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Volume 126, Number 1
March 30, 2012
ISSN 0028-1344
A quarterly devoted
to malacology.
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National Museum of
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Department of Invertebrates
Field Museum of
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North Carolina State Museum of
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Florida Museum of Natural History
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Institution
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Museum of Zoology and Department
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Florida Museum of Natural History
University of Florida
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The Academy of Natural Sciences
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Dr. Geerat J. Vermeij
Department of Geology
University of California at Davis
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Dr. G. Thomas Watters
Aquatic Ecology Laboratory
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Columbus, OH 43212-1194
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THEC'NAUTILU S
Volume 126 , Number 1
March 30, 2012
ISSN 0028-1344
CONTENTS
R.E. Petit
Silvio Felipe Barbosa de Lima
Book Review
43
Notice
46
THE NAUTILUS 126(1): 1-14, 2012
Page 1
Hispaniolan Annulariidae (Gastropoda), primarily from the
Barahona Peninsula: New taxa and notes
G. Thomas Watters
Department of Evolution, Ecology, and Organismal Biology
Ohio State University
Columbus, OH 43212 USA
[email protected]
ABSTRACT
Eleven new taxa of Hispaniolan Annulariidae are described:
Abbottella ( Abbottella ) urbana new species, Chondropoma
( Chondropoma ) crijstallinum new species, Chondropoma
( Chondropoma ) duffyhooksorum new species, Chondropoma
( Chondropoma ) vanattae pohjchroma new subspecies,
Chondropoma ( Wetmorepoma ) morsecodex new species,
Parachondria ( Parachondria ) gettlemani new species, Articulipoma
rhodei new species, Chondropominm blaineorum new species,
Chondropomium lynx new species, Chondropominm eusarciim
saonaense new subspecies, and Licina bartschi new species.
Based on new material, Chondropoma ( Chondropoma ) manielen.se
montivagum Bartseh, 1946, is synonymized with the nominal
subspecies Chondropoma ( Chondropoma ) manielen.se manielen.se
Bartseh, 1946 and Chondropoma (Chondropoma) quisquen.se
sculptior Bartseh, 1946 is synonymized with the nominal sub-
species Chondropoma ( Chondropoma ) qui.squen.se qui.squen.se
Bartseh, 1946. Chondropoma (Chondropomium) venniculatum
sensu lato Bartseh, 1946 is moved to Crossepoma based on opercu-
lar features. The type locality of Chondropoma (Chondropomium)
venniculatum sallei Bartseh, 1946 is restricted to Cabral,
Barahona Province, Dominican Republic. The type locality of
Choanopoma solution Pfeiffer, 1S52 is restricted to the southern
edge of Los Haitises Mountains, N of Majagual, ca. 12 km NW
of Sabana Grande de Boya, Monte Plata Province, Dominican
Republic. The type locality of Chondropoma loweanum Pfeiffer,
1852 is restricted to Loma del Aguaeate, Sierra Martin Gracia,
Barahona Province, Dominican Republic. Distributional and
habitat notes are given for additional species.
Additional keywords: Hispaniola, Dominican Republic, habitat
INTRODUCTION
Bartseh (1946) and Watters (2006) recognized the
Barahona Peninsula of the Dominican Republic as an
area of high endemism for the Annulariidae. Bartseh
(1946) described 21 species of Annulariidae from the
peninsula out ol 67 for all of the Dominican Republic.
As impressive as this seems, on closer inspection it is
apparent that Bartseh’s species all came from relatively
few sites in the eastern half of the peninsula; the western
half has never been adequately sampled or reviewed.
Recent collections there and re-examination of older col-
lections have revealed a wealth of previously unknown
species (Watters and Duffy (2010a, 2010b), Watters
(2010)), the description of which is continued here.
MATERIALS AND METHODS
Length is measured from the tip of the apex or decol-
lated spire to the end of the outer aperture. Width is
measured as the maximum dimension in a plane with
the aperture perpendicular to the axis of coiling. Spiral
sculpture is counted from the suture to the inside of the
umbilicus. Dimensions in captions refer to shell length.
Shells are solute if a portion of the final whorl is
detached from the previous whorl; shells are adnate if
they are not detached. Inner apertural lips are exserted
if they protrude beyond the plane of the outer lips.
Abbreviations used in the text are: BMSM, The Bailey-
Matthews Shell Museum, Sanibel, Florida, USA; GTW,
collection of the author, Columbus, Ohio, USA; NHMUK,
Natural History Museum, London, UK; OSUM, Ohio
State University Museum of Biological Diversity,
Columbus, Ohio, LISA; UF, Florida Museum of Natural
History, Gainesville, Florida, USA.
SYSTEMATICS
Family Annulariidae Henderson and Bartseh, 1921
Subfamily Annulariinae Henderson and Bartlsch, 1921
Genus Abbottella Henderson and Bartseh, 1921
Subgenus Abbottella Henderson and Bartseh, 1921
Type Species: Chondropoma moreletianum Crosse,
1873, by original designation.
Abbottella (Abbottella) urbana new species
(Figures 1-3)
Description (2 specimens): Shell ca. 5 mm in length,
turbinoid, umbilicus wide (holotype 35% of maximum
width), circular, open to apex. Nuclear whorls IVz, scarcely
demarcated from teleoconch, smooth, minute and erect.
Page 2
THE NAUTILUS, Vol. 126, No. 1
Figures 1—21. Abhottella , Leiahhottella, and Chondropoma species. 1-3. Abbottella ( Abbottella ) urbana new species. Holotype,
UF 446061, 4.9 mm. 4-7. Leiahhottella soluta (Pfeiffer, 1852). 4. Illustration from Pfeiffer (1854b: pi. 39, fig. 8). 5-7. UF 249105,
2 km N of Majagual, Monte Plata Province, Dominican Republic, 11.3 mm width. 8-9. Chondropoma ( Chondropoma ) cnjstallinum
new species. Holotype, UF 446062, 15.0 mm. 10-12. Chondropoma {Chondropoma) duffijhooksorum new species. 10-11. Holotype,
UF 446063, 9.3 mm. 12. Paratype, OSUM 36510, 8.5 mm. 13-14. Chondropoma ( Chondropoma ) manielense Bartsch, 1946. GTYV
71671), 13 km NNE of Pedernales, Pedemales Province, Dominican Republic, 13 mm. 15-17. Chondropoma ( Chondropoma )
qum/uense Bartsch, 1946. 15-16. GTYV 7082b, 3—4 km NW of Oviedo, Pedernales Province, Dominican Republic, 14.0 mm. 17.
GTYV 7082c, 10 km NW of Oviedo on Highway 44, Pedernales Province, Dominican Republic, 13.8 mm. 18-20. Chondropoma
( Chondropoma ) vanattae poli/chroma new subspecies. 18-19. Holotype, UF 446064. 11.4 mm. 20. Paratype, OSUM 36511, 10 km
ENE of Las Mercedes, Pedernales Province, Dominican Republic, 12.2 mm. 21. Chondropoma {Chondropoma) vanattae vanattae
Pilsbry, 1933. GTW 7090a, Polo, Barahona Province, Dominican Republic, 13.5 mm.
G. T. Watters, 2012
Page 3
Teleoeoncii of 3 1/4 whorls, adnate, last 1/8 whorl
scarcely deflected anteriorly. Suture impressed. Peri-
stome double, circular (holotype 1.9 mm inner peristome
diameter; 2.5 mm outer peristome diameter). Outer lip
thin, lamellate, evenly but narrowly expanded with a
barely perceptible auricle, lip deflected abaperturally.
Inner lip exserted, thick, short. Spiral sculpture of
numerous, weak spiral threads (ca. 24 on final whorl),
stronger and more widely spaced in umbilicus, weakest
on final whorl. Axial sculpture of widely spaced threads
and thicker ribs, ca. 70 on final whorl. Intersections of
spiral and axial sculpture form minute pustules, some-
what stronger on spire. Axial sculpture forms weak cusps
at suture. Shell dirty white with ca. 7 faint, tan narrow
bands. Outer lip white with faint bands, which are
apparent on both sides of lip. Operculum, anatomy, and
radula unknown.
Holotype: UF 446061 .
Type Locality: Parque Central, off Jose Contreras Blvd.,
N of the Loteria suburb of western Santo Domingo, Distrito
Nacional, Dominican Republic, 18.44° N, -69.97° W.
Paratype: OSUM 36509, from the type locality
(1 specimen).
Distribution: Lowlands just west of Santo Domingo,
Dominican Republic.
Habitat: Limestone slabs.
Comparison with Other Species: Only one other
species has presumably been found in the vicinity of
Santo Domingo, Abbottella moreletiana domingoensis
Bartsch, 1946. That taxon differs from A. urbana in its
much larger adult size (ca. 12 mm diameter vs. ca. 6 mm
for A. urbana), in having a much wider outer lip, and in
having coarser sculpture (ca. 60 axial ribs vs. ca. 70 in
A. urbana). Abbottella urbana is similar to Abbottella
moreletiana gabriella Bartsch, 1946, but is even smaller
(ca. 8.6 mm diameter for A. m. gabriella), has more
numerous axial ribs (53 in A. m. gabriella), and has
more spiral threads (ca. 24 in A. urbana vs. 17 in A. m.
gabriella). Additionally, Abbottella moreletiana gabriella
is only known from Isla San Gabriel in Samana Bay in
the easternmost part of the island.
Etymology: L. urbanus, of the city, in reference to the
type locality of the species.
Genus Leiabbottella Watters, 20 1 0
Type Species: Leiabbottella galaxius Watters, 2010,
by original designation.
Leiabbottella solutus (Pfeiffer, IS52)
(Figures 4-7)
Choanopoma solution “Richard” Pfeiffer, 1852: 167.
Cyclostoma solution (Pfeiffer, 1852). Pfeiffer, 1854b: 295, pi. 39,
figs. 8-10.
Abbottella solution (Pfeiffer, 1852). Henderson and Bartsch,
1921: 75.
Incertipoma solution (Pfeiffer, 1852). Bartsch, 1946: 171, 173-174,
pi. 30, figs. 2-4.
Rolleia? soluta (Pfeiffer, 1852). Watters, 2006: 93, 487-488.
Leiabbottella solutus (Pfeiffer, 1852). Watters, 2010: 17-18.
Distribution: Described from the “Island of Santo
Domingo” Bartsch (1946) had no examples beyond
Pfeiffer’s description and illustration and he seemed to
hint that he believed the locality was spurious. However,
specimens (UF 249105) have since been collected at 150 m,
southern edge of Los 1 1 ai rises Mountains, N of Majagual,
ca. 12 km NW of Sabana Grande de Boya, Monte Plata
Province, Dominican Republic. Specimens were collected
in a coffee grove at the base of a limestone ledge in leaf
mulch and under rocks (F.G. Thompson, UF, pers. comm.,
January, 2011). The type locality of Choanopoma solution
Pfeiffer, 1852 is herein restricted to 150 nr, southern
edge ol Los Haitises Mountains, N of Majagual, ca. 12 km
NW of Sabana Grande de Boya, Monte Plata Province,
Dominican Republic, 19.05° N, -69.83° W.
Subfamily Chondropomatinae Henderson and Bartsch, 1921
Genus Chondropoma ( Chondropoma ) Pfeiffer, 1847
Subgenus Chondropoma Pfeiffer, 1847
Type Species: Cijclostoma sagra d’Orbigny, 1842, by
subsequent designation of Petit de la Saussaye, 1850.
Chondropoma (Chondropoma) crystallinwn new species
(Figures 8, 9)
Description (Holotype): Shell 15.0 mm maximum
length, including peristome x 13.2 mm maximum width,
including peristome, fragile, translucent, turbinate, umbi-
licus moderately wide (20% of maximum width).
Protoconch whorls present in adult, IV2 whorls, not
demarcated from teleoeonch, smooth, minute but erect,
with a diffuse brown peripheral band. Teleoeonch ol
4 whorls, adnate. Suture deeply impressed. Peristome
double, oval (6.2 mm diameter maximum inner aper-
ture height x 5.3 mm diameter maximum inner aper-
ture width; 8.8 mm diameter maximum outer peristome
height x 7.3 mm diameter maximum outer peristome
width). Outer lip thin, expanded perpendicular to whorl,
nearly evenly produced around peristome, adnate, lack-
ing an auricle. Inner lip barely exserted, simple. Spiral
sculpture present as sharp, narrow threads, weakest at
periphery, ca. 39 on final whorl including umbilicus. Axial
sculpture present only as numerous, minute beads on
spiral threads. Suture with minute denticles. Teleoeonch
ol faint bronze color, with 3 faint rows ol narrow, widely
spaced bands of darker spots at, just above, and below
midline, most prominent on earlier whorls; umbilicus
brown. Outer lip of peristome and portion of final whorl
immediately before lip are white, unpattemed. Inner peri-
stome bronze. Operculum, anatomy, and radula unknown.
Holotype: UF 446062.
Page 4
THE NAUTILUS, Vol. 126, No. 1
Type Locality: 22 km N of Pedemales, off road HH to
Agua Negra, at 460 m, Pedernales Province, Barahona
Peninsula, Dominican Republic, in cleared field on south-
ern slope of the Sierra de Baoruco. 18.10° N, -71.64° W.
Distribution: Known only from the type locality.
Habitat: This species was found with live and dead
individuals of Chondropoma manielen.se Bartsch, 1946
and Articulipoma rhodei new species in a cleared field
of rocks with little top soil.
Comparison with Other Species: This remarkable
species is unlike any other Hispaniolan Chondropoma.
The combination ol large size, translucent shell, and
nearly unicolored whorls with contrasting white peri-
stome is unique. It is similar to C. manielense , with which
it is found, in possessing the similar sculpture of spiral
threads regularly produced into minute points; this sculp-
ture is finer and less coarse in C. crystallinum than in
C. manielense. In addition, Chondropoma manielense is
smaller, more solid, and usually boldly patterned.
Etymology: L. crystallinum , of crystal, in reference to
the glassy, translucent shell.
Chondropoma ( Chondropoma ) duffyhooksorum
new species
(Figures 1 0-12)
Description (2 specimens): Shells S.5-9.3 mm in
length (holotype 9.3 mm maximum length decollate,
including peristome x 4.8 mm maximum width, including
peristome), high-spired, umbilicus narrow. Protoconch
whorls decollate in all examples studied. Teleoconch of
4 whorls, solute just before peristome. Suture minutely
channeled. Peristome double but fused to form a single
lip, tear drop-shaped (holotype 3.0 mm diameter maxi-
mum inner aperture height x 2.7 mm diameter maxi-
mum inner aperture width). Outer lip thin, not expanded
except for a minute auricle. Inner lip adherent to outer
lip, simple. Spiral sculpture of ca. 25 narrow, low threads,
widely spaced. Axial sculpture of irregularly spaced, nar-
row threads (ca. 58 on final whorl), produced into min-
ute, vertical pustules at intersections with spiral threads.
Suture serrate but without cusps or tufts. Teleoconch tan
with 6 darker, interrupted spiral bands equally spaced from
suture through umbilicus. Operculum thin, pauci spiral with
a fine granular deposit. Anatomy and radula unknown.
Holotype: UF 446063.
Type Locality: On an old house, Boca Chica, at nearly
sea level, Santo Domingo Province, Dominican Republic,
18.46° N, -69.61° W.
Paratype: OSUM 35610, from the type locality
(I specimen).
Distribution: Known only from the typo locality.
Comparison with Other Species: Chondropoma
duffyhooksorum is related to Chondropoma catalinen.se
Bartsch, 1946 from Isla Catalina off La Rornana, ca.
64 km E of Boca Chica. Chondropoma duffyhooksorum
differs in being smaller (ca. 9 vs. ca. 12 mm in length),
in the more pronounced spiral sculpture, in the nearly
completely adnate final whorl (solute for l/6fl1 of the
final whorl in C. catalinense) , and in the better defined
and continuous brown spiral bands. No similar forms
have been found between Isla Catalina and Boca Chica.
The specimens also greatly resemble the crude illus-
trations of Cyclostoma clominicense Pfeiffer, 1850 given
by Pfeiffer in 1854b (pi. 38, figs. 9, 10). However,
Watters (2006) placed that species in Colonina based on
examination of the syntype lot (NHMUK 1996132) and
synonymized Haitipoma catalinense Bartsch, 1946 with
it. Thus Chondropoma catalinense Bartsch, 1946 and
Colonina clominicense (Pfeiffer, 1850) are both endemic
to Isla Catalina and are veiy similar in appearance but
actually belong to different genera.
Etymology: Named for Glenn Duffy and Randy Hooks,
who have generously contributed many annulariid speci-
mens for study.
