54?43'W, 210-220 m, RN Eltanin,
cr. 12, stn. 1003, 15 Mar 1964; USNM 881728, I L, off
Visokoi Island, South Sandwich Islands, 56?42/18"S,
27?00/24"W, 93-121 m, RN Islas Orcadas, cr. 575, stn.
61,30 May 1975; USNM 881906,1 L, D'Urville Island,
Bransfield Strait, Antarctic Peninsula, 62?39'S,
56?10'W, 426-311 m, RN Eltanin, cr. 6, stn. 418, 2
Jan 1963; USNM 881907, 1 L, D'Urville Island,
Bransfield Strait, Antarctic Peninsula, 62?39'S,
56?10'W, 426-311 m, RN Eltanin, cr. 6, stn. 418, 2
Jan 1963; USNM881939, 1 L 2 D, off South Georgia
Island, 54?41 'S, 38?38/W, 220-320 m, RN Eltanin, cr.
9, stn. 671, 23 Aug 1963; USNM 896041, 2 L, NW of
Brabant Island, Palmer Archipelago, Antarctic Penin?
sula, 63?5l/S, 62?38/W, in 128-165 m, RN Eltanin, cr.
6, stn. 439, 9 Jan 1963; USNM 896056, 2 L, E of South
Orkney Islands, 600 51'S, 42?57'W, in 155 m, RN
Eltanin, cr. 12, stn. 1083, 14 Apr 1964; USNM 901636,
2 D, off Zavodovski Island, South Sandwich Islands,
56?28.8'S, 27?24.6'W, in 161-210 m, RN Islas Orca-
M. G. Harasewych & G. Pastorino, 2008 Page 93
2010o
50 m
Om
200m
150m
400 m
250 m
350m
100m
300 m
450m
500m
T. echinolamellatus (T)
type locality (\7)
O'
I
0 2 3
Om
50m
100m
150m
200m
250m
300m
350m
ISO. 400m
T. enderbyensis (.)
type locality (0)
Figure 21. Geographic and bathymetric distributions of Trophonella echinolamellatus and T enderbyensis.
das, cr. 575, stn. 70, 2 lun 1975; USNM 901638, 1 L,
Gibbs Island, Bransfield Strait, South Shetland Islands,
Antarctica, 61?2YS, 56?30'W, in 300 ill, RN Eltanin,
cr. 12, stn. 998, 14 Mar 1964; USNM 901639, 1 L, Low
Island, South Shetland Islands, Antarctica, 63?26'S,
62?15'W, in 119-124 ill, RN Hero, cr. 691, stn. 26, 10
Feb 1969; USNM 901640, 1 L, Aspland Island, South
Shetland Islands, Antarctica, 61?1TS, 56?26'W, in
421--462 ill, RN Polarstern, cr. 2, stn. 22, 21 Nov
1996; USNM 901641, 4 L, Elephant Island, South
Shetland Islands, Antarctica, 6Oo 53'S, 55?32'W, in
178-120 ill, RN Polarstern, cr. 2, stn. 79, 9 Dec 1996;
col. P. Arnaud, 200-220 ill, South Shetland Islands,
Antarctica; ZMH wIn, 1 L, NW of Elephant Island,
South Shetland Islands, Antarctica, 600 56'S,
55?32.9'W 81 ill, RN Walther Herwig, cr. 68, stn. 68;
ZMH wIn, 2 L, E of Laurie Island, South Orkney
Islands, Antarctica, 600 48.9'S, 43?34'W 257 ill, RN
Walther Herwig, cr. 68, stn. 87; ZMH wIn, 2 L 2 D, E
of Laurie Island, South Orkney Islands, Antarctica,
600 41.6'S, 43?57.1'W 290 ill, RN Walther Herwig, cr.
68, stn. 88; ZMH wIn, 1 D, NNE of Laurie Island,
South Orkney Islands, Antarctica, 600 34.9'S,
44?17.3'W 240 ill, RN Walther Herwig, cr. 68, stn.