Chondropoma (Chondropoma) manielense
Bartsch, 1946
(Figures 13, 14)
Chondropoma (Chondropoma) manielense manielense Bartsch,
1946: 53, pi. 10, fig. 3.
Chondropoma (Chondropoma) manielense rnontivagurn Bartsch,
1946: 53, pi. 10, fig. 2.
Discussion: Bartsch (1946) described two subspecies
based on differences in size and degree of coloration; both
subspecies originated from the region of Manuel Viejo
in the Barahona Peninsula. While variously known as
Maniel Viejo to Bartsch and appearing as such on some
current maps, the official name is Manuel Viejo. In the
several collections reported here (GTW 7167a-d) adult
specimens from the same locality range in size from 10 to
15 mm maximum length and vaiy from darkly colored to
nearly white. There is no reason to maintain these two
taxa as distinct and the subspecies C. (C.) manielense
rnontivagurn Bartsch, 1946 is herein synonymized with the
nominal subspecies. It is curious to note that some spec-
imens have a single peristome whereas odiers have a dou-
ble peristome.
The records here extend the range of this species to
the southern slope of the Sierra de Baoruco north of
Pedernales, where it occurs under rocks in fields and in
outcrops at 500-700 m. The overall range appears to be
the southern slopes of the Sierra de Baoruco across the
entire Barahona Peninsula.
Chondropoma ( Chondropoma) quisquense
Bartsch, 1946
(Figures 15-17)
G. T. Watters, 2012
Page 5
Chondropoma (Chondropoma) quisquense quisquense Bartsch,
1946: 61, pi. 11, fig. 1.
Chondropoma (Chondropoma) quisquense sculptior Bartsch,
1946: 61-62, pi. 11, fig. 2.
Discussion: Bartsch (1946) described C. quisquense
quisquense from Trujin and C. quisquense sculptior from
Manuel Viejo, both in the southeastern portion of the
Barahona Peninsula. Recent collections from the Oviedo
region (near Bartsch’s Trujin, GTW 7082b) and Manuel
Golla (15 km W of Oviedo, GTW 7082c) indicate that
this species occurs in the lowlands around the south-
eastern edge of the Sierra de Baoruco. Specimens were
found in fields with large hardwood trees. Bartsch dif-
ferentiated the two subspecies based on the degree to
which the final whorl was solute and the strength of the
sculpture. The new collections reported here include
both subspecies within a single lot as well as intergrades.
Therefore these two taxa are not considered distinct
and C. (C. ) quisquense sculptior Bartsch, 1946 is herein
synonymized with the nominal subspecies.
Chondropoma (Chondropoma) ranattae sensu Into
Pilsbry, 1933
Chondropoma vanattae Pilsbry, 1933: 124, pi. 6, figs. 8, 9.
Chondropoma ( Chondropoma ) vanattae Pilsbry, 1933. Bartsch,
1946: 52, 70-71, pi. 12, fig. 3.
This species breaks up into three subspecies across
the Chaine de la Selle mountain range, from east to
west: C. vanattae vanattae Pilsbry 1933 in the extreme
eastern part of the Sierra de Baoruco; C. vanattae
polychroma new subspecies on the southern slope of
the western Sierra de Baoruco; and C. vanattae
verettense Bartsch, 1946 from the Fonds-Verette region
of the Massif de la Selle in Haiti.
Chondropoma ( Chondropoma) vanattae pohjchroma
new subspecies
(Figures 18-20)
Description (4 specimens): Shell 11.4-12.3 mm in
length (holotype 11.4 mm maximum length, including
peristome x 6.8 mm maximum width, including peri-
stome), solid, conic, umbilicus narrow (holotype 7% of
maximum width). Protoconch whorls retained in adults,
114 whorls, scarcely demarcated from teleoeoneh, smooth,
minute but erect, a diffuse brown band at periphery
and a darker band at suture. Teleoeoneh of 4 whorls,
solute just before peristome. Suture impressed but not
channeled. Peristome single, tear drop-shaped (holo-
type 4.0 mm diameter maximum inner aperture height x
3.2 mm diameter maximum inner aperture width), nar-
row, barely expanded, lacking an auricle but posterior
edge of lip forming 90° angle, inner lip just touching
previous whorl. Spiral sculpture of irregularly spaced low
cords of various widths, ca. 25 on final whorl, cords absent
immediately below suture. Axial sculpture of irregularly
spaced, wide, rounded cords of various widths with occa-
sional fine threads in between (ca. 70 cords on final
whorl). Intersections of axial and spiral sculpture barely
produced into pustules, strongest on spire. Sculpture has
texture of coarse cloth. Suture serrate but lacks cusps
or tufts. Teleoeoneh with a light tan base with irregular
dark brown blotches arranged in loose axial anti spiral
bands. Final l/S1^11 of final whorl with a pinkish hue. Inside
of peristome orange. Operculum thin, paucispiral with a
fine granular deposit. Anatomy and radula unknown.
Holotype: UF 446064.
Type Locality: 9 km NE of Las Mercedes, Pedernales
Province, Dominican Republic, at 1000 m on trees in
pine forest, 18.12° N, -71,57° W.
Paratypes: BMSM 1 7944 ( 1 specimen); OSUM 3651 1
(1 specimen); NHMUK 20110335 (1 specimen). All
paratypes from 10 km NE of Las Mercedes, Pedernales
Province, Dominican Republic, at 1300 m in pine forest,
18.13° N, -71,56° W.
Distribution: Chondropoma vanattae pohjchroma is
known from 9-10 km NE of Las Mercedes at 1000-
1300 m. Chondropoma vanattae vanattae Pilsbry, 1933
was described from “Station 85, Sr. Del Monte’s planta-
tion,” which was in a “verdant gully. . . 5-6 miles west of
Barahona as the crow flies” at 1000 m. Bartsch (1946)
added records from 3 km N of Manuel Viejo at 1000 m.
The specimen of C. vanattae vanattae illustrated here
(Figure 21, GTW 7090a) is from Polo at 610 m.
Chondropoma vanattae verettense Bartsch, 1946, was
described from Fonds-Verette, Haiti, at ca. 800 m. As a
whole, this species occurs in the Chaine de la Selle
mountain range from ca. 600-1300 m but intergrades
are not known between the subspecies.
Habitat: On trees in pine forests at ca. 1000-1300 m.
Comparison with Other Species: Chondropoma
vanattae pohjchroma differs from other subspecies of
C. vanattae in its coarser sculpture, more vivid colora-
tion, and in particular its orange aperture.
Etymology: Gr. poly, many + Gr. chroma , color, many
colored, in reference to the colorful appearance of the shell.
Genus Chondropoma Bartsch, 1946
Subgenus Wetmorepoma Bartsch, 1946
Type Species: Chondropoma ( Wetmorepoma ) wetmorei
Bartsch, 1946, by original designation.
Discussion: This subgenus was based on a single spe-
cies, Chondropoma wetmorei Bartsch, 1946 from Isla
Beata off the southern tip ol the Barahona Peninsula.
Watters and Dully (2010a) added C. (W.) oculeum from
the Cabo Rojo/Pedernales area ol the western peninsula.
A third species is described here from E of Cabo Rojo
westward to Oviedo. The subgenus was created by
Bartsch (1946) for elongate Chondropoma lacking any
Page 6
THE NAUTILUS, Vol. 126, No. 1
spiral sculpture except in the umbilicus. This subgenus
appears to be a true endemic of this peninsula occurring
in the lowlands of the south and west, including; Isla
Beata. Further genetic research will probably show
Wetmorepoma to be a subgenus of Chondropomium
rather than Chondropoma.
Chondropoma (Wetmorepoma) morsecodex new species
(Figures 22, 23)
Description (21 specimens): Shells 10.0-13.0 min in
length, decollate (holotype 13.0 mm maximum length,
decollate, including peristome x 6.0 mm maximum
width, including peristome), translucent, high-spired,
umbilicus narrow. Protoconch whorls decollate in all
specimens examined. Teleoconch of 4 whorls, last 1/4* 1
turn free from previous whorl. Suture sharply impressed.
Peristome double, tear drop-shaped (holotype 3.6 mm
diameter maximum inner aperture height x 2.8 mm
diameter maximum inner aperture width; holotype
4.9 mm diameter maximum outer peristome height, includ-
ing auricle x 4.3 mm diameter maximum outer peristome
width). Outer lip thin, expanded perpendicular to whorl,
narrowest facing umbilicus, widest anterior and medial,
solute from previous whorls, minute auricle present.
Inner lip barely exserted, simple. Spiral sculpture pres-
ent only as two very feeble cords within umbilicus. Axial
sculpture of very weak, wide, low ribs, nearly obsolete on
most of shell, strongest on last lA of final whorl. Suture
smooth. Teleoconch glossy white with irregularly spaced
axial bands of tan spots and dashes, four to a band; fused
into a solid axial band in some specimens. Operculum
paucispiral, corneous. Anatomy and radula unknown.
Holotype: UF 446065.
Type Locality: Ca. 9.3 km NW of Manuel Golla, ea.
140 m, off Highway 44, Pedernales Province, Barahona
Peninsula, Dominican Republic, under rocks, 17.94° N,
-71.65° W.
Paratypes: OSUM 36512 from the ripe locality
(1 specimen); UF 249171, from 210 m, 17 km NW of
Oviedo, Pedernales Province, Dominican Republic.
17.92° N, -71,51° W (15 specimens).
Other Material Examined: Jesus Santana Benitez
coll., 13.5 km W of Oviedo, Pedernales Province,
Dominican Republic (4 specimens).
Distribution: Lowland valley aloi ig Higl lway 44 from
Oviedo to at least Manuel Golla, southern Barahona
Peninsula.
Habitat: Specimens were found under rocks on a hill
slope with limestone outcrops next to recently cleared
agricultural land.
Comparison with Other Species: This is the third
species described in this subgenus, all from the
Barahona Peninsula. It differs from both C. wetmorei
(Figures 24, 25) and C. oculeum (Figure 50) in being
larger, having a wider peristome, and having a distinct
pattern of dark brown rectangular spots and dashes
arranged in irregularly spaced axial bands, occasionally
as a single unbroken band (C. wetmorei lias large, bold,
round spots, C. oculeum has smaller, fainter spots).
Etymology: Morse + L. codex , writing, Morse code,
in reference to the linear series of dots and dashes on the
shell (on the holotype, from anterior to posterior, dot,
dot, dot, dash = “V”). A noun in apposition.
Genus Parachondria Dali, 1905
Subgenus Parachondria Dali, 1905
T>pe Species: Turbo fascia Wood, 1828, by original
designation .
Parachondria (Parachondria) gettlemani new species
(Figures 2G-28, 58)
Description (7 specimens): Shell 13.9-14.6 mm in
length (holotype 13.9 mm maximum length including
peristome, decollate x 7.7 mm maximum width includ-
ing peristome), solid, high-spired, umbilicus narrow (holo-
type 8% of maximum width). Protoconch whorls decollate
in some adult specimens, retained in others, IV2 whorls,
scarcely demarcated from teleoconch, smooth, minute but
erect, white. Teleoconch of 5 whorls, adnate except for just
behind peristome. Suture minutely channeled. Peristome
single, tear drop-shaped (holotype 4.8 mm diameter max-
imum inner aperture height x 3.5 mm diameter maxi-
mum inner aperture width; holotype 6.0 mm diameter
maximum outer peristome height, including auricle x
4.6 mm diameter maximum outer peristome width). Lip
thin, expanded perpendicular to whorl, narrowest facing
umbilicus, barely adnate to previous whorls. Small trian-
gular auricle. Spiral sculpture present only as 2 barely
perceptible cords within umbilicus. Axial sculpture of
regularly spaced, narrow threads (ca. 90 on final whorl),
interstices smooth. Suture broken by axial lamella as
minute cusps, groups of 3-6 cusps slightly larger.
Teleoconch tan with or without a pattern of small spots
and fine zigzag tan markings that may coalesce to form
vague, narrow bands; all specimens seen have a wide
brown basal band composed of blurred, dark brown spots;
there may be narrow bands between this basal band and
umbilicus; wide brown band deep within umbilicus;
pseudotufts white. Operculum thin, paucispiral with a
granular deposit. Anatomy and radula unknown.
Holotype: UF 446066.
Type Locality: Virgen de San Rafael, ca. 200 m, ca.
7 km NNE of Parafso, Barahona Province, Barahona
Peninsula, Dominican Republic, on trees. 18.05° N,
-71.12° W.
Paratypes: BMSM 17945 (1 specimen); UF 446067, juve-
nile (1 specimen), OSUM 36514, juveniles (2 specimens);
G. T. Watters, 2012
Page 7
Figures 22-42. Chondropoma, Parachondria , Articulipoma, and Chondropomium species. 22-23. Chondropoma
( Wetmorepoma ) morsecodex new species. Holotype, UF 446065, 13.0 mm. 24-25. Chondropoma ( Wetmorepoma ) wetmorei Bartsch,
1946. GTW 7161a, Isla Beata, Pedernales Province, Dominican Republic, 1 1.3 mm. 26-28. Parachondria ( Parachondria ) gettlemani
new species. 26-27. Holotype, UF 446066, 13.9 mm. 28. Paratype, NHMUK 20110336, 14.3 mm, from type locality. 29-31.
Articulipoma rliodei new species. 29-30. Holotype, UF 446068, 17.0 mm. 31. Paratype, OSUM 36515, 15-20 km NE of Cabo Rojo,
Pedernales Province, Dominican Republic, 14.5 mm. 32-33. Articulipoma loweanum (Pfeiffer, 1852). 32. Syntype, NHMUK 42/10,
“St. Domingo,” 17 mm. 33. UF 249153, W slope of Loma del Aguacate, Sierra Martin Gracia, Barahona Peninsula, Dominican
Republic, 14.8 mm. 34-35. Chondropomium blaineorum new species. Holotype, UF 446070, 13.0 mm. 36-37. Chondropomium
eusarcum saonaense new subspecies. Holotype, UF 446071, 13.2 mm. 38. Chondropomium eusarcum cataliniten.se (Bartsch, 1946).
GTW 7464a, Isla Catalinita, Dominican Republic, 12.2 mm. 39-41. Chondropomium lynx new species. 39-40. Holotype, UF 446072,
19.6 mm. 41. Paratype, OSUM 36519, Playa Los Patos, Barahona Province, Dominican Republic, 19.0 mm. 42. Chondropomium
swiftii weinlandi (Pfeiffer, 1862). GTW 7087c, 14 km N of Barahona, Barahona Province, Dominican Republic, 20.7 mm.
Page 8
THE NAUTILUS, Vol. 126, No. 1
NHMUK 20110336, adult (1 specimen); all from die type
locality; OSUM 36513, from 15 km S of Barahona, Barahona
Province, Barahona Peninsula, Dominican Republic, on
trees, 18.08° N, -7 1 .08° W. Adult (1 specimen).
Distribution: Known only from the area from
La Cienaga to Paraiso, in the narrow coastal plain
between the mountains and the sea in the eastern
Barahona Peninsula.
Habitat: On trees and rock fences; some specimens
were found near a waterfall (Figure 58).
Comparison with Other Species: Parachondria
gettlemani differs from P. kazikus (Bartsch, 1946) and
P. lindenianus sensu lato (Weinland, 1880) in lacking
any spiral sculpture outside of the umbilicus and in
the presence of false tufts. Parachondria kazikus was
described from “Dominican Republic” but has not
been further localized. Parachondria lindenianus has a
narrow range on the Massif de la Selle from Port-au-
Prince to Fonds-Verette, Haiti. Parachondria gettlemani
is only known from the area of La Cienaga, specifically
Virgen de San Rafael, ca. 15 km south of Barahona, on
the eastern coast of the Barahona Peninsula of the
Dominican Republic.
This new species resembles members of Bartsch’s
(1946) Lindenipoma, which was synonymized with
Parachondria by Watters (2006). This species is a good
example ol the reasoning behind that action.
Lindenipoma was described by a small suite of features
including the presence of spiral threads over the whorls.
Parachondria gettlemani has all of the characteristics of
Lindenipoma but lacks spiral sculpture except for a few
cords within the umbilicus. Bartsch (1946) further noted
that Lindenipoma lacked false tufting on the suture,
which P. gettlemani possesses. It was this patch-work
and inconsistent assortment of features that Bartsch used
to delimit some of his taxa that resulted in their synon-
ymy and inclusion into more broadly defined genera by
Watters (2006).
Etymology: Named for Alan Gettlenran of Merritt
Island, Florida, tireless and generous collector of land
snails, who collected part of the type lot.
Subfamily Tudorinae Watters, 2006
Genus Articulipoma Bartsch, 1940
Type Species: Chondropoma ( Articulipoma ) caroli
Bartsch, 1946, by original designation.
Discussion: Watters (2006) raised Articulipoma to
generic status. Species occur from the Chaine de la Selle
mountain range, including the Sierra de Baoruco in the
Dominican Republic, east almost to Punta Salinas. They
are characterized by axial lamellae that have an undulat-
ing or scalloped aspect. Most Articulipoma species are
quite rare in collections.
Articulipoma rhodei new species
(Figures 29-31)
Description (11 specimens): Shells 13.0-17.0 mm in
length, decollate (holotype 17.0 mm maximum length,
decollate, including peristome x 11.7 mm maximum
width, including peristome), translucent, ovate-conic,
umbilicus moderately narrow (holotype 13% of maximum
width), partially covered by inner margin of outer lip.
Protoconch whorls decollate in adults, IV2 whorls, scarcely
demarcated from teleoconch, smooth, minute but promi-
nent, a diffuse, brown, peripheral band may be present.
Teleoconch of 4 whorls, last 1/4* turn free from previous
whorl. Suture impressed and minutely channeled. Peri-
stome double, tear drop-shaped (holotype 6.8 mm diame-
ter maximum inner aperture height x 5.2 mm diameter
maximum inner aperture width; holotype 9.8 mm diameter
maximum outer peristome height, including auricle x 8.2 mm
diameter maximum outer peristome width). Outer lip
thin, expanded perpendicular to whorl, widest facing
umbilicus, which it partially covers, solute from previous
whorls in holotype but adnate in other examples. Promi-
nent triangular auricle and wide inner portion of outer lip
reflected forward, cup-like. Inner lip somewhat exserted,
simple, prominent. Spiral sculpture present only as faint,
wide undulations of axial threads, strongest near suture,
present as 4-5 barely perceptible cords within umbili-
cus. Axial sculpture of regularly spaced, narrow threads
(ca. 190 on final whorl), interstices smooth, having a
wavy or scalloped appearance. Suture broken by axial
lamella as minute cusps. Every 6-10 small cusps are
followed by 3-4 enlarged but unfused cusps. Teleoconch
silky in texture, bronze, purplish, tan, or orange with or
without a pattern of fine zigzag tan markings that
may coalesce to form bands. Suture brown with white
pseudotufts. Operculum thin, paueispiral with a granular
deposit. Anatomy and radula unknown.
Holotype: UF 446068.
Type Locality: 15 km NNE of Pedernales, on road to
Agua Negra, Pedernales Province, Barahona Peninsula,
Dominican Republic, on moist limestone cliff on the
southern slope of the Sierra de Baoruco. 18.10° N,
-71.64° W.
Paratypes: OSUM 36515, 15-20 km NE of Cabo
Rojo, off Road HH (2 specimens), ca. 18.09° N, -71.72°
W; BMSM 17946, 13 km NNE of Pedernales, on road to
Agua Negra. 18.15° N, -71.70° W (2 specimens);
NHMUK 20110337, 13 km NNE of Pedernales, on road
to Agua Negra. 18.15° N, -71.70° W (2 specimens); UF
446069, 9.5 km NNE of Pedernales, on road to Agua
Negra. 18.12° N, -71.72° W (4 specimens).
Distribution: Known from several localities on the
southern slope of the Sierra de Baoruco N of Pedernales,
Pedernales Province, Dominican Republic.
G. T. Watters, 2012
Page 9
Habitat: Associated with limestone outcrops. Speci-
mens have been found to ca. 1000 m. Dead spec-
imens have been found in cleared land adjacent to
agricultural fields.
Comparison with Other Species: Articulipoma
rhodei is most similar to A. cciroli (Bartsch, 1946), but
differs in being more rotund, more compact, and gener-
ally larger (13-17 mm in length in A. rhodei , 11-13 mm
in A. caroli). Articulipoma caroli occurs in a narrow
range on the northern side of the Massif de la Selle from
Croix des Bouquets to Bodarie, Haiti. Articulipoma
rhodei is only known from the region ca. 10-15 km north
of Pedernales on the southern side of the Sierra de
Baorueo, Dominican Bepublic. Articulipoma woodringi
Bartsch, 1946 differs from A. rhodei in lacking the pro-
nounced auricle and in having a very narrowly expanded
peristome on the inner lip (in contrast this is the widest
portion in A. rhodei). Articulipoma rhodei occurs in
association with Chondropoma maneilense Bartsch, 1946
and Chondropoma cnj.stallinum new species.
Etymology: Named for Homer Rhode of Englewood,
Florida, accomplished collector of land snails for nearly
60 years (he accompanied such luminaries as Archie
Jones and Edmund Winte), who collected portions of
the type lot of this species.
Articulipoma loweanum (Pfeiffer, 1852)
(Figure 32, 33)
Chondropoma loweanum Pfeiffer, 1852: 281.
Cyclostoma loweanum (Pfeiffer, 1852). Pfeiffer, 1854c: pi. 47,
figs. 15-16.
Chondropoma ( Chondropomium ) loweanum Pfeiffer, 1852.
Henderson and Bartsch, 1921: 60.
Chondropoma ( Articulipoma ) loweanum (Pfeiffer, 1852). Bartsch,
1946: 40, 48-49, pi. 8, fig. 3.
Articulipoma loweanum (Pfeiffer, 1852). Watters, 2006: 58, 337.
Distribution: Described from “Haiti,” the label to the
type (Figure 32, NI4MUK 42/10) reads “St. Domingo.”
Both localities refer to the island of Hispaniola rather
than to a more specific place. Bartsch ( 1946) had no
examples beyond a photo of the type. Specimens (UF
249153) have since been collected on the western slope
of Lonia del Aguaeate at 850 m in the Sierra Martin
Gracia E of the Barahona Peninsula, Barahona Province,
Dominican Republic. The type locality of Chondropoma
loweanum Pfeiffer, 1852 is herein restricted to Loma del
Aguaeate, Sierra Martin Gracia, Barahona Province,
Dominican Republic, ca. 19.10° N, -71.37° W.
Genus Chondropomium Henderson and Bartsch, 1921
Type Species: Chondropoma weinlandi Pfeiffer, 1862,
by original designation.
Discussion: As more material becomes available from
the Barahona Peninsula the lines between Chondropomium
and Chondropoma have become increasingly blurred.
Chondropomium was erected lor Chondropoma that
lacked spiral sculpture outside of the umbilical region.
But the suite of ‘'Chondropomium " -type features that
has emerged based on additional material — typically
large sized with a tear drop-shaped, narrowly expanded
peristome; high spire; usually unfused sutural serrations;
and undulating axial sculpture — is present in species,
unknown to Henderson or Bartsch, with spiral sculpture.
It is clear that a reorganization of species will be necessary
in the future based on phylogenetic studies. For now it
appears that Chondropoma marmoreum Watters and
Duffy, 2010 may also be a member of Chondropomium.
Chondropomium hooksi and C. ah/shae , both Watters
and Duffy (2010a), may belong to an unnamed genus,
as suggested by Watters and Duffy (2010a).
Chondropomium hlaineorum new species
(Figures 34, 35)
Description (2 specimens): Shells ca. 13.0 mm in
length, decollate (holotype 13.0 mm maximum length,
decollate, including peristome x 8.8 mm maximum width,
including peristome), solid, conic, umbilicus rather narrow
(holotype 11% of maximum width). Protoconch whorls
decollate in all specimens examined. Teleoeonch of
314 whorls, final whorl solute for last l/6fl1 of whorl.
Suture minutely channeled. Peristome single, tear
drop-shaped (holotype, 5.1 mm diameter maximum
inner aperture height x 3.5 mm diameter maximum
inner aperture width). Outer lip narrow, not expanded,
minute auricle present. Spiral sculpture present only as
7-8 very feeble cords within umbilicus. Axial sculpture of
narrow, closely spaced, low ribs, nearly obsolete on final
whorl, forming minute cusps at suture. Suture serrate.
Teleoeonch base color pale tan with narrow darker bands
having zigzag markings between them axially arranged;
umbilicus and peristome white. In paratype bands are
wider and markings more diffuse. Operculum, anatomy,
and radula unknown.
Holotype: UF 446070.
Type Locality: ca. 9.3 km NW of Manuel Golla, ca.
140 m, off Highway 44, Pedernales Province, Barahona
Peninsula, Dominican Republic, under rocks. 17.94° N,
-71.65° W.
Paratype: OSUM 36516, from the type locality
(1 specimen).
Distribution: Known only from the type locality.
Habitat: Specimens were found under rocks on a hill
slope with limestone outcrops next to recently cleared
agricultural land. No live-taken specimens are known.
Etymology: Named for Matt and Dona Blaine, who
helped collect the type lot.
Page 10
THE NAUTILUS, Vol. 126, No. 1
Comparison with Other Species: This small species
resembles Chondropoma eusarcum sensu lato (Pfeiffer,
1854), which on the southern coast is only known from
Isla Catalinita and Isla Saona, some 300 km E of
the range of C. blaineorum. The nominal subspecies,
C. eusacrum eusarcum , has not been localized, but
differs from C. blaineorum in having much coarser sculp-
ture, particularly on the body whorl where sculpture is
nearly absent in C. blaineorum, and in having a different
color pattern.
Chondropomium eusarcum sensu lato (Pfeiffer, 1852)
Chondropoma eusarcum Pfeiffer, 1852: 281.
Cyclostoma ( Chondropoma ) eusarcum (Pfeiffer, 1852). Pfeiffer,
1854a: 143.
Cyclostoma eusarcum (Pfeiffer, 1852). Pfeiffer, 1854e: pi. 48,
figs. 1, 2.
Chondropoma ( Chondropomium ) eusarcum Pfeiffer, 1852.
Henderson and Bartsch, 1921: 60.
Chondropomium eusarcum (Pfeiffer, 1852). Watters, 2006: 254.
Chondropomium eusarcum saonaense new subspecies
(Figures 36, 37)
Description (2 specimens): Shells 13.2-14.1 m in
length, decollate (holotype 13.2 mm maximum length,
decollate, including peristome x 8.1 mm maximum width,
including peristome), solid, conic, umbilicus rather nar-
row (holotype 13% of maximum width). Protoconch
whorls decollate in all specimens examined. Teleoeoneh
of 4 whorls, final whorl adnate. Suture minutely chan-
neled. Peristome double but outer and inner lips fused
to form a single lip, tear drop-shaped (holotype 5.5 mm
diameter maximum inner aperture height x 4.0 mm diam-
eter maximum inner aperture width). Outer lip narrowly
expanded, narrowest on inner margin, small, wide auricle
present. Spiral sculpture present only as 4-5 very feeble
cords within umbilicus. Axial sculpture of narrow, closely
spaced, low ribs, nearly obsolete on final whorl, forming
minute cusps at suture. Suture serrate. Teleoeoneh base
color glossy white with three broken spiral bands, one
above, one at, and one below periphery, composed of
brown rectangular spots, nearly forming a continuous
band at base; peristome white with bands showing
through. Operculum thin, paucispiral with a fine gran-
ular deposit. Anatomy and radula unknown.
Holotype: UF 446071.
Type Locality: Northeastern Isla Saona, Dominican
Republic, ca. 18.15° N, -68.60° W.
Paratype: OSUM 36517, from the type' locality
(1 specimen).
Distribution: Known only from the type locality.
Habitat: Specimens were found on a limestone cliff
after rain.
Comparison with Other Species: Chondropomium
eusarcum saonaense is somewhat similar to
Chondropomium eusarcum catalinitense (Bartsch, 1946)
(Figure 38), which occurs less than 4 km away on Isla
Catalinita. Chondropomium eusarcum catalinitense is
colored a uniform pinkish brown with only the faintest
trace of color bands (C. e. saonaense is white with obvious
bands), has an orange aperture (white in C. e. saonaense),
and has axial ribs that are much narrower and more
thread-like than those of C. e. saonaense.
Chondropomium eusarcum is among the smallest spe-
cies of the genus. If the subspecies described by Bartsch
(1946) truly are conspecific then it also has one of the
widest and most fragmented ranges of any Hispaniolan
annulariid. The nominal subspecies was described
from “Santo Domingo” and has not been localized.
Chondropomium eusarcum puertoplatense (Bartsch,
1946) from Puerto Plata is the only Chondropomium
known from the northern half of the island. Described
from Isla Catalinita, Chondropomium eusarcum
catalinitense is found 280 km away on the southeastern
end of Hispaniola (Watters (2006) speculated that
Bartsehs locality of Catalinita Island was an error for
Catalina Island, however, Catalinita Island is a separate
place). The subspecies described here is from Isla Saona,
adjacent to Isla Catalinita. Genetic analysis may eventually
determine that these four subspecies are not conspecific.
Chondropomium lynx new species
(Figures 39-41, 58)
Description (3 specimens): Shells 19.1-19.6 mm in
length, decollate (holotype 19.6 mm maximum length,
decollate, including peristome x 10.4 mm maximum
width, including peristome), solid, high-spired conic,
umbilicus rather narrow (holotype 11% of maximum
width). Protoconch whorls decollate in all specimens
examined. Teleoeoneh of 4 whorls, last 1/411' turn solute.
Suture minutely channeled. Peristome single, tear drop-
shaped (holotype 7.2 nun diameter maximum inner aper-
ture height x 5.0 mm diameter maximum inner aperture
width). Outer lip narrow, solute from previous whorls,
minute auricle present. Spiral sculpture present only as
three very feeble cords within umbilicus. Axial sculpture
of wide, closely spaced, low ribs, strongest on last 14 of
final whorl, ea. 90 on final whorl, forming minute cusps
at suture. Suture serrate. Teleoeoneh base eolor tan with
five broken spiral bands, three above, one at, and one
below periphery, composed of dark brown spots, blurred
and repeated; first teleoeoneh whorl dark brown; peri-
stome white. Operculum thin, paucispiral with a fine
granular deposit. Anatomy and radula unknown.
Holotype: UF 446072.
Type Locality: Virgen de San Rafael, ea. 200 m, ea.
7 km NNE of Paraiso, Barahona Province, Barahona
Peninsula, Dominican Republic, under rocks. 18.05° N,
-71.12° W.
G. T. Watters, 2012
Page 11
Paratypes: OSUM 3651S, Playa Los Patos, just off
Highway 44, Barahona Province, Dominican Republic,
under limestone rocks. 17.96° N, -71.19° W (1 speci-
men); OSUM 36519, from the type locality (1 specimen).
Distribution: Known from a narrow coastal range from
Virgen de San Rafael to Playa Los Patos, in the vicinity of
Paraiso, Barahona Province, Dominican Republic.
Habitat: Under limestone rocks, one specimen found
near a waterfall (Figure 58).
Comparison with Other Species: Chondropomium
lynx is part of the large complex of Chondropomium
species in the Barahona Peninsula. In its large size,
sculpture, and color pattern it resembles some iorms ol
C. swiftii (Figure 42). Chondropomium swiftii is a low-
land species distributed between the Massif de la Selle
to the south and the Sierra de Neiba to the north.
Chondropomium lynx occurs to the SE in the narrow
band of lowland between the ocean and the Sierra de
Baorueo. It differs from C. swiftii in having a unique
color pattern composed of spots that are overlaid and
repeated slightly out of alignment. It superficially resem-
bles C. swiftii harahonense (Bartseh, 1946) from ca. 25 km
to the north but does not have the nearly flat-sided
pupoid shape of the C. swiftii complex.
Etymology: Lynx rufus, the American bobcat or lynx;
the shell has a similar color pattern. Used as a noun
in apposition.
Genus Crossepoma Bartseh, 1946
Type species: Cyclostoma emilianum Weinlnnd, 1862,
by original designation.
Crossepoma vermiculatum sensu Into (Bartseh, 1946)
new combination
Discussion: For whatever reason Bartseh (1946) rarely
illustrated specimens with opercula although he always
included an account of the operculum in the description
of the species. Bartseh must not have had any specimens
with opercula of his Chondropoma vermiculatum
vermiculatum or any of the three other subspecies as he
did not mention any opercula in the descriptions. He
apparently relegated the species to Chondropoma
( Chondropomium ) based solely on shell features, a rea-
sonable conclusion given the shell's characteristics. But
examination of dive-taken material (GTW 7088b) reveals
an operculum consisting of a pseudolamellum that does
not completely cover the basal, corneous portion — the
hallmark of Crossepoma. Bartseh (1946: 72) defined a
pseudolamellum as a ‘ flat plate formed by the expansion
and fusion of the retractively curved opercular riblets on
the outer edge.” Although Bartseh (1946) considered
sutural tufts a characteristic of Crossepoma , this feature
does not occur in all taxa (i.e., C. emilianum insulanum
Bartseh, 1946) and it does not occur in C. vermiculatum.