89; ZMH wIn, 4 L, 2 D, SE of Laurie Island, South
Orkney Islands, Antarctica, 6Oo51.3'S, 44?12'W 178 ill,
RN Walther Herwig, cr. 68, stn. 90; ZMH wIn, 3 L,
1 D, WNW of Coronation Island, South Orkney
Islands, Antarctica, 600 25'S, 46?25.7'W 150 ill, RN
Walther Herwig, cr. 68, stn. 118; ZMH wIn, lL, NW of
Aspland Island, South Shetland Islands, Antarctica,
61?21.8'S, 56?0.6'W 368 ill, RN Walther Herwig, cr.
68, stn. 138; ZMH wIn, 1 L, 3 D, NW of Aspland
Island, South Shetland Islands, Antarctica, 61?19.8'S,
56?9.9'W 328 ill, RN Walther Herwig, cr. 68, stn. 139;
ZMH wIn, 1 L, 1 D, NW of Aspland Island, South
Shetland Islands, Antarctica, 61?12.5'S, 56?23.4'W
460 ill, RN Walther Herwig, cr. 68, stn. 141; ZMH
Page 94 The Ve1iger, Vol. 51, No.1
Figures 22-28. Trophonella echinolamellata (Powell, 1951). 22. Apertural, 23. lateral, and 24. dorsal views of a large specimen,
USNM 901636, Off Zavodovski Island, South Sandwich Islands, 56?28.8'S, 27?24.6'W; 161-210 m. 25. Lateral, 26. abapertural,
and 27. apical views of the protoconch, USNM 870593, NE of loinville Island, Antarctic Peninsula, 62?41'S, 54?43'W, 210-220 m.
28. Shell ultrastructure. Fracture surface parallel to growing edge. USNM 901638, off Gibbs Island, Bransfield Strait, South
Shetland Islands, Antarctica, 61?25'S, 56?30'W, in 300 m.
M. G. Harasewych & G. Pastorino, 2008
win, 2L, W of Elephant Island, South Shetland Islands,
Antarctica, 61?12.7'S, 55?56A'W 134 m, RN Walther
Herwig, cr. 68, stn. 148; ZMH win, 2 L, W of Elephant
Island, South Shetland Islands, Antarctica, 61?8'S,
55?56.2'W 125 m, RN Walther Herwig, cr. 68, stn.
150; ZMH win, I LID, NW of Elephant Island, South
Shetland Islands, Antarctica, 600 53.9'S, 55?30A'W
135 m, RN Walther Herwig, cr. 68, stn. 159.
Literature records: Only the type locality has been
reported in the literature previously.
Distribution: Known from Antarctic Peninsula, South
Shetland Islands, South Georgia Island, South Sand?
wich Islands and South Orkneys Islands, at depths
ranging from 93 m to 460 m (Figure 21).
Remarks: The characteristic sculpture of the shell,
consisting of sharply raised spiral cords overlain by closely
spaced lamellae to produce a scabrous surface, distinguish?
es T. echinolamellata from all other known species of
Trophonella. Trophonella echinolamellata and T. scotiana
have similar bathymetric ranges, and they overlap in
portions of their geographical ranges. However, T.
echinolamellata appears to be restricted to the Scotia
tectonic plate, based on the material thus far available.
Trophonella enderbyensis (Powell, 1958),
new combination
(Figures 21, 29-31)
Synonymy: Trophon enderbyensis Powell, 1958: 197, pI.
3, fig. 1.