These taxa (Chondropoma ( Chondropomium ) vermiculatum
domingense , C. v. nubilum , C. v. sallei , and C. v.
vermiculatum, all Bartseh, 1946) are herein allocated
to Crossepoma Bartseh, 1946. This is the first species
of Crossepoma to be found in the Dominican Republic.
However, other Haitian species of Crossepoma occur
in the Chaine de la Selle mountain range, which con-
tinues into the Dominican Republic as the Sierra de
Baorueo where C. vermiculatum is found. Other genera
have the same distribution and this range extension is
not unexpected.
Crossepoma vermiculatum sensu lato (Bartseh, 1946)
ranges along the eastern Barahona Peninsula from
Cabral south to Trujin, although two of the four sub-
species were unlocalized in Bartsch’s 1946 treatment. A
locality for one of these two, Crossepoma vermiculatum
sallei (Bartseh, 1946), is recorded below. This is the
most eastern species of Crossepoma. It apparently inhab-
its lowlands.
Crossepoma vermiculatum vermiculatum (Bartseh,
1946) new combination
(Figures 43, 57)
Chondropoma ( Chondropomium ) vermiculatum vermiculatum
Bartseh, 1946: 29-30, pi. 3, fig. 1 .
Chondropomium vermiculatum vermiculatum (Bartseh, 1946).
Watters, 2006: 63, 533.
Distribution: These examples (GTW 7088b) came
from the area of Colonia Juancho off Highway 44, NE
of Oviedo, at 30 nr along an unpaved road lined with
trees, fences, and occasional rocks, 17.85° N, -71 .34° W.
Some individuals were found on trees (Figure 57). This
area is just north of the type locality of Trujin.
Crossepoma vermiculatum sallei (Bartseh, 1946)
new combination
(Figure 44)
Chondropoma ( Chondropomium ) vermiculatum sallei Bartseh,
1946: 30, pi. 3, fig. 6.
Chondropomium vermiculatum sallei (Bartseh, 1946). Watters,
2006: 533.
[Distribution: This subspecies was described from
“Haiti” although Bartseh (1946) clearly did not believe
that locality was correct. A specimen from Cabral (GTW
7088a), ca. 13 lan WNW of Barahona, at ca. 40 m, matches
the type of C. vermiculatum sallei. The type locality of
Chondropoma (Chondropomium) vermiculatum sallei
Bartseh, 1946 is herein restricted to Cabral, Barahona
Province, Dominican Republic, ca. 18.25° N, -71.22° W.
This is the northern-most subspecies.
Genus Licina Gray, 1847
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THE NAUTILUS, Vol. 126, No. 1
Figures 43-58. Crossepoma , Licina, Chondropomium, and Chondropoma species and habitats. 43. Crossepoma vermiculatum
vermiculatum (Bartsch, 1946). GTW 7088b, 8 km NE of Oviedo, Pedernales Province, Dominican Republic, 15.0 mm. 44. Crossepoma
vermiculatum sallei (Bartsch, 1946). GTW 7088a, Cabral, Barahona Province. Dominican Republic, 18.6 mm. 45-47. Licina bartschi
new species. 45-46. Holotype, UF 446073, 20.1 mm. 47. Paratype, OSUM 36520, 20.6 mm, from type locality'. 48, 53, 55.
Chondropomium nobile (Pfeiffer, 1852). GTW 7089b, Virgen de San Rafael, Barahona Province, Barahona Peninsula, 26.9 mm. 53.
Living animal from same locality. 55. Habitat (photo courtesy of Alan Gettleman). 49, 54. Chondropomium pumilum Watters and
Duffy, 2010. GTW 7172b, 10 km NE of Las Mercedes, Pedernales Province, Dominican Republic, 19.9 mm. 54. Living animal from
same locality. 50. Chondropoma ( Wetmorepoma ) oculeum Watters and Duffy, 2010. Holotype UF 420737, 9.0 mm. 51, 52, 56.
Chondropoma marmoreum Watters and Duffy, 2010. Holotype UF 420735, 18.3 mm. 56. Habitat (photo courtesy of Alan Getdeman).
57. Habitat of Crossepoma vermiculatum vermiculatum (Bartsch, 1946). Area of Colonia Juancho, NE of Oviedo, Pedernales Province,
Dominican Republic (photo courtesy of Alan Gettleman). 58. Habitat of Parachondria gettlemani new species and Chondropomium
lynx new species, Virgen de San Rafael, Barahona Province, Dominican Republic (photo courtesy of Alan Gettleman).
G. T. Watters, 2012
Page 13
Type Species: Nerita labeo Miiller, 1774, by original
designation.
Discussion: Bartsch (1946) separated his genus
Kisslingia from Licina based solely on the presence of a
single peristome in the former and a double peristome in
the latter. Based on the fact that among species in other
genera ( Chondropoma , for example), and occasionally
even within species, taxa may have both single and dou-
ble peristomes, Watters (2006) synonymized Kisslingia
with Licina.
Licina bartschi new species
(Figures 45-47)
Description (4 specimens): Shells 18.4-20.6 mm in
length, decollate (holotype 20. 1 mm maximum length,
decollate, including peristome x 12.7 mm maximum
width, including peristome), solid, conic, umbilicus rather
narrow (holotype 12% of maximum width). Protoconch
whorls decollate in adults, consisting of 1 Vz smooth whorls,
well-delimited from teleoconeh. Teleoeoneh of 4 whorls,
final whorl solute just before peristome. Suture deeply
channeled. Peristome single, thick, tear drop-shaped
(holotype 7.6 mm diameter maximum inner aperture
height x 5.5 mm diameter maximum inner aperture
width). Outer lip narrow, barely expanded on outer side,
auricle absent. Spiral sculpture present only as 3-5 very
feeble cords below suture rendering axial sculpture
scalloped, 7-9 cords in umbilicus. Axial sculpture of nar-
row, fine ribs separated by 2-3 times their width, ca. 150
on final whorl, forming minute cusps at suture. Suture
serrate. Teleoconeh base color glossy pale tan with a
complex pattern of dense, brown, very fine, zigzag mark-
ings axially aligned, sometimes forming interrupted
spiral bands on base and within umbilicus; growth rests
marked by axial rows of tiny dark spots; suture with pat-
tern of alternating white and dark brown patches; mark-
ings persist on both sides of peristome and are visible
through inside of shell. Operculum paucispiral with a
pseudolamella that extends 3/5ths of way to outer mar-
gin, corneous portion with a fine granular deposit. Anat-
omy and radula unknown.
Holotype: UF 446073.
Type Locality: 9.5 km ENE of Las Mercedes,
Pedernales Province, Dominican Republic, on a mountain
top at 1,300 m. 18.12° N, -71,57° W.
Paratypes: BMSM 17947, (1 specimen); OSUM
36520, (1 specimen); NHMUK 20110338, (1 specimen);
all from the type locality.
Distribution: Known only from the type locality.
Habitat: In pine forest with large rock outcroppings.
Comparison with Other Species: Licina bartschi
resembles other members of Bartsch’s Kisslingia group
(those having a simple peristome), distributed from east to
west: Licina clencki (Pilsbry, 1933), Licina baharucensis
(Bartsch, 1946), Licina poloensis (Bartsch, 1946), Licina
bartschi , and Licina hincliensis (Bartsch, 1946). Licina
clench i (Pilsbry, 1933) is known only from the Barahona
area and most closely resembles L. bartschi , even though
it is the most geographically distant (50 km) of the penin-
sular Kisslingia group. Licina bartschi differs in being
slightly larger (18-20 vs. 15-18 mm in length), having a
higher spire, having more rounded whorls, and possessing
thinner and more widely separated axial ribs that have a
scalloped or undulating appearance over the entire body
whorl. Licina baharucensis (Bartsch, 1946) occurs in the
eastern half of the Barahona Peninsula in the Sierra
de Baharuco. It is smaller, more ovate, more coarsely
sculptured, and the last whorl is more solute than in
L. bartschi. Licina poloensis (Bartsch, 1946) is known
only from Polo, located in a valley in east central
Barahona Peninsula. It is more broadly turbinate than
L. bartschi , with a more circular aperture (teardrop-
shaped in L. bartschi). Licina liinchensis (Bartsch, 1946)
is from Basin Sin (Zim) near Hinche in the Massif du
Nord of central Haiti and is probably not closely related
to the remaining Kisslingia -like taxa in the Barahona
Peninsula. It is more turbinate in outline than L. bartschi
and has a spotted rather than vermiculated color pattern.
No other member of the Kisslingia group has such an
intricate color pattern as does L. bartschi.
Etymology: Named for Paul Bartsch (1871-1960) of
the U.S. National Museum (now National Museum of
Natural Histoiy, Smithsonian Institution), who wrote the
definitive works on the Annulariidae (Torre and Bartsch,
1938, 1941 ; Bartsch, 1946), describing nearly 800 taxa.
Often vilified as a splitter, his works were nevertheless
Herculean, thorough, and professional. It is surprising
and disappointing that only one taxon of this family to
which he devoted so much of his time (Annnlarella
torrebartschi Jaume, 1984), had ever been named to
honor him.
Additional Records
Chondropoma ( Welmorepoma ) oculeum Watters
anti Duffy, 2010. Originally described from 14.5 km N
of Cabo Rojo, new records are from ca. 7 km S of the
type locality (GTW 7166b, 7166c, Figure 50). This spe-
cies seems to have a narrow range confined to the
Pedernales/Cabo Rojo area on the west coast of the
Barahona Peninsula.
Chondropomium nobile (Pfeiffer, 1852) (Figures 48,
53, 55). One of the largest of the Annulariidae, this
species was known from a few records from the coastal
town of Paradis (? Paralso), 26 km SSW of Barahona,
and another record from 10 km S of Barahona. Records
reported here are for Virgen de San Rafael, less than
7 km up the coast from Paraiso (GTW 7089b). Spe-
cimens were common at 200 m under rocks in dirt on
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THE NAUTILUS, Vol. 126, No. 1
hillside crests in a scrub cactus area. It appears that this
species has a very narrow range on the coastal eastern
edge of the Sierra de Baoruco. A live individual is illus-
trated here (Figure 53).
Chondropomium pumilum Watters and Duffy, 2010
(Figures 49, 54). Described from 19-32 km N of Cabo
Rojo, new records give a more precise locality at 10 km
NE of Las Mercedes at 1,300 m in pine forests, 18.12° N,
-71.56° W (GTW 7172b). It co-occurs with Chondropoma
vanattae polychroma new subspecies.
Chondropomium swift ii weinlandi (Pfeiffer, 1862).
This subspecies occupies a rather wide range in the xeric
lowlands between the Massif de la Selle and the Sierra
de Neiba, occurring from Thomazeau, Haiti, to Azua,
Dominican Republic. One live example was found
ca. 14 km N of Barahona at the base of the Sierra de
Neiba at 37 m elevation under rocks and plant roots in
xeric shrub (GTW 7087c, Figure 42). Other live-taken
specimens have been found at Mella (GTW 7087a) and
Fordi Negro (GTW 7087b).
Chondropoma marmoreum Watters and Duffy,
20 1 0. This species is now known from several sites along
the foothills ol the southern Sierra de Baharuco (GTW
7170c-f). It occurs under rocks in xeric areas where it is
locally common (Figures 51, 52, 56).
Leiabbottella galaxius Watters, 2010. Described
from the Sierra de Samana of the Samana Peninsula,
additional records extend its range westward along the
north coast to at least Sosua, Puerto Plata Province (UF
249107, 249112). This is a range extension of over
100 km and illustrates how poorly the annulariid fauna
ol the northern coast is known.
ACKNOWLEDGMENTS
I am grateful for the opportunity to study specimens
from these new localities generously made possible by
Matt and Dona Blaine, Glenn Duffy, Alan Gettleman,
Randy Hooks, Homer Rhode, and Jesus Santana
Benitez. Fred Thompson, John Slapcinsky, and Gustav
Paulay graciously allowed me access to the UF
collection. The manuscript was greatly improved by an
anonymous reviewer.
LITERATURE CITED
Bartsch, P. 1946. The opereulate land mollusks of the family
Annulariidae of the island ol Hispaniola and the Bahama
Archipelago. Bulletin of the U.S. National Museum 192:
264 pp., 38 pis.
Dali, W. H. 1905. An arrangement of the American Cyelo-
stomatidae, with a revision of the nomenclature. Proceedings
of Malaeological Society of London 6: 208-210.
Gray, J.E. 1847. A list of the genera of Recent Mollusca, their
synonyma and types. Proceedings of the Zoological Soci-
ety for 1847 (15): 129-219.
Henderson, J.B. and P. Bartsch. 1921. A classification of the
American opereulate land mollusks of the family
Annulariidae. Proceedings of the United States National
Museum 58 (2327): 49-82.
jaume, M.L. 1941. Reetificaciones a nonrbres de molluscos
terrestres. Miseelanea Zoologica, Academia de Ciencias
de Cuba (20): 1-4.
Pfeiffer. L. 1847. Uebersieht aller bekannten Arten von Cyclo-
stomaceen. Zeitsehrift fur Malakozoologie 4: 101-112.
Pfeiffer, L. 1852. Monographia pneumonoporum viventum.
T. Fischer, Cassellis, 435 pp.
Pfeiffer, L. 1854a. Descriptions of twenty-four new species of
land shells, collected by M. Salle on the island of St.
Domingo, from Mr. Cuming’s collection. Proceedings of
the Zoological Society for 1852 (20): 138-144, pi. 13.
Pfeiffer, L. 1854b. Die gedeckelten Lungensehneeken.
(Helicinacea et Cyclostomacea). Systematisches Conehylien-
Cabinet von Martini und Chemnitz (Kiister ed.). Part 19.
Installment 133: pis. 37-42, pp. 269-308.
Pfeiffer, L. 1854c. Die gedeckelten Lungensehneeken. (Helicinacea
et Cyclostomacea). Systematisches Conchylien-Cabinet von
Martini und Chemnitz (Kiister ed.). Part 19. Installment
136: pis. 43-48, pp. 309—356.
Pilsbry, Id. A. 1933. Santo Domingo land mollusks collected by
Samuel C. Pease, 1932, and by A. A. Olsson, 1916. Proceedings
of the Academy of Natural Sciences 85: 121-162, pis. 6-11.
Watters, G.T. 2006. The Caribbean land snail family
Annulariidae: a revision of the higher taxa and a catalog
of the species. Backhuys Publishers, Leiden, 557 pp.
Watters, G.T. 2010. New taxa of Annulariidae from Dominican
Republic (Gastropoda: Littorinoidea). VisayaSl: 16-20.
Watters, G.T. and G. Duffy. 2010a. New species of Annulariidae
(Gastropoda) from the Bahamas and Dominican Republic.
Novapex 11: 1-12.
Watters, G.T. and G. Duffy. 2010b. Rolleia oberi , new species-
first record of the genus from the Dominican
Republic, with a lectotype designation of Cyclotus martensi
Maltzan, 1888 (Gastropoda: Annulariidae). The Nautilus
124: 185-187.
THE NAUTILUS 126(1): 15-24, 2012
Page 15
Americardia lightboumi new species and A. columbella new
species compared to A. media (Linnaeus, 1758), A. speciosa
(A. Adams and Reeve, 1850), and the extinct A. columba
(Heilprin, 1886) (Bivalvia: Cardiidae)
Harry G. Lee
4132 Ortega Forest Drive
Jacksonville, FL 32210 USA
[email protected]
Markus Huber
Winterthurerstrasse 190
8057 Zurich, SWITZERLAND
[email protected]. ch
ABSTRACT
A comparatively small and many ribbed Americardia living
from Bermuda, off Florida to Caribbean Panama and Brazil is
described and named after J.R.H. (Jack) Lightbourn. The new
species has more anterior ribs and reaches a significantly
smaller size than the two other Recent western Atlantic spe-
cies. It also lives in deeper water, particularly in the northern
part of its distribution. It is well represented in Bermuda and
Brazil but has been confounded with the much larger, lower
ribbed and shallow water Americardia media , best known from
the West Inches. A comparison of the Pliocene Americardia
columba (Heilprin, 1886) with Recent material revealed gross
similarities, yet its marked distinct rib sculpture did not allow
consideration of the latter material conspeeific. Tims Americardia
columbella, living from North Carolina to Yucatan, Mexico, is
also described as new herein. These two new species are com-
pared with the type species, A. media , living from Florida to
Ascension, and its congener A. speciosa, living exclusively at
St. Helena. Instead of one, in Florida there are three and in
Brazil two, extant Americardia. There is only one species living
in the waters of Bermuda, North Carolina, Ascension, and
St. Helena, but each of these populations represents a distinct
Americardia species. Based on the most recent studies, the
application of Americardia is genetically, biogeographically, and
morphologically supported for this Atlantic and Eastern Pacific
species complex. Consequently, Ctenocardia is restricted to
the superficially similar but spiny Indo-Pacific species-group.
According to Linnaeus, 1758 and 1767, his “Ofceano] Indico”
type of Cardium medium should be at Uppsala only. However,
no type material or trace thereof could be found there. In reality,
the true identity of the Linnaean species is simply unknown and
con specificity with the Indo-Pacific Ctenocardia fornicata could
not be excluded. To stabilize one of the “best known” Caribbean
cardiids a neotype from material in the Linnean Society, London
is selected, and McLeans supposed type locality, Cuba, Havana
is herein confirmed.
The Natural History Museum, London [NHMUK] type
material of Cardium speciosum is comprised of four spec-
imens, which, however, represent two species. Only one
matches the original description and is confirmed as the holo-
type. The other three specimens were added after 1890.
Additional keywords: Western Atlantic, Bermuda, Florida,
Brazil, St. Helena, Ctenocardia, neotype Cardium medium
Linnaeus, 1758, Cardium fornication G.B. Sowerby II, 1840.
INTRODUCTION
In conformity with all cardiid experts, e.g. Vidal (2000:
642), and ter Poorten (2005: 6), and most regional spe-
cialists, e.g. Clench and Smith (1944), Abbott (1974:
484), and Mikkelsen and Bieler (2008: 298), Huber
(2010: 300) also recognized only one large Americardia ,
living from North Carolina to Ascension, exceptionally
reaching 60 mm in the Bahamas (Redfern, 2001: 226).
This species was uniformly understood as highly vari-
able, living from shallow to rather deep water and being
widely distributed, including North Carolina (Porter
and Houser, 1994: 19), Bermuda (Jensen and Pearce,
2009: 344), Brazil (Rios, 2009: 1479), and Ascension
(Rosewater, 1975: 33).
However, Lee (2009: 33) instead recognized
Americardia columba from NE. Florida as valid recent
species and separated it from the West Indian A. media.
Lee further mentioned some “atypical populations" of
Americardia from Bermuda and Brazil. Subsequently,
the present authors combined their respective Americardia
holdings resulting in nearly 30 lots with over 100 spec-
imens collected throughout the Western Atlantic. Imme-
diately it became obvious that not only two, but in
fact three, distinct Recent species from Florida were
involved. These differed in the prominence of the poste-
rior sulcus, maximum size, number of ribs, color, and
notably also in biogeography and habitat. Subsequent
review of material at the Florida Museum of Natural
Histoiy (FLMNII, Gainesville) by the senior author
revealed 27 and 28 lots respectively of the new taxa
described below. In addition, he studied 12 lots of the
latter new taxon in the E.F. Garcia Collection.
Two distinct Americardia species were detected in a
small collection of Pliocene fossils from Florida. This led
to further inquiries into that fauna, including a review
of the holdings at the FLMNH. Moreover, the “well
known” Americardia having been sorted out, the identity
of Ascension Island specimens was at stake inasmuch as
the obscure Cardium speciosum A. Adams and Reeve,
1850 recognized by E.A. Smith (1890a) as living in the
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THE NAUTILUS, Vol. 126, No. 1
S. Atlantic, is a valid congener (Huber, 2010: Chapter 5).
Thus, additionally the identity and exact distribution of
A. speciosa had to be clarified. Moreover, the N11MPIK
type material contained four specimens and two species,
which necessitated further inquires and solutions.
Distinction from such a well known species as
Americardia media obviously requires an unambiguous
opinion on the true identity of that Linnaeus’ species. Most
surprisingly, this investigation led to the fact that the
London material identified by Hanley (1855) cannot rep-
resent Linnaeus’ true type. In Uppsala, where, according
to Linnaeus (1758 and 1767), his type should be, no spec-
imen or traces thereof could be located. Consequently,
the true identity of Cardium medium Linnaeus, 1758 from
“Oceano Indico” is simply unknown; one could not exclude
it being an earlier name for the well known Indo-Pacifie
Ctenocardia fomicata. The “firm” understanding of this
“well known” American species proved neither firm nor
even known, but ultimately based on mere assumptions.
As a final consideration, many modern authors use
instead of Americardia , as proposed by Stewart (1930)
and recognized by Vidal (2000), the Indo-Pacific
Ctenocardia for this group. This is also supported by the
genetic analyses performed by Kirkendale (2009).
Consequently, the authors went beyond the simple
question of whether indeed a second Americardia lives
in Florida, to investigate all Atlantic species and their
generic allocation, including fossil records. A neotype
for A. media had to be designated, the holotype for A.
speciosa had to be identified and confirmed, and two
species had to be described as new to science.
All these issues are addressed below, and two species,
Americardia lightboumi and Americardia columbella,
are described as new to science and compared with their
two Recent Atlantic congeners.
Abbreviations used in the text are: UF: Florida
Museum of Natural History, Gainesville, Florida; BMAZ:
Bermuda Aquarium, Museum and Zoo, Hamilton,
Bermuda; NHMUK: Natural History Museum, London;
USNM: National Museum of Natural Histoiy, Smithsonian
Institution, Washington, DC.
SYSTEMATICS
Family Cardiidae Lamarck, 1809
Subfamily Fraginae Stewart, 1930
Genus Americardia Stewart, 1930
Remarks: Type species by original designation is
Cardium medium Linnaeus, 1758 as understood by
“Reeve pi. 6 fig. 30”. Stewart explicitly noted: “Should
the identity of Cardium medium Linne with C. medium
Reeve ever be questioned, the name Americardia is to go
with the latter ". Obviously, Stewart himself had notice-
able doubts on the true identity of Linnaeus’ species.
Linnaeus (1758: 678, 1767: 1122) gave 0[ceano]
Indico as type locality. However, this was changed early
on by Gmelin (1791: 3247) to Oceano Americano. Sub-
sequently, McLean (1939: 167) even specified Havana,
Cuba. Neither of the latter authors saw or designated a
type. Neither discussed or excluded the quite common,
spined, white-purple Indo-Pacific Ctenocardia fomicata
(G.B. Sowerby II, 1840). This species was unknown to
Gmelin, but matches Linnaeus’ original description
precisely at least in context as sp. 61, in shape, in colors,
in biogeography, and eventually even in sculpture “sulcis
laevibus absque aculeis, angulisque obsoletis”. Why
should Linnaeus mention a not present feature within
the very few words available? Is it indeed “without
spines” or is it instead, “slightly angular ribs, smooth,
where the spines are abraded”?
Linnaeus had no personal Cardium medium material
but described this species without any references from
the Swedish Queen Ulrike collection “M.L.U.” (1758)
and “M.L.U. 485 n. 34” (1767). It was not until later
(Gmelin, 1791) that references leading to today’s inter-
pretation of the species were provided. Linnaeus based
his species solely on a Swedish specimen, of which he
apparently could find no satisfactory figure. Lister (1685:
pi. 316, fig. 153) as proposed by Clench and Smith (1944:
21) was well known to Linnaeus, but not explicitly cited,
so this leetotype designation is invalid. Linnaeus clearly
excluded any type material in his personal collection
(now in the Linnean Society, London) as he left no indi-
cation, which was his custom, with that material. Despite
this, Hanley (1855: 47) isolated two unmarked valves in a
box in Linnaeus' personal collection, though noting some
differences. These, a right and a left valve, are illustrated
online as Cardium medium <www. Linnean-online.org/
16920> (Figures 18, 19). They conform well to spec-
imens collected in Cuba and to the common understand-
ing of Americardia media but definitely do not represent
true Linnaean material contemporary with the descrip-
tion of C. medium. No doubt these specimens were
introduced later, possibly by the son of Linnaeus or by
the subsequent owner of the collection. Sir J.E. Smith.
Quite unfortunately, the Museum of Evolution, Upp-
sala University, could not deliver the requested species.
E. Sjolin. Curator, wrote 19 October, 2010: “Regarding
your request for photos of Cardium medium Linnaeus,
1758; I’m sorry to inform you that 1 couldn’t find this
specimen in the Museum of Evolution, or information of
its whereabouts.” Thus, one of the “best known” Western
Atlantic cardiids has no type material; even the type local-
ity and the true identity of C. medium are unsubstantiated.
To eliminate any doubts of the identity of Cardium
medium Linnaeus, 1758, the unmarked right valve from
the Linnean Society London, S.P. Dance label image
G-M 0010260, well illustrated at <www.Linnean-online.
org/16920/> is selected as neotvpe (Figure 19).
McLean’s ( 1939) assumed type locality, Havana, Cuba is
supported. As such, Cardium medium Linnaeus, 1758
conforms, as anticipated by Stewart, to Reeve’s interpre-
tation and becomes the true type species of Americardia .
No solution other than that presented here would
better serve taxonomic stability and the consistent inter-
pretation of C. medium over the last 200 years. For some
H.G. Lee and M. Huber, 2012
Page 17
time the Linnaean species was retained in Cardium-,
later it was transferred to Fragum (e.g. E.A. Smith,
1890a, b). Stewart (1930: 267) separated Americardia at
the subgenus level from the New World Trigoniocardia
and designated Cardium medium as its type. Stewart
explicitly included the Panamic Cardium biangulatum
Broderip and G.B. Sowerby I, 1829 but not the Carib-
bean Cardium guppyi Thiele, 1910 in Americardia.
Stewart somewhat questioned his own snbgenerie place-
ment of Americardia under Trigoniocardia but clearly
separated Americardia from the spiny sculptured Indo-
Pacific Ctenocardia II. and A. Adams, 1857. In fact,
narrow spined ribs are typical for the Indo-Pacific
Ctenocardia, strongest expressed in the type species
Cardium hystrix Reeve, 1844 non Lightfoot, 1786
( =Cardium virgo Reeve, 1845) weakest in Ctenocardia
gustavi Vidal and Kirkendale, 2007. The western Atlantic
Americardia- group, including the two well known
Panamic A. biangulata (Broderip and G.B. Sowerby I
1829) and A. planicostata (Broderip and G.B. Sowerby I
1833) display broader and smoother ribs. In general,
Americardia also grow larger and more solid than
the usually smaller and more fragile Indo-Pacific
Ctenocardia. Most conspicuous among the eonchologieal
differences is the ornamentation of the ribs: corn-
marginal bars/knobs vs. spines respectively.
Vidal (2000) applied Americardia at generic level,
distinct from the New World Trigoniocardia , and also
separated from the Indo-Pacific Ctenocardia. Due to
morphological affinities, however, Vidal placed both
genera in the same group 2 within Fraginae. Ter Poorten
(2005) and Mikkelsen and Bieler (2008) synonymized
Americardia into Ctenocardia and combined the New
World with the Indo-Pacific species. Lee (2009) and
Huber (2010) did not follow those latter works, but
applied Americardia to American species, generically
distinct from the Indo-Pacific Ctenocardia.
Finally, Kirkendale (2009) genetically studied Fraginae.
Her results are consistent with Stewarts and Vidal’s view.
Americardia media is in all clades more closely related
to Trigoniocardia than Ctenocardia. Aside from disjunct
biogeography and distinct morphology, the genetic data
are consistent with the view here presented. The morpho-
logical ly distinct "Ctenocardia" victor was recently recog-
nized as unrelated Freneixicardia by ter Poorten (2009:
31). Pending further studies with additional Ctenocardia
species and resolution of some inconsistent evidence
of paraphyly among these generic units Americardia
is herein treated as a valid genus within the cardiid
subfamily Fraginae.
Americardia lightbourni new species
(Figures 1-11, 21)
Diagnosis: Rather small and thin-shelled, deeper
water Americardia with very numerous ribs, narrow
interstices and a marked posterior sulcus.
Description: Shell thin and rather fragile, inflated
to strongly inflated in older specimens; moderately to
strongly oblique, with a rather strong posterior sulcus.
Smallest Atlantic Americardia , exceptionally up to 31 mm,
but usually around 20 mm. Anterior margin obliquely
rounded; ventral margin oblique, descending toward
carina, almost imperceptibly sinuate in the posterior
third. Carina sharply expressed, moderately rounded.
Shell equivalve and inequilateral with rounded, weakly
prosogyrous umbones placed in anterior half and extend-
ing above dorsal line. Shell surface densely radially sculp-
tured, with 27-31, often 28 or 29, anterior ribs. Anterior
ribs low, rather flat, and separated by narrow inter-
spaces less than 14 rib in breadth. Beneath intritacalx ribs
are glossy smooth without spines, or elevated scales.
Dehiscent intritacalx composed of a whitish calcareous
layer, veiy densely and regularly commarginally lined,
often forming small, irregularly arranged knobs. Liga-
ment external, very small, dark brown, situated just
posterior to umbones. External coloration whitish with
irregularly arranged rose and brownish blotches and
streaks. Internal shell surface porcellaneous white,
usually suffused with yellow and occasionally with pur-
plish streaks. Pallial line entire, positioned veiy close
to the ventral margin and connecting two medium-sized,
nearly homomyarian, subovate scars. Shell margin finely
crenulate ventrally, but coarser on posterior margin.
I linge line strong and rather thick, with two cardinal
teeth and two prominent lateral teeth. Posterior cardi-
nal tooth in right valve and anterior lateral in left valve
strong. Valves close tightly.
Variations: Size, thickness and color intensity vaiy
remarkably within its range. Bermuda and S. Florida spec-
imens are in general smaller with maximum sizes around
21 mm, paler and thinner, characters possibly induced by
their deeper habitat. Louisiana, Caribbean Panama, and
Brazilian specimens grow larger, with maximum sizes up
to 32.7 mm, are often more solid, usually more brightly
colored, often deep yellow inside. In the Caribbean and
Brazil the species lives in much shallower water. Brazilian
shells may be more elongate than the others. However,
base colors and rib characters remain identical, and their
habitat is still deeper than that of the type species.
Etymology: The new species name honors John (Jack)
R.H. Lightbourn, who pioneered deep-water shelling
in his native Bermuda and who, with the late Arthur T.
Guest, discovered the holotype.
Type Locality: Bermuda Island, S. of Castle Roads,
dredged in 82 m, collected by J.R.II. Lightbourn and
A. T. Guest, September, 1976.
Type Material: Holotype: UF 447279, one paired
specimen, 20.5 mm [Figures 1 2], from the type locality,
ex Coll. H.G. Lee. Paratype suite 1: BMAZ 2011 274
019, two paired specimens and one single right valve,
20.7 [Figures 6, 7], 20.9, and 16.3 mm respectively, from
type locality, ex Coll. H.G. Lee. Paratype suite 2: Coll.
M. Huber, QQ2691, one paired specimen, 25.3 mm.
Page 18
THE NAUTILUS, Vol. 126, No. 1
Figures 1—11. Americardia lightbourni new species. 1-2. Holotype Bermuda 20.5 mm. 1. External view. L, R valve. 2. Internal
view. L, R valve. 3-5. Paratype suite 5, Brazil, 23.5 mm. 3. External view L, R valve. 4. Internal view, L, R valve. 5. Posterior view,
paired valves. 6-7. Paratype suite 1, Bermuda, 20.7 mm. 6. External view, L, R valve. 7. Internal view L, R valve. 8-9. Paratype
suite I, Brazil, 31.3 mm. 8. External view, L, R valve. 9. Internal view, L, R valve. 10-11. Paratype suite 6, Louisiana, 32.0 mm.
10. External view, L, R valve. 11. Internal view L, R valve. Seale bar (all views) = 1 cm.
Caribbean Panama, off Portobello, 15 m, coral sand,
dived February, 2002. Paratype suite 3: Coll. M. Huber,
QQ269, one paired specimen, 21.1 mm, N. Brazil, Bahia,
Alcobaga, 10-20 m, coral sand, dived March, 2005.
Paratype suite 4: Coll. H.G. Lee, two paired speci-
mens 13.1 and 15.8 mm, S. Florida, SW. Dry Tortugas,
24.736°N, 85.639°W, dredged 62.5-66.5 m, collected by
E.F. Garcia, NSF I 13. Paratype suite 5: Coll. H.G. Lee,
two paired specimens, 23.5 and 31.3 mm [Figures 3, 4, 5;
8, 9 respectively], N. Brazil, Bahia, Alcobaga, 10-20 m,
coral sand, dived February, 1999. Paratype suite 6:
Coll. E.F. Garcia 24405, two paired specimens 21.1 and
32.0 mm [Figures 10, 11], Louisiana 28°5.85'N, 91°1.28'W,
dredged 68.3 m, collected by E.F. Garcia 3 July, 2003.