Description: Shell (Figures 29-31) large (to 70 mm),
fusiform, moderately thin-shelled, with regular spiral
sculpture and widely spaced lamellose varices. Proto?
conch and transition to teleoconch unknown. Tele?
oconch with 7 or more evenly convex whorls; sutural
shelf obsolete; spire tall, narrow, conical, approximate?
ly 50% of shell length; suture abutting; spire angle
about 55?. Aperture broadly ovate, deflected from the
coiling axis by 25?, interior glossy white. Siphonal canal
short; siphonal fasciole conspicuous, with recurved,
scabrous processes; pseudoumbilical chink reduced to a
narrow slit. Outer lip rounded, reflected; columellar lip
thin. Axial sculpture of low, generally distantly spaced
thin lamellae covering entire whorl, obsolete on early
whorls, approximately 3 per whorl except for the final
whorl, where 3 lamellae are closely adjacent in the
holotype. Spiral sculpture of low, rounded cords (3 or 4
on early whorls, 20 on last whorl), about as wide as
interspaces, which may have 1-4 fine spiral threads
between adjacent cords. Shell whitish. Shell composed
of three layers, as in T. scotiana. Operculum (Fig?
ure 44) oval, with terminal nucleus; outer surface with
Page 95
regular growth lines, inner surface with broad, thickly
glazed rim along posterior edge.
A single, male animal was examined and found to be
anatomically similar to T. scotiana. The radula
(Figures 16-17) is similar to that of T. scotiana but
has more prominent denticles between the central and
lateral cusps of the rachidian.
Type locality: (Figure 21: 0), off Enderby Land,
Antarctica, 65?48/S, 71?24'E in 193 m. [British, Aus?
tralian, and New Zealand Antarctic Research Expedi?
tion, St. 41, January 24/25 1930].
Type material: Holotype, SAM D15497.
Material examined: Holotype; SMF win, I D, Off the
Princess Martha Coast, Queen Maud Land, Antarc?
tica, 700 30/S, 8?4'W, 261-263 m; SMF win, 1 L, off the
Princess Martha Coast, Queen Maud Land, Antarc?
tica,71?23/S, 13?58/W, 293-357 m.
Literature records: None, apart from the original
description.
Distribution: Presently known from three specimens,
one from type locality (off Enderby Land) and two
from off Queen Maud Land, Antarctica.
Remarks: In his description of Trophon enderbyensis,
Powell (1958:198) noted that this species resembled T.
scotianus, which, at the time, was known only from
South Georgia Island. The specimens presently avail?
able indicate that the geographic and bathymetric
ranges of these two species overlap. Both Dell
(1990:208) and Numanami (1996:140) commented that
larger specimens of T. scotiana tend to have an increased
number of axial lamellae and a more inflated final whorl
of the shell. The adult holotype of Trophonella
enderbyensis is comparable in size to the largest
specimens of T. scotiana, yet it has far fewer, shorter,
axial lamellae and a narrower, more fusiform shell. The
intermediate denticle on the rachidian teeth of T.
endebryensis is pronounced, but it is reduced or absent
in T. scotiana. As more material becomes available,
especially from off Enderby Land and Wilkes Land, the
status of T. enderbyensis may require reevaluation.
Trophonella eversoni (Houart, 1997)
new combination
(Figures 32-34)
Synonymy: Trophon eversoni Houart, 1997:9, figs. 1-2,
4, 6.
Description: Shell (Figures 32-34) large (to 75.8 mm),
fusiform, of medium thickness, strongly sculptured
with spiral cords and axial lamellae. Protoconch and
transition to teleoconch unknown. Teleoconch with 6
Page 96 The Veliger, Vol. 51, No.1
Figures 29-37. Trophonella enderbyensis (Powell, 1958). 29. Apertural, 30. lateral, and 31. dorsal views of the holotype, SAM
D15497, off Enderby Land, Antarctica, 65?48'S, 71?24'E in 193 m. 32-34. Trophonella eversoni (Houart, 1997). 32. Apertural, 33.
lateral, and 34. dorsal views of the holotype USNM 880177, South Atlantic Ocean, ? Antarctica, ? Kerguelen Is. 35-37. Trophonella
rugosolamellata new species. 35. Apertural, 36. lateral, and 37. dorsal views of the holotype, ZMH 2777, off South Orkney Islands,
600 52.2'S, 44?28.4'W, in 205 m.