Distribution: The new species is widely distributed
within the Caribbean Province, known from Bermuda,
S. Florida, the Gulf of Mexico, Caribbean Panama, and
Brazil. Americardia lightbourni is uncommon in the
north but quite commonly found in Brazil.
Habitat: Americardia lightbourni is a sublittoral spe-
cies, living from 10-100 m depth. In the southern part of
its range, it is found in shallower water, 10-70 m,
whereas in the north it is usually found at 50-100 m.
The substrates are sand, coral-sand, and mixed carbon-
ate rubble (off Louisiana). It is of relevance that in all
collections studied the type species, A. media, is limited
to much shallower habitats.
H.G. Lee and M. Huber, 2012
Page 19
Discussion and Comparison: Americardia lightboumi
can be separated from A. media by its higher rib number,
and consequently by finer internal shell surface crenula-
tion. The interstices are narrower, the posterior sulcus is
stronger, and the valves are in general thinner and more
fragile. In addition, A. lightboumi reaches only half the
size of A. media. Whereas A. media is a common, shallow
water species, often found beached, A. lightboumi is
deeper dwelling, especially in the northern part of its
distribution, and is less frequently found. The distribution
of A. lightboumi surpasses the known range of A. media ,
which does not reach Bermuda or central Brazil.
Similarly, the new species is separated from
A. columbella new species by its greater number of ante-
rior ribs and a much finer crenulation of the internal
margin. In addition, the area adjacent to the poste-
rior sulcus is flat, not sunken as characteristic of
A. columbella. The sulcus here is less marked com-
pared to the very strong expression in A. columbella.
Americardia lightboumi is more oblique compared to
the rather square, upright shape ol A. columbella.
Americardia lightboumi is also markedly smaller and less
solid than that species. Their distribution is close in
Florida; otherwise A. lightboumi is much more widely
distributed. Both species live in the sublittoral although in
the northern part of its distribution A. lightboumi dwells
in deeper water, living in general below 50 m, whereas
A. columbella is most often found from 20 to 40 m.
From the Pliocene of Florida (e.g., APAC Pit, Sara-
sota) there is an apparently unnamed Americardia sp.
(Figure 24), which has a comparable number of about
29 ribs and similar outline and surface features. Com-
pared to Recent material, the valves of this fossil taxon
are more compressed. Additionally, it is less oblique, the
intercostal areas are broader, the beaks are lower and
more posterior (central), and the posterior angulation is
weaker. The senior author found this taxon in about ten
percent of the approximately 200 Plio-Pleistocene
Americardia lots housed at the UF collection. About
half of these (approximately 10) lots also contained
A. columba , and the two seemed to be morphologically
distinct without overlap. Specimens were present in
material from the Bermont, Caloosahatchee, Late
Duplin, and Tamiami formations. Interestingly, only
one lot referable to A. media was encountered [UF
35360 USA: Florida: Charlotte Co., Pnnta Gorda Quad,
Tamiami Formation, Pinecrest Beds R. Portell! 12/23/
88], This station produced several other unusual taxa
(R. Portell, pers. comm., 21 Dec., 2010). Nonetheless
this extinct Americardia sp., apparently coeval with
A. columba (Figure 23), may well represent the ancestor
of A. lightboumi.
Cardium speciosum A. Adams and Reeve, 1850 was
described b om China in the notorious Samarang Report
(see Huber, 2010: 75). As with nearly half of the bivalves
newly described therein, C. speciosum was never again
collected in Chinese waters. E.A. Smith (1890a: 302)
recognized its misallocation, compared it with Atlantic
specimens, and accepted it as a St. Helena species, likely
collected on the return “voyage” from China.
The type material is difficult to analyze. NHMUK
74.12.11.392 represents a specimen, undoubtedly from
St. Helena and labeled type. This “type” specimen is
accompanied by three specimens labeled “syntypes”. Thus,
the whole lot contains four specimens, which, however,
represent two distinct species. Originally, Adams and
Reeve did not give any size or indicate more than one
specimen. Furthermore, E.A. Smith (1890a: 302) only
referred to “the type of this species preserved in the British
Museum." Adams and Reeve clearly indicated 26 ribs.
None of the present four specimens has 26 ribs. All of the
three “syntypes" have far in excess of 30, up to 35 ribs.
Together with shape and color, these three “syntypes” can
be dismissed as subsequently introduced A. media, neither
originating from the original lot nor St. Helena. The “type”
itself has 28 ribs, two thereof admittedly rather weak.
Otherwise it matches the original description well with its
strongly oblique shape, moderately marked sulcus, solid
texture, presence of reddish marks, and commarginally
sculptured interstices. It is here recognized as the holotype
(Figure 20). Consequently, the other NHMUK “syntypes”
should be removed from the type collection.
In the general collection there is an unambiguous lot,
NHMUK 1889.10.1.1960-4, comprised of five well pre-
served specimens collected in the 19th Century by Lt. W.H.
Turton at St. Helena. These, together with recently
collected specimens, allowed us to finally characterize
A. speciosa as expressed herein. The exact depth of occur-
rence is still unknown, but it is presumed to be sublittoral.
Another lot, NHMUK 1889.10.9.38, collected by
Conry at Ascension Is., is of importance. These shells
confirm E.A. Smith’s (1890b: 322) identification: they
match A. media in having 25-26 anterior ribs, a solid
shell, a rather broad and weakly sulcate shape, and the
same color pattern. This record confirms the southern-
most occurrence of this species.
In many respects A. speciosa and A. lightboumi rep-
resent extremes of this small Atlantic Americardia-
group. A. lightboumi lias the highest rib count;
A. speciosa has the lowest; A. lightboumi is rather fragile,
whereas A. speciosa is solid. Their maximal sizes differ
markedly, and their distribution is disjunct.
Americardia columbella new species
(Figures 12-17, 22)
Americardia media “Linnaeus” Porter and Houser (1994: 19,
North Carolina) non Linnaeus, 1758.
Americardia columba “Heilprin” Lee (2009: 33, pi. 7, sp. 119,
NE. Florida) non Heilprin, 1886.
Americardia media “Linnaeus” Huber (2010: 300 fig. If, left
specimen, NE. Florida) non Linnaeus, 1758.
Americardia media “Linnaeus” Tunnell et al. (2010: 353, Texas)
non Linnaeus, 1758.
Diagnosis: Large, quadrangular, orange-brown Americardia
with distinct posterior sulcus and intermediate number of
ribs, closely adjacent and without any interstices anteriorly.
Page 20
THE NAUTILUS, Vol. 126, No. 1
Figures 12-17. Americardia columbella new species. 12-13. Holotype, NE Florida, 42.2 mm 12. External view L, R valve.
13. Internal view L, R valve. 14. Paratype suite 1, NE Florida, 44.5 mm. External view, R valve. 15-17. Paratype suite 1, NE Florida,
30.1 mm. 15. External view L. R valve. 16. Internal view L, R valve. 17. Posterior view, paired valves. Scale bar (all views) = 1 cm.
Description: Shell solid and thick, large for genus,
reaching 44.5 mm in NE Florida, inflated to strongly
inflated in old specimens; subquadrate, with a strong
posterior sulcus, which is strongest extended medially.
Anterior margin evenly rounded; ventral margin nearly
straight, weakly sinuate in posterior third. Posterior
angulation marked but moderately rounded. Shells
equivalve and subequilateral, with rounded, orthogyrous
umbones nearly centrally placed and clearly extending
above the dorsal line. Surface radially sculptured, with
23-25, usually with 24 anterior ribs. Anterior ribs
touching each other, eliminating any flat interspace as
found in all other Americardia. Ribs subtrigonal in
cross-section, with highest elevation in posterior third.
Ribs beneath intritacalx glossy, smooth, without spines
or elevated scales. Dehiscent intritacalx composed of
orange-brown calcareous layer, very densely and regu-
larly eommarginally lined, often forming small knobs
on highest part of ribs. External ligament veiy small,
light brownish, situated just posterior to umbones.
External coloration whitish with irregularly arranged
orange-brown blotches and streaks. Overall shell appears
brownish-orange. Internal shell surface in general
glossy-white, brown along posterior margin. Pallial line
entire, positioned close to ventral margin and connect-
ing two mediums-sized, nearly homomyarian, subovate
scars. Margin strongly crenulate. Crenulations broader
centrally and narrower on both sides. Hinge line strong
and rather thick with two cardinal teeth and two pro-
minent lateral teeth. Posterior cardinal in right valve
and anterior lateral in left valve strong. Valves close tightly.
Variations: Externally, rarely a weak purplish-red band
is present, in addition to the typical orange-brown
blotches; inside, occasionally purplish spots or streaks are
found; rarely the 3-4 ribs immediately anterior to the
carina may be separated by very narrow interstices; invari-
ably, however, the anterior portion never has interstices.
In general, soudiem and western Florida specimens are
smaller, lighter colored, and occur in deeper water.
Etymology: This new Americardia is named as dimin-
utive of columba (from the extinct A. columba.) The
Pliocene Florida fossil shares morphological features
with and may represent the ancestor of A. columbella .
Type Locality: Eastern USA, NE Florida, St. Johns
Co., Nine Mile Reef, 9 miles ESE St. Augustine, 22 m,
near reef, on sand; 1 September, 1979, H.G. Lee (dive).
Type Material: Holotype: UF 447280, one paired
specimen, 42.2 mm [Figures 12. 13], from the type local-
ity, ex Coll. H.G. Lee. Paratype suite 1: USNM 1 156949,
one single valve, 44.5 mm [Figure 14], and three paired
specimens, 9.9 to 30. 1 mm [Figures 15, 16, 17], from the
type locality, ex Coll. H.G. Lee. Paratype suite 2; Coll.
M. Huber, QQ232, two paired specimens, NE. Florida,
60 miles SSE St. Augustine, dredged in 40 - 43 m, June,
1978; larger specimen, 29.2 mm illustrated in Huber
(2010: 300 fig. 11, left specimen). Paratype suite 3: Coll.
H.G. Lee, five paired specimens, 15.5 to 38.5 mm, same
data as holotype. Paratype 4: Coll. H.G. Lee, one paired
specimen, 30.6 mm; NE. Florida, Mayport, dredged in
21 .5 m, November, 2009.
Distribution: This new species is narrowly restricted
geographically, only known from the southeast coast of
North America, from North Carolina to Yucatan. It is
H.G. Lee and M. Huber, 2012
Page 21
Figures 18-22. Western Atlantic Americardia species. 18. Cardium medium Linnaeus, 1758, 29 mm, Linnean Society Collection.
19. Cardium medium , neotype, Linnean Society' Collection 16920 [online] (Dance image G-M 0010260), 37 mm. 20. Cardium
speciosum A. Adams and Reeve, 1850, holotype, NHMUK 74.12.11.392, 28.5 mm. 21. Americardia lightbounii, holotype, 20.5 mm.
22. Americardia cohimbeJla . holotype, 42.2 mm. Scale bar (all views) = 1 cm.
rather uncommonly encountered and rarely, if ever,
cast ashore (personal observations; Andrews, 1971, 1992;
Ode, 1973, 1975; Rice and Komicker, 1962; Vokes and
Vokes, 1984; UF 328133 H.E. Vokes!).
Habitat: Americardia columbella lives sublittorally
offshore, with many records from 20-40 m, occasionally
down to 60 m; the substrates vary from shell grit to
coarse and fine siliceous sands, rarely on carbonate bot-
toms. It is ol relevance that, in all collections studied,
the type species, A media , is restricted to much
shallower habitats.
Discussion and Comparison: With the type species,
Americardia columbella shares a comparatively large
size, a solid shell, but also a broad hinge and an identi-
cal dentition. The rib count is comparable, although
A. media often has one or two more anterior ribs. The
carina in A. columbella is generally more prominent,
weaker, and more rounded in A. media. The shape is
more quadrangular upright, whereas A. media shape
is more oblique anteriorly. Americardia columbella is
most easily separated by its more prominent posterior
sulcus and its more vivid orange red color. Moreover,
the area adjacent to the sulcus is typically sunken
in columbella , whereas in A. media this area is flat.
Unmistakably distinct is the rib sculpture; A. media has
rather flat- topped ribs with rather broad interstices, but
the interstices are completely missing in A. columbella ,
and the ribs are subtrigonal in cross-section. The two
Figures 23-24. Americardia fossil species. 23. Americardia columba , Pliocene, Florida, external view L, R valve. 24. Americardia
sp.. Pliocene, Florida, external view L, R valve. Scale bar (all views) = 1 cm.
Page 22
TPIE NAUTILUS, Vol. 126, No. 1
differ in geographic distribution: A. media is mainly a
West Indian species, occasionally found in southern
Florida and southeastern Mexico, and A. columbella
is not known outside of eastern North America, in
the Carolinian portion of the Caribbean Province.
Americardia media lives just subtidally and is commonly
beached, whereas A. columbella is a sublittoral species,
rarely, it ever, cast ashore. The common A. media and
the rather uncommon A. columbella are often found in
sizes between 25-30 mm; however, the maximum size
of A. media is 61.3 mm (West Indies, NHMUK, while A.
columbella is not known to exceed 44.5 mm (paratype 1).
Hemicardium columba Heilprin (1886: 93) was
described as a Pliocene Floridian fossil from two left
valves, illustrated on his pi. 11 figs. 26 and 26a, up
to 20 mm. Heilprin compared it to the Recent
Hemicardium medium of southern Florida. He noted
23 anterior ribs and additionally 13 on the posterior
slope. Compared to H. medium , Heilprin noted that
H. columba “its more upright form, the deeper hollowing
of the posterior face, and the more pronounced
carination of the umbonal slope”. In this regard
A. columba neatly shares the features found in the
Recent A. columbella.
However, well preserved Pliocene fossils from Florida
revealed a character not mentioned by Heilprin, namely
a veiy distinct rib sculpture (Figure 23). In Americardia
columba , the comparatively broad ribs are low and
rather flat, separated by a deep flat channel. Thus the
rib structure of A. columba closely approaches that of
A. media while it is quite different from A. columbella
with its lack of interstices and higher, subtrigonal ribs.
Moreover, the Recent material is broader in shape and
clearly sinuate at the margin of the valves posterior
third. The original coloring of the extinct species is
unknown. Due to similarities in shell shape, it is hypoth-
esized that A. columbella is a comparatively young
species derived from an old Floridian stock represented
by A. columba.
Cardium speciosum is only known from St. Helena,
and A. columbella only from the southeast coast of North
America. If the locality is known, these two cannot
Table 1. Comparison of the four Recent Atlantic Americardia species.
H.G. Lee and M. Huber, 2012
Page 23
be confused. They possess a similar exterior color, reach
similar sizes, and both have solid shells. However,
typically their shape is markedly distinct: narrow and
oblique, with a moderately expressed sulcus in
A. speciosa and broad, subquadrangular, with a promi-
nent sulcus in A. columbella. A. speciosa has a signifi-
cantly lower rib count, with interstices of about hall
a rib’s breadth, while in A. columbella the number of
anterior ribs is greater, and interstices are lacking. Inter-
nally, A. speciosa is yellowish, even deep yellow in some
specimens; A. columbella is glossy white within.
CONCLUSIONS
The simple question, whether the “Recent" Floridian
A. columba is indeed distinct from “A. media'’ led to many
unexpected results. Identification ot material from various
locations proved straightforward, with one exception.
Smaller south Florida material was difficult. The cause of
this predicament appears to be threefold: firstly, three
species may be found there, secondly, cardiids globally
develop many of their taxonomic characteristics only at
medium to large size, and third, it could not be completely
excluded that some specimens represent hybrids. The
authors are convinced that instead of one, three species
of Americardia live in Florida waters. A fourth Atlantic
species evolved in St. Helena. Evidence suggests that
a second undescribed and now extinct Americardia lived
coevally with the well known Pliocene A. columba in
Florida waters. Based on the diversity encountered in
this study, it is suggested that Americardia originated
in, and radiated from, this area.
ACKNOWLEDGMENTS
Thanks are due to Kathie Way and Andreia Salvador,
Natural Histoiy Museum, London (NHMUK), for allow-
ing the junior author to study the NHMUK material of
Americardia species. Phil Hurst (NHMUK) produced
the photograph of the holotype of Cardium speciosum ,
and Ms. Way facilitated the delivery of the images taken
by S. Peter Dance of C. medium in the Linnean Society
Collection. Dr. Gustav Paulay, Roger Portell, and John
Slapeinsky of the Florida Museum of Natural Histoiy
kindly provided space, instrumentation, and specimen
material for the senior author. Dr. Emilio Garcia
(Lafayette, LA) and Sue Hobbs (Cape May, NJ) provided
valuable material, and Bill Frank (Jacksonville, FL) lent
technical assistance in assembling the plates.
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Biodiversity Project 2004 and the Panglao 2005 Deep-Sea
Cruise with descriptions ol four new species (Bivalvia).
Vita Malacologica 8: 9-96.
Tunnell, J.W. Jr., J. Andrews, N.C. Barrera, and F. Moretzsohn.
2010. Encyclopedia of Texas Seashells. Texas A&M Uni-
versity', College Station, xi + 512 pp.
Vidal, J. 2000. Classification of Cardiidae. Phuket Marine Bio-
logical Center Special Publication 21(3): 639-644.
Vokes, II. E. and E.H. Vokes. "1983” [1984], Distribution
of shallow-water marine mollusca, Yucatan Peninsula,
Mexico. Mesoameriean Ecological Institute, Monograph 1,
Middle American Research Institute, Publication 54:
i-viii + 1-183.
THE NAUTILUS 126(l):25-32, 2012
Page 25
New Naticidae (Gastropoda) from Brazil
Paulo Marcio Santos Costa
Museu Nacional, Universidade Federal do Rio de Janeiro
Departamento de Invertebrados
Setor de Malaeologia
Rio de Janeiro, CEP 29940-040, RRAZIL
[email protected]
Guido Pastorino
Museo Argentino de Ciencias Naturales
Av. Angel Gallardo 470, 3° piso, lab. SO
C1405DJR Buenos Aires, ARGENTINA
[email protected]
ABSTRACT
Two new species are described from Brazil. Notocochlis Jmirae
new species from the Canopus Bank, a seamount located 160 Km
off Fortaleza, Ceara, is characterized by its large size, die umbil-
ical and parietal calluses separated by a notch, a straight sulcus,
and a moderately developed funiele, brownish-white color, with
axial and oblique bands extending from the suture to the lower
third of the whorl, and the subsutural region widi a thin white
spiral band. Notocochlis isabelleana , a shallow water species, is
comparable and can be differentiated from N. laurae by its
small size, rounded shape, coloration pattern and operculum
with only one marginal rib. Notocochlis guesti , a deep-water
species that occurs in Bermuda and the Caribbean, can be
differentiated by its rounded profile, pale coloration pattern,
and radular characters. Natica juani new species, distributed
from southern Brazil to Uruguay, has a small, smooth shell, with
an obsolete funiele. Its color pattern has axial brownish zig-zag,
interconnected, flammules over a white background. The internal
margin of the operculum is serrate and the external margin has
two thick cords. It is similar to N. perlineata and N. menkeana,
both Caribbean species with different coloration pattern and a
thicker funiele in the latter.
Additional keywords: Natica, Notocochlis, Canopus Bank
Seamount, Notocochlis guesti, Natica menkeana, Natica castrensis
INTRODUCTION
About 17 species of Naticidae are known from the
Brazilian coast. Except for a few rare species (e.g., Euspira
radiata (Watson, 1881), found below 500 m) most natieid
species known from this region occur in shallow waters
(Rios, 1994). Nevertheless, a systematic revision of the
entire family from Brazilian waters is needed. One prob-
lem in studying this family is that some genera remain
inadequately defined. Many of the species are easy to
identify, but some of tire subfamilies and genera require
better phylogenetic resolution, as some are grades, not
clades. Traditionally, the genus Natica sensu lato has been
used as a catch-all name including the numerous species
belonging to the subfamily Naticinae. Pastorino (2005)
recently reviewed all the species of naticids living along
the Argentine coast, restricting the use of Natica, and using
the genus Notocochlis Powell, 1933 for several southwest-
ern Atlantic species hitherto placed in Natica.
In 2005, during a dredging expedition on the north-
eastern coast of Brazil, nrollusks were collected between
240 and 260 m depth from Canopus Bank, off Ceara,
including a new species ol Naticidae, which is described
in the present paper.
MATERIALS AND METHODS
Several shells and live specimens were dredged from
Canopus Bank, off Ceara, Brazil, by commercial fishing
boats; others are from the stomachs of the fish Ogcoccphalus
vespertilio (Linnaeus, 1758) and of tire starfish Astropecten
sp. Additionally, material from tire collections of the Museu
Nacional do Rio de Janeiro (MNRJ), Museu Oceanografico
Professor Eliezer de Carvalho Rios (MORG), and Museo
Nacional de Historia Natural y Antropologia, Montevideo
(MNHNM) (collected during several cruises of the B/I
Aldebaran and the R/V Academik Knipovich), were
also studied.
Radulae and jaws were prepared by as in Solem
(1972). Scanning Electron Microscope (SEM) photo-
graphs were taken with a Philips XL30 at the Museo
Argentino de Ciencias Naturales “Bernardino Rivadavia”
(MACN) and a |EOL JMS - 6390 LV at the Centro de
Microscopia Eletronica de Varredura at the Museu
N acional/UFRJ .
SYSTEMATICS
Superfamily Naticoidea Guilding, 1834
Family Naticidae Guilding, 1834
Subfamily Naticinae Guilding, 1834
Genus Notocochlis Powell, 1933
Type Species: Cochlis migratoria Powell, 1927, by
original designation (-N. gualteriana Reeluz, 1844).
Notocochlis laurae new species
(Figures 1-8, 15, 17-20)
Diagnosis: Shell ol large size for genus, reaching
about 40mm. Color brownish white, axial and oblique
bands, from suture to lower third ol whorl; white thin
Page 26
O
THE NAUTILUS, Vol. 126, No. 1
Figures 1-16. Natieid species. 1-8, 15. Notocochlis laurae new species. 1-7. Holotype MNRJ 11866, 33.2 x 31.0 mm. 5. External
view of the operculum. 6. Internal view of the operculum. 7. Detail of the umbilical area. Scales bars = 1 cm. 8. Paratype MNRJ
11867, SEM detail of the protoconch. Scale bar 500 pm. 15. Detail of the operculum. Scale bar = 2.4 mm. 9-1 1, 16. Natica gnesti
Harasewych and Jensen, 1984. 9-10. Two views of the holotype, USNM 765087. 11. External view of operculum. Scale bar = 1 cm.
12-14. Notocochlis isabelleana MNRJ 17873, 19.5 x 18.8 mm. 12-13. Views of shell. 14. External view of the operculum. 16. Natica
guesti Harasewych and Jensen, 1984, detail of the operculum showing the smooth inner margin. Scale bar = 2.4 mm.
P. M.S. Costa and G. Pastorino, 2012
Page 27
Figures 17-20. Notocochlis laurae new species. Holotype MNRJ L1S66. 17. Radulae. Scale bar = 100 pm. 18. Detail of raehidian
teeth. Seale bar = 50 pm. 19. Jaws. Seale bar = 500 pm. 20. Detail of the rods of the jaws. Scale bar=100 pm.
subsutural, spiral band always present. Operculum with
a smooth internal margin.
Description: Large shell (Figures 1-4, 7-8), holotype
33.2 mm in height (40.0 mm maximum height), globose,
spire moderately elevated; shell thickness average for
genus; protoconch (Figure 8) of about 3.0 whorls (1.4 mm
of diameter), smooth, without ornamentation, transition
to teleoconch defined. Suture distinctly impressed. Tele-
oconeh with up to three rounded whorls; axial sculpture
of incised furrows on first teleoconch whorls, turning into
oblique incremental growth lines in succeeding whorls;
suture adpressed. Parietal callus very thin, separated from
umbilical callus by notch; lobe of anterior parietal callus
large, weakly defined. Umbilicus (Figure 7) moderately
narrow, always open; umbilical callus moderate in size,
always present; sulcus deep, excavated, straight (channel
wide, overtaking anterior internal lip without notch);
funicle moderately developed; basal lip sharp. Aperture
veiy large, semicircular.
Color of fresh specimens brownish white, axial and
oblique bands present from suture to lower third of
whorl; white thin subsutural spiral band always present.
Periostracum brownish, very thin.
Operculum calcareous (Figures 5-6, 15), semicircu-
lar, solid, paucispiral, closing entire aperture. Internal
opercular sculpture of growth lines, covering whole sur-
face; two well defined marginal ribs of angular profile,
after them, a spiral groove along outer margin as wide as
both marginal ribs together; calcified granulose zone on
center of operculum; inner and outer margins smooth.
Radulae taenioglossate (2-1-R-I-2), raehidian teeth
trapezoidal with three sharp cusps, central cusp larger
than lateral cusps, anterior edge of base straight, poste-
rior edge slightly convex centrally and concave at tips,
ending in two sharp lateral processes. One lateral tooth
Page 28
THE NAUTILUS, Vol. 126, No. 1
Figures 21-36. Natica species. 21-28, 30. Natica juani new species. 21-27. Holotype MNRJ 17876, 9.0 x 9.0 mm. 25-26.
External and internal views of the operculum scale bar = 2.5 mm. 27. Detail of the umbilical area, scale bar = 5 mm. 28. Paratype
MNRJ 17877, 9.5 x 9.6 mm. 30. Natica juani new species, Paratype MNRJ 17875, detail of the protoconch SEM picture, scale bar =
500 pm. 31-33. Natica perlineata. Syntype USNM 87201, 18.0 x 18.2 mm. 34-36. Natica castrensis. Syntype USNM 87198. 12.2 x
12.8 mm. 29. Natica menkeana , original illustration (Philippi, 1851, pi. 15, fig. 8).
on each side, each with three cusps, central cusp four
times larger than almost-obsolete lateral cusps, inner
edge with two conspicuous basal prolongations. Two
marginal teeth curved towards rachidian tooth; inner
marginal tooth bifid with cusps of different sizes, inner
cusp smaller (Figures 17-18).
Jaws trapezoidal; rods elongated, virgule shaped, diag-
onally arranged (Figures 19-20).
Type Material: Holotype MNRJ 11866, collected
alive on 11/2005 by P. M. S. Costa and J. Coltro Jr.;
Paratypes, MNR) 11867, 12 shells and 3 opereula.
P. M.S. Costa and G. Pastorino, 2012
Page 29
Figures 37-40. Natica juani new species. Holotype MNRJ 17876. 37. Radula, frontal view. Scale bar = 20 pm. 38. Two details of
lateral teeth. Scale bar = 20 pm. 39. Jaws. Scale bar = 200 pm. 40. Detail of the rods of the jaws. Scale bar = 20 pm.
08/2005; MZSP 53715, 6 shells; MZSP 53926, 2 shells;
MZSP 55525, 1 specimen; MZSP 70293, 7 shells; MZSP
71794, 4 shells; all paratypes from the type locality.
Type Locality: Canopus Bank, off Fortaleza, Ceara,
Brazil, 02°14,25" S, 38°22'50" W, dredged between
240-260 m depth, from biogenic substratum, collected
by P.M.S. Costa and J. Coltro Jr.
Distribution: Known from the type locality only (see
map Figure 41).
Etymology: Dedicated to Laura Gomes Costa, daugh-
ter of the senior author.
Remarks: The new species has the diagnostic charac-
ters of the genus Notocochlis , an operculum with two
Page 30
THE NAUTILUS, Vol. 126, No. 1
Figure 41. General map showing most ot the localities
mentioned in text. The gray area indicates the distribution of
N.juani new species. On top, detail of Canopus Bank area, the
black arrow indicates the type locality of N. laurae new species.
marginal ribs and smooth margins, as well as the fused
umbilical and parietal calluses. Notocochlis laurae new
species is comparable to Notocochlis guesti (Harasewyeh
and Jensen, 1984) (Figures 9-11, 16), which occur from
Bermuda to Colombia in moderate deep water (live
201 m). The two species differ, however, in shell shape,
color pattern, and operculum sculpture. Individuals in
the new species have a higher spire, with the color pat-
tern consisting of an irregular brown band above milk
white background without flammules. Notocochlis guesti
shows one or two spiral bands of brown dots or flam-
mules. The operculum of N. laurae new species has
two marginal ribs of the same size with an angulated
profile, a groove as wide as both marginal ribs together,
and the inner margin not serrate. Notocochlis guesti has
the second marginal rib three times wider than the first,
forming a broad plateau and a groove slightly narrower
than the second rib. In addition, the inner margin is
strongly serrate.
The comparable South Atlantic species belonging to
the same genus is Notocochlis isabelleana (d'Orbigny,
1840) (Figures 12-14). In addition to the different color-
ation, N. laurae new species is almost twice as large as
N. isabelleana , which also has a quadrangular profile
with a prominent shoulder and higher spire, while the
new species is higher than wider, with a shallower suture
and thicker basal lip. The opercula in the two species
are also quite different, with the operculum in the new
species having two well defined marginal ribs, while
the operculum ol N. isabelleana has only one. Finally,
there is a significant geographical disjunction between
both species, since N. laurae new species is a deep
water species (~250 m) from off northeastern Brazil,
while N. isabelleana is a common shallow water species
(0—100 m) from the Argentine malacological province.
Genus Natica Scopoli, 1777
Type Species: Natica vitellus (Linnaeus, 1758) by
subsequent designation (Anton, 1838).
Natica juani new species
(Figures 21-28, 30, 37-40)
Type Species: Natica menkeana auct.: Costa, Fortes
and Freitas, 1997: 14; Costa, Rios and Calvo, 1997: 16
(non Philippi, 1851).
Diagnosis: Shell small, smooth. Umbilicus wide,
open; umbilical callus weak fused with parietal callus;
sulcus straight, well developed; funicle obsolete; basal
lip thick. Color of white background with axial brownish
zig-zag flammules, irregularly spaced and interconnected,
covering whole shell. Calcareous operculum with a serrate
internal margin, and two thick, sometimes one thinner,
cords on external margin.
Description: Shell (Figures 21-24, 27-28, 30) small,
thick-walled, up to 14 mm of maximum height, globose,
spire short, with about two and half whorls; body whorl
well developed; protoconch (Figure 30) of about 1.5 whorls
P. M.S. Costa and G. Pastorino, 2012
Page 31
(950 pm of diameter), smooth, without ornamentation,
transition to teleoconeh well defined; Suture distinctly
impressed. Axial sculpture of oblique incremental growth
lines. Parietal callus thin; lobe of anterior parietal callus
weakly defined but large. Umbilicus (Figure 27) wide,
always open; umbilical callus moderate in size, fused with
parietal callus, always present; sulcus deep, excavated,
straight; funicle moderately developed; basal lip sharp.
Aperture very large, semicircular.
Color (fresh specimens) white background with axial
brownish zig-zag flammules, irregularly spaced, intercon-
nected, forming a network, particularly on middle of last
whorl. Some specimens with thin, dark periostracum.
Operculum (Figures 25-26) semicircular, solid, com-
pletely calcified, paucispiral, closing entire aperture. Inter-
nal opercular sculpture of growth lines covering whole
surface; two well defined marginal ribs, sometimes one
extra, thinner, after them, a spiral groove along outer
margin; calcified granulose zone on center of operculum;
inner margin serrate, outer margin smooth.
Radulae taenioglossate (2-1-R-1-2), rachidian teeth
trapezoidal with three cusps, central one larger, almost
obsolete, lateral cusps smaller, anterior edge of base
straight, posterior edge slightly convex centrally and con-
cave at tips, ending in two weak lateral processes. One
lateral tooth on each side; central cusp of lateral teeth
twice as large as lateral cusps, inner edge of lateral teeth
with conspicuous basal prolongation toward center. Two
marginal teeth long, curved toward rachidian tooth;
inner marginal tooth bifid with cusps of different sizes,
inner cusp smaller (Figures 37-38).
Jaws pyriform; rods elongated, comma-shaped, diago-
nally arranged (Figures 39-40).
Type Locality: Southern littoral of Sao Paulo State,
Rrazil, in ~ 17—35 m.
Distribution: Rio de Janeiro, Brazil to Uruguay (Figure 41).