M. G. Harasewych & G. Pastorino, 2008 Page 97
Figures 38--46. Trophonella enderbyensis (Powell, 1958).38. Apertural, 39. lateral, and 40. dorsal views of a young specimen, SMF
win, off the Princess Martha Coast, Queen Maud Land, Antarctica, 71?23'S, 13?58'W, in 293-357 m. 41. T. rugosolamellata new
species, operculum of holotype, Figures 35-37. 42. T. echinolamellata (Powell, 1951), operculum ofspecimen in Figures 22-24.43. T.
scotiana (Powell, 1951), operculum ofspecimen in Figure 4.44. T. enderbyensis (Powell, 1958) operculum ofspecimen in Figures 38--40.
45--46. Critical-point dried penis of T scotiana (Powell, 1951).45. Dorsal and left lateral views of penis. 46. Detail oftenninal papilla.
Page 98 The Ve1iger, Vol. 51, No.1
53
pp
fo
asg
51
52
rno
vi
\~sto-
asg
E
E
LO
50
48
47
Figures 47-53. Trophonella scotiana (Powell, 1951).47. Anterior portion of male specimen, mantle reflected. 48. Ctenidium and
osphradium. 49. Anterior alimentary system. 50. Dorsal and 51. right lateral views of stomach. 52. Distal tip of penis. 53. Pallial
oviduct. Abbreviations for these figures: a, anus; asg, accessory salivary gland; bc, bursa copulatrix; cg, capsule gland; ct, ctenidium;
dd, ducts to digestive diverticula; e, esophagus; fo, female opening; i, intestine; mo, mouth; os, osphradium; p, penis; pp, penial
papilla; r, rectum; rg, rectal gland; s, siphon; sg, salivary gland; sto, stomach; vi = valve of Leiblein.
M. G. Harasewych & G. Pastorino, 2008
or more globose whorls. Sutural shelf present in last
whorls. Spire conical, about 33% of total shell length.
Suture abutting. Spire angle about 56?-59?. Aperture
large, suboval, deflected from the coiling axis by 27?;
interior glossy white; siphonal canal moderately short,
slightly inclined; siphonal fasciole scaly. Outer lip
rounded with reflected edges; columellar lip thin,
tightly adpressed, pseudoumbilical chink absent. Axial
sculpture of pronounced, regularly spaced, thin lamel?
lae, 12-14 on last whorl, running along whorl surface
from adapical suture to the siphonal fasciole. Spiral
sculpture of low, rounded, spiral cords: 3, 8, and 24 on
the first, second, and last whorls, respectively. Cords
most pronounced along shell periphery, reduced near
suture and siphonal canal. Cords cover the lamellae but
became obsolete along their edges. Shell opaque white.
Regular growth lines covering whorls and lamellae.
Operculum, radula, and anatomy unknown.
Type locality: (Figure 20:.), Antarctica, 185 m.
Material examined: Holotype, USNM 880177.
Literature records: Kerguelen Islands (Paratype 2);
submarine bank "South" off the Kerguelen Islands, in
"approximately" 250 m (Houart, 1997).
Distribution: This species is based on a holotype
collected by a Russian trawler "in the Antarctic," from
a depth of 185 m. Paratype 2 is reported, with some
certainty, to have been taken off the Kerguelen Islands.
Houart (1997:9) mentions two additional specimens
from a submarine bank south of the Kerguelen Islands,
at depths of 250 m.
Remarks: Trophonella everson! IS known from five
specimens, four of which are in private collections. The
geographic and bathymetric distributions of this species
are apocryphal, and they have yet to be confirmed by
subsequent collections. Powell (1957:113) reported the
occurrence of five species of Trophon in the Kerguelen
Islands, although all but two of the stations sampled by
the British, Australian, and New Zealand Antarctic
Research Expedition were from substantially shallower
depths. Houart (1997: 11) noted that T albolabratus
Smith, 1875, one of the Kerguelen Island species,
differs from T eversoni in having "low, thin, and more
numerous axial lamellae, stronger spiral cords, and a
shorter siphonal canal. He distinguished T eversoni
from T scotiana on the basis of the former having more
numerous spiral cords, more numerous and abapertu?
rally sloping axial lamellae that are not as strongly
produced adapically. As with T enderbyensis, addi?
tional collections from off Enderby Land, Wilkes
Land, and the islands off their coasts will be required
to assess more accurately the distribution and relation?
ships of T eversoni.
Page 99
Trophonella rugosolamellata new species
(Figures 20, 35-37)
Description: Shell (Figures 35-37) large (to 76 mm),
fusiform, solid, chalky, with strong spiral cords and
thick axial lamellae that are furrowed and reflected at
the shoulder. Protoconch unknown. Teleoconch of at
least six narrowly convex whorls; spire 33% of total
shell length. Suture abutting. Spire angle about 59?.
Aperture large, oval, deflected from the coiling axis by
about 22?, interior glossy white. Siphonal canal
moderately long and narrow, slightly inclined, siphonal
fasciole strongly scabrous, pseudoumbilical chink
closed. Outer lip rounded with reflected edge. Colu?
mellar lip broad, with thick, expanded callus. Axial
lamellae thick, more or less evenly spaced (lIon last
whorl, 8 on the penultimate whorl), running along
whorl surface from suture to the siphonal fasciole,
narrowest at midwhorl, most pronounced near and on
the siphon and especially at a position corresponding to
the shoulder, where they are enlarged, furrowed and
strongly reflected. Regular growth lines evident on the
whorls and lamellae. Spiral sculpture of rounded,
closely spaced cords (3 or 4 on early whorls, 20 on
last whorl) that extend onto the outer surfaces of the
lamellae. Cords most pronounced from siphonal canal
to region corresponding to the shoulder. Shell color
whitish. Shell composed of three layers, as in T
scotiana. Operculum (Figure 41) subpolygonal, thick,
brownish, with terminal nucleus. Growth lines on
external surface; broad, glazed rim covering nucleus;
attachment area smaller.
Animal large. Foot short, broadly rectangular.
Tentacles very wide, blunt, with small black eyes;
mantle edge smooth, siphon very short. Osphradium
small, asymmetrical, about 50% ctenidium length.
Pleurembolic proboscis short and broad. Radular
ribbon (Figures 18, 19) short (radular length ~ 0.44
aperture length), thin, extending beyond rear of buccal
mass. Rachidian tooth subrectangular, and basal plate
weakly concave anteriorly. Central cusp broadly
triangular, flanked by short stout denticles. Outer
cusps shorter and narrower, slightly outwardly direct?
ed. Marginal cusps absent, marginal area smooth.
Lateral teeth L-shaped, narrower than rachidian teeth.
Esophagus short. Salivary glands large, ascinous,
whitish, with ducts entering esophagus just anterior
to the wide valve of Leiblein. Accessory salivary glands
small, spherical, located in the anterior part of the
buccal mass, totally embedded in salivary glands.
Gland of Leiblein compact, without lobules, brownish
in color, overlaying esophagus. Penis similar to that of
T scotiana; large, wide, flat, > 4 times tentacle length;
with terminal, conical papilla surrounded by narrow
collar.
Page 100 The Veliger, Vol. 51, No.1
I
?
MP
Muricanthus
Forreria
Nucella CDeu
Acanthina C
?C
Trochia ~
Ocenebra erinacea C
Trophon geversianus ~
Haustrum 0
Trophonella scotiana
Boreotrophon aculeatus
Paziella pazi
Morula
Cronia
Vexilla
Orupa
Nassa
Pinaxia
Oicathais
Vasula
Tribulus
Thais
Neorapana
Mancinella
Purpura
Plicopurpura
Rapana
Stramonita
Concholepas
Cymia NJ
Figure 54. Strict consensus maximum parsimony [MP] and neighbor-joining [NJ] trees depicting the relationships of genera based
on data derived from their type species. Data from Kool (1993b:Table 3) have been supplemented by data for Ocenebra erinaceus,
Trophon geversianus, Trophonella scotiana, Boreotrophon aculeatus, and Paziella pazi.
Type locality: S of Laurie Island, South Orkney Islands,
6Oo 52.2'S, 44?28.4'W, in 205 m.
Type material: Holotype, ZMH 2777.
Material examined: This species is presently known
only from the holotype.
Distribution: Known only from the type locality,
collected by the RN Walther Herwig cr. 68, stn. 91
(Figure 20).
Etymology: rugosolamellata from Latin rugosus and
Latin lamellata (with lamellae, or thin plates) referes to
the extension of the spiral cords onto the axial lamellae.
Remarks: Trophonella rugosolamellata most closely
resembles T eversoni, but it can be distinguished by
its thicker shell, with slightly fewer lamellae and fewer,
coarser spiral cords. The axial lamellae of T rugoso?
lamellata are distinctive in that they are reflected along
the outer edges, especially at a position corresponding
to the shoulder, where they are enlarged, furrowed, and
M. G. Harasewych & G. Pastorino, 2008 Page 101
Table 1
Taxa, characters, and character states used in an analysis of the relationships of Trophoninae. The data matrix is that
of Kool (1993b:table 3), to which have been added the taxa Trophon geversianus, Ocenebra erinaceus, Trophonella
scotiana, Boreotrophon aculeatus, and Paziella pazi. See Kool (1993b) for detailed descriptions of characters and
character states.
Character 2 3 4 5 6 7 8 9 La 11 12 13 14 15 16 17 J8
Taxa [rom Kool (1993b):
Muricanthus a a a a a a a a a a a a a a a a a a
Forreria b b b a b a b a a a b a a a b ? b c
Nucella b b b a b a b a a a b b c a b b b c
Haustrum b ? c a b a a a a a b b b a e b a b
Morula a a a a c b a b b b c c d b c a a e
Crania c a a a c b a b b b d c d b b a a [
Rapana a a c a d a a a b c d e d b b a a [
Cymia ? ? c a e a b a b a ? d d b b a a d
Stramonita a a b a e a a a b c d e d b b a a g
Concholepas a a b a e a a a b c d e d b b a a g
Dicathais c a b a e a a b b c ? e d b c a a g
Vasula ? ? ? b e a a b b c d e d b c ? a j
Vexilla d a a a [ a c b b c d f d b d a a ?
Nassa a a a a e a a b b c d f d b c a a ?
Pinaxia a a a a e a a b b c d f d b c a a h
Drupa ? ? ? a e a a b b c d e d b d a a h
Plicopurpura a a ? a e a a b b c d e d b c a a ?
Thais ? a c b e a a b b c d e d b c a a j
Purpura ? a c b e a a b b c d e d b c a a h
Mancinella ? ? c b e a a b b c ? e d b c a a
Neorapana ? a c b e a a b b c d e d b c a a J
Tribulus ? a c b e ? a b ? ? ? e d ? ? ? a J
Acanthina b b ? a b ? b ? a ? ? b ? ? ? ? ? ?
Trochia b b b a ? ? ? ? ? ? ? ? ? ? ? ? ? ?
Newly added taxa:
Ocenebra erinaceus b b b a a ? b a a a b b d a b b b c
Trophon geversianus b a b a a ? a a a a b b c a c b b c
Trophonella scotiana d a b a a ? a a a a b b a a c b a a
Boreotrophon aculeatus d a c a a ? a a a a b b a a c b a a
Paziella pazi b a c a a ? a a a a a b a a c b a a
strongly reflected to produce a tapering structure
similar to the open shoulder "spine" of Pterochelus
Jousseaume, 1880. The spiral cords of T eversoni are
more numerous and thinner, and they are not as
pronounced along the siphonal canal or on the axial
lamellae as in T rugosolamellata. Trophonella ender?
byensis has a higher spire, fewer, smaller, and more
widely spaced lamellae, and fainter spiral cords than T
rugosolamellata.
PHYLOGENETIC RELATIONSHIPS
A maximum parsimony analysis of the data matrix
shown in Table I [PAUP* 4.0 beta version 10 Heuristic
search, Accelerated transformation, TBR Swapping
algorithm] produced 475,972 equally parsimonious
trees (length = 60; consistency index = 0.783; retention
index = 0.903]. A strict consensus of these trees is
shown in Figure 54 (MP). Figure 54 (NJ) shows the
neighbor joining tree based on the same data.
Ocenebra erinaceus, the type species of the type gen us
of Ocenebrinae, emerges in an unresolved pentatomy
with the genera Nucella, Trochia, Forreria, and
Acanthina in the strict-consensus tree and as sister
taxon to these genera in the neighbor-joining tree.
Trophon geversianus, the type species of the type genus
Trophoninae, emerges as the sister group to this clade
in both the MP analysis and the NJ analysis. While the
NJ tree shows Haustrum to be the sister taxon to this
clade, which, in turn, was joined by the clade
[(Trophonella + Boreotrophon) Paziella], the MP
analysis groups the genera Haustrum, Trophonella,
Boreotrophon, and Paziella into an unresolved polyt?
omy that is sister to Trophon and the polytomy
containing Ocenebra.
These results indicate that the type species of
Page 102
Trophonella is more closely related to Boreotrophon
than to the type species of Trophon and thus sup?
port the segregation of Trophonella from Trophon
at the generic level. They also confirm the conten?
tions by various authors that Trophoninae, as
defined by Cossmann (1903:9), is not monophyletic.
Based on the relationships of the type species of their
respective type genera, either Trophoninae is the
sister taxon of Ocenebrinae or both are part of a larger
clade.
In his phylogenetic study, Kool (1993b:fig. 30)
considered Ocenebrinae to include Haustrum. Tan
(2003) subsequently erected the subfamily Haustrinae
(with Haustrum as the type genus) to encompass a clade
of nonrapanine muricids endemic to Australia and
New Zealand. However, the relationship of this clade
to Ocenebrinae and Rapaninae varied significantly,
depending on the choice of outgroup. Beu (2004:216)
concluded that it seemed preferable to retain the taxa
included by Tan in Haustrinae within the subfamily
Ocenebrinae. This broader delineation of Ocenebrinae
includes the type species of Trophoninae in both
MP and NJ trees. The genera Trophonella and
Boreotrophon emerged as part of an unresolved
polytomy with Haustrum in the strict-consensus MP
tree, but were depicted as being more distantly related
in the NJ tree.
Both Ocenebrinae and Trophoninae were proposed
in the same publication (Cossmann, 1903: 10), so
neither has priority. Although it is likely that these
two subfamilies will be synonymized, based on the
close relationship between the type species of their type
genera, a better resolved phylogeny, including a more
comprehensive sampling of the more than 20 genus?
level taxa attributed to each of these two subfamilies,
will be required to delineate more precisely the
membership of the clade and to identify its diagnostic
synapomorphies.
Acknowledgments. We thank the following curators and
collection managers for access to specimens in their collec?
tions: K. Way (NHM); A. Waren (NHRM); P. Bouchet
and V. Heros (MNHN); P. Mikelsen (AMNH); B. Hausdorf
(ZMH), A. Tablado (MACN); C. Ituarte; (MLP) and R.
Janssen (SM); P. Arnaud, Endoume, France. This work was
supported by a grant from the Consejo Nacional de
Investigaciones Cientificas y Tecnicas (CONICET), Argen?
tina, which enabled the junior author to work in the Divi?
sion of Mollusks, United States National Museum of
Natural History, Smithsonian Institution. Additional sup?
port was provided in part by a Research Award from the
NSF-USAP United States Antarctic Program Grant
[ANT0636408] and a grant-in-aid from the Conchologists
of America and the Walter E. Sage Memorial Award. We
are grateful to Dr. Yu. Kantor for his many helpful and
insightful comments during the course of this research, and to
Roland Houart and Dr. Geerat Vermeij for their helpful
reviews.
The Ve1iger, Vol. 51, No.1
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