Type Material: Holotype, MNRJ 17876, southern lit-
toral of Sao Paulo State, Brazil, in ~17-35 m, from the
digestive tract of Astropecten sp., shrimp net; Paratypes:
MNRJ 15029, off Buzios, Rio de Janeiro State, Brazil,
22°42'39.4" S , 40°40'50.2" W, 3 shells collected on 03/
2007; MNRJ 15030, off Buzios, Rio de Janeiro State,
Brazil, 22°4T49" S, 40° 40' 29. 8" W, 3 shells collected on
03/2007; MNRJ 15028, off Ilha de Cabo Frio, Arraial
do Cabo, Rio de Janeiro State, Brazil, 23° 18' 00" S,
40o00'00" W, fishing vessel Muriae III coll., 2 specimens
collected on 04/1993; MNR| 17875, between Rasa Is.
and Ilha Grande, Rio de Janeiro State, Brazil, fishing
vessel Marques Torres coll., 2 shells; collected on 02/
1994, from the digestive tract of bat fish Ogcocephalus
vespertilio (Linnaeus, 1758); MNRJ 17877, from the
type locality, 1 shell; MACN-In 39255, from the type
locality, 10 specimens; MZSP 32274, off Ilha Grande,
Rio de Janeiro State, 1 specimen, dredged on 07/11/1969;
MZSP 31946, east of Ilha de Vitoria, Sao Paulo State,
36-40 m, 24 specimens, collected with shrimp net, on
06/1999; MZSP 32103, southeast of Lage de Santos, Sao
Paulo State, 55-62 m, 2 shells, collected with shrimp
net, on 10/1999; MZSP 32201, off Itajai, Santa Catarina
State, 26°37' W, 48° 15' S, 85 specimens, collected by otter
trawl. Carlo Magenta leg., 04/1997; MNHNM 15517
MARE, 35°38.2' S, 53°43' W to 35° 39. 4' S, 53°44.7' W,
61,5-63 m, 1 specimen collected on 14/11/05; MNIINM
15520, MARE, 35° 24.6' S, 53° 24' W to 35°26' S, 53°25.7' W,
60-61 m, 1 shell collected on 1/10/07; MNHNM 15521,
MARE, 35°58.T S, 53° 52. 3' W to 35°59.4' S, 53°53.5' W,
62-63,5 m, 2 shells collected on 3/10/07.
Etymology: Dedicated to Juan Pastorino, son of the
junior author.
Other Material Examined: MNHNM 10989, R/V
Academik Knipovich, St. 1067, 22-29/4/1967, Okean
Dredge, 34° 26' S, 5U48.7' W, 166 m, (3 shells) collected
by V. Searabino; MNHNM 15514, 12/11/05, MARE,
34°41.7' S, 52° 35. 6' W to 34°43.T S, 52°37.0' W, 66-68
m, (2 shells); MNHNM 15513, collected on 12/1 1/05,
MARE, 34° 46,5' S-52°23.7' W to 34°44.7' S, 52°23,5' W,
101-104,5 m, (1 shell); MNHNM 15515, 12/11/05, MARE,
34°55.8/ S, 52°39,5' W to 34° 57,5' S, 52°40.0' W, 80.5-
82.5 m, (1 shell); MNHNM 15519, MARE, 34° 58. 2' S,
52°30.9/ W to 34°59.4' S, 52°32.0' W, 108-106 m, l shell
collected on 12/11/05; MNHNM 11233 R/V Academik
Knipovich, St. 1061, 22-29/4/1967, Okean Dredge,
35°04.0' S, 52° 13.6' W, 175 m, (2 shells), collected by
V. Searabino; MNHNM 15516 13/11/05, MARE,
35° 22. 9' S, 53° 18.0' W to 35°24,5' S, 53° 19,3' W, 62-
63.5 m, (1 shell); MNHNM 15518, 14/11/05, MARE,
35° 38.2' S, 53° 43' W to 35°39.4' S, 53°44.7' W, 61,5-63 m,
(1 shell); all material, except noted, collected by F.
Searabino on board B/I Aldebaran.
Remarks: The complex coloration pattern, fused
umbilical and parietal calluses, operculum calcareous
with serrated internal margin, and up to three marginal
ribs, are the features of the new species that allow the
inclusion in the genus Natica. The new species is com-
parable to Natica menkeana Philippi, 1851, which is
common in Puerto Rico and rare elsewhere in the West
Indies. According to comparative material examined,
the specimen illustrated by Warmke and Abbott (1962:
96, pi. 17 fig. e), and the original illustration (Philippi, 1851b:
pi. 15, fig. 8, reproduced here in Figure 29), the colora-
tion pattern is the main difference. While N. menkeana
presents a network of reddish brown markings below the
suture and a spiral band below the middle of the body
whorl, the new species has a subsutural white field, and,
after it, a brownish network of zig-zag lines. In addition,
the spire is shorter and the funicle appears to be larger.
Natica castrensis Dali, 1889 appears to be a synonym of
N. menkeana (A. Kabat pers. comm., 2011). A syntype is
illustrated here in Figures 34-36. The subsutural color-
ation and a remnant of spiral band below the middle of
the body whorl are similar to the original illustration of
N. menkeana and the specimen illustrated by Warmke
and Abbott (1962).
THE NAUTILUS, Vol. 126, No. 1
Page 32
In that same publication, Dali (1889: 294) described
Natica perlineata as a possible variety of N. castrensis.
The type material is illustrated here in Figures 31-33.
The coloration pattern clearly indicates that N. perlineata
is not conspecific with N. menkeana or N. juani
new species.
ACKNOWLEDGMENTS
We want to thank to Jose Carlos Tarasconi, Porto
Alegre, Brazil, for the loan of the specimens of Natica
juani and the usual support to our research. A special
thanks to Antonio Gil Bezerra and Elisa Gradvohl
Bezerra, owners of IN ACE Shipyard (Industria Naval
do Ceara) for use of their fishing boat and to Jose and
M arcus Coltro by financial support to the field trips to
Canopus Bank in August and November 2005. Com-
ments by Renata Gomes greatly improved an earlier
draft of this paper. Fabrizio Scarabino (MNHNM) read
an early draft of this paper and found several specimens
in the old collection of the MNHNM that helped
extend the distribution of N. juani. Elivaldo de Lima
from “Centro de Miscroscopia Eletronica de Varre-
dura”, Museu Nacional/UFRJ (supported by CENPES/
PETROBRAS), helped with SEM micrographs of
protoconchs. The manuscript improved dramatically with
the revision of Alan Kabat and an anonymous reviewer. Ellen
Strong and Yolanda Villacampa (USNM) helped us with
photographs from Dali’s type material. Emilio Garcia
(Lafayette, LA) provided samples of rare specimens from
his own collection. Conselho National de Desenvolvi-
mento Cientifico e Tecnologico, Brazil (CNPq) provided a
PROTAX postdoctoral grant support No. 155263/2006-4
to P. M.S. Costa.
LITERATURE CITED
Anton, 11. E. 1838. Verzeiehnis der Conchylien. Halle, xvi + 110 pp.
Costa, P.M.S., E.C. Rios, and I S. Calvo. 1997. Moluscos
Eneontrados no Trato Digestivo do Peixe-Moreego,
Ogcocephalus vespei~tilio (Linnaeus, 1758) Coletados
na Costa do Rio de Janeiro. Siratus, Sao Paulo 3(13):
16-19.
Costa, P. M.S., R.R. Fortes, and C.A. Freitas. 1997. Lista de
Macromoluscos Coletados por Barcos de Pesca de
Arrastao ao Largo da Costa do Estado do Rio de Janeiro,
Sudeste do Brasil. Siratus, Sao Paulo 3(13): 14-15.
Dali, H. 1889. Reports on the results of dredgings, under the
supervision of Alexander Agassiz, in the Gull of Mexico
(1877-78) and in the Caribbean Sea (1879-80), by the
U. S. Coast Survey Steamer Blake’. Bulletin of the Museum
of Comparative Zoology 18: 1-492, pis. 10-40.
Pastorino, G. 2005. Recent Naticidae (Mollusca: Gastropoda)
from the Patagonian coast. The Veliger 47: 225-258.
Philippi, R.A. 1851a. Centuria quarta testaeeorum novorum.
Zeitsehrift fur Malakozoologie 8: 39-48.
Philippi, R.A. 1851b. Die Gattung Natica und Amaura. In:
Systematisches Conchylien-Cabinet von Martini und
Chemnitz, vol. 2, pis. A, 13-18.
Rios, E.C. 1994. Seashells of Brazil. 2nd. ed.. Fundayao
da Universidade do Rio Grande, Rio Grande. 368
pp.,113 pis.
THE NAUTILUS 126(l):33-37, 2012
Page 33
Eosipho zephyrus , a new species (Gastropoda: Buecinidae)
from deep water off Chile
Koen Fraussen
Leuvensestraat 25
B-3200 Aarschot, BELGIUM
[email protected]
Javier Sellanes
Universidad Catolica del Norte
Faeultad de Ciencias del Mar
Larrondo 1281, Coquimbo, CHILE
[email protected]
and
Centro de Investigation Oceanografica en
el Paclfico Sur-Oriental (COPAS)
Universidad de Concepcion
Casilla 160-C, Concepcion, CHILE
Peter Stahlsehmiclt
Institute for Environmental Sciences
U niversitiit Koblenz- Landau
Fortstrasse 7, D-76S29 Landau, GERMANY
[email protected]
ABSTRACT
Eosipho zephyrus, a new deep-water species from off the
Chilean coast is described. Protoconch morphology distinguishes
die new species from Cantharus aldermenensis Powell, 1971, a
species commonly placed in die genus Eosipho Thiele, 1929. It is
the first record of a member of the “ Eosipho aldermenensis
group” in the eastern Pacific. Generic allocation in Eosipho
Thiele, 1929 versus in Manaria Smith, 1906 is briefly dis-
cussed. The presence of a third cusp on the lateral teeth ol
the radula is observed, compared to the same phenomenon in
Manaria Smith, 1906 and Coins Roding, 1786, and considered
to be an atypical morphology within the group rather than a
feature warranting additional taxonomic separation.
Additional keywords : Manaria, radula, variability, biodiversity.
Eastern Pacific
INTRODUCTION
The coastal zone ol north to south-central Chile, strongly
influenced by wind-driven upwelling, is one of the areas
with the highest known primary production rates in the
world (Daneri et ah, 2000). Consequently, this area of
the southeastern Pacific Ocean harbors a vast pelagic
and benthic biomass. However, in spite of proof that the
benthic fauna is rich in endemic species, many species
are still unstudied, undescribed, or unknown. For a brief
history and overview of past and recent expeditions and
malacological investigations offshore the Chilean coast
we refer to Fraussen and Sellanes (2008: 97).
Fraussen and Sellanes (2008) treated some deep-water
species of the genus Aeneator Finlay, 1927 and a spe-
cies from the Concepcion Methane Seep area (or CMSA)
belonging to the genus Kn/ptos Jeffreys in Dautzenberg
and Fischer, 1896. The goal of this second paper is to
continue adding to the knowledge of the Buecinidae
from off Chile with the description of a new species and
interpreting it in a wider context by comparing with Indo-
West Pacific relatives. It belongs to what we call the
“ Eosipho aldermenensis group”. Species belonging to, or
assigned to, Eosipho are known from the western Atlantic
and from the Indo-West Pacific; tire Eosipho aldennenensis
group, however, was until now known only from the Indo-
West Pacific. The present new species is the first mem-
ber of the genus known to occur in the eastern Pacific.
Tropical deep-water Buecinidae are taxonomicaly a
rather puzzling group, not the least the species belonging
to Eosipho Thiele, 1929 and Manaria Smith, 1906. Both
genera are very similar and it is still a question whether
Cantharus aldermenensis Powell, 1971 and related spe-
cies, including the new? species described herein, belongs
to Eosipho or to Manaria. We follow the opinion of
Bouehet and Waren (1986: 466, 469), retaining this spe-
cies tentatively in the genus Eosipho. Strong evidence
exists that the Eosipho aldermenensis group may consist
of more than one species (Fraussen and Stahlschmidt,
unpublished), and the description of the present new
species is a first step towards answering this question.
ABBREVIATIONS
JS: collection of Javier Sellanes, Chile; KBIN: Koninklijk
Belgisch Instituut voor, Natuurwetenschappen, Brussels,
Belgium; KF: collection of Koen Fraussen, Belgium;
MNHN: Museum national d’llistoire naturelle, Paris,
France; MNPINCL: Museo Nacional de Historia Natu-
ral, Santiago, Chile; PS: collection of Peter Stahlschmidt,
Rohrbach, Germany; lv: live collected specimen; dd:
empty shell.
SYSTEMATICS
Class Gastropoda Cuvier, 1797
Order Neogastropoda Wenz, 1938
Subfamily Buecinoidea Rafinesque, 1815
Page 34
THE NAUTILUS, Vol. 126, No. I
Family Buccinidae Rafmesque, 1815
Genus Eosipho Thiele, 1929
Type Species: Chrysodomus (Sipho) smithi Schepman,
1911 (by original designation) (type locality: Indonesia,
north of Pulau Talisei, Celebes Sea, 01°58' N, 125°00' E,
1 165-1264 m, Siboga stn 122).
Remarks: Members of the genus Eosipho mainly live
in the Indo-West Pacific, with species known from the
Mozambique Channel in the west, along Indonesia to
the Philippines in the east, from Japan in the north to
New Zealand in the south. Eosipho smithi (Schepman,
1911) and Eosipho aldermenensis (Powell, 1971) are
both widespread species. The latter was tentatively
placed in this genus by Bouchet and Waren (1986: 469)
because of conchological affinities. In the present paper
we follow this opinion. Further study (Fraussen and
Stahlschmidt, unpublished) may involve the placement
of Eosipho aldermenensis in the genus Manaria Smith,
1906 (type species: Manaria thurstoni Smith, 1906, by
original designation, from “Gulf of Manar”, India). More
species with conchological characteristics similar to
Eosipho were included by Shikama (1977: 16, pygmaeus),
Bouchet and Waren (1986: 467-469, coriolis, engonia ,
thorybopus), Okutani and Iwahori (1992: 149-250,
tosaensis), Fraussen (2001: 1-5, poppet) and Fraussen
and Hadorn (2005: 107-109, atlanticus) as well as mem-
bers from hydrothermal vents (Okutani and Ohta,
1993: 217-218, desbruyeresi ; Waren and Bouchet, 2001:
191, auzendai).
Eosipho zephyrus new species
(Figures 1-10, 13-14, 17-21)
Description: Shell small for genus (up to 11.7 mm),
thin, rather fragile, snow white. Shape broad with short
spire, whorls convex, suture distinct. Teleoconch con-
sisting of 4 convex whorls with distinct suture. Protoconch
consisting of 1 1/2 glossy, rather convex whorls, tip small,
whorls rapidly increasing, last whorl rather inflated,
resulting in a flattened protoconch. First teleoconch
whorl with 6 broad spiral cords, interspaces of fine but
deep lines. Second whorl with 8, penultimate whorl with
10 or 11, broad, flat spiral cords; subsutural spiral cords
slightly narrower than abapieal ones. Body whorl with
22 spiral cords, 3 or 4 cords on siphonal canal; adapical
interspaces fine, abapieal interspaces gradually becom-
ing broader; interspaces on base half as broad as spiral
cords. Last half of body whorl occasionally smooth above
periphery (paratype 2, Figures 3, 4) or entirely smooth
(paratype 4, Figures 9, 10), but possibly with some traces
of interspaces visible near suture.
First teleoconch whorl with 22, second whorl with 27,
fine, slightly curved axial ribs, subsuturally rather weak,
abapically gradually becoming stronger; interspaces shal-
low, rather broad. Penultimate whorl with 18 (paratype 5)
to 28 (paratype 2), body whorl with 25-27 such axial
cords, gradually becoming weaker, last part of body
whorl almost smooth. Aperture oval, columella smooth,
glossy, slightly curved; outer lip thin, simple, edge sharp.
Siphonal canal short, broad, open.
Operculum (Figure 8) small, thin, transparent, yellow-
ish brown, elongate; nucleus terminal, tip sharp.
Radula (Figures 19-21) typically buccinid. Central
tooth tricuspid with rather rectangular base. Lateral teeth
variable, atypical of genus, principally bicuspid but with a
small additional intermediate cusp.
Type Material: Holotype (MNI4NCL 6677) (10.5 mm),
southern Chile, off Chiloe, about 42°S, in 500 m, del;
Paratype 1 (MNHNCL 6678) (11.7 mm), same locality
as holotype, eld; paratype 2 (KF 5442) (10.5 mm), same
locality as holotype, dd. Paratypes 3-4 (MNHNCL
6679-6680) (9.8-10.7 mm), Chile, off Concepcion, "Ant-
arctic Intermediate Water’ Cruise, R/Y Vidal Gormdz,
36024.I2/ S, 73°36.44// W, in 606 m, lv; paratype 5
(KF 5443) (9.7 mm), same locality as paratypes 3 and 4;
paratype 6 (MNI1N 24775) (11.8 mm), southern Chile,
off Taitao peninsula, INSPIRE Cruise, R/Y Melville,
46 54 15' S, 75°35.99/ W, in 497 m, lv; paratype 7 (PS
150133) (11.0 mm), same locality as paratype 6, lv;
paratypes 8-10 (KF 6579-6581) (10.7-12.2 mm), same
locality as paratype 6, 2 lv, 1 dd; paratype 11 (KBIN)
(11.5 mm), same locality as paratype 6, lv; paratypes
12-28 (JS) (8.8-13.2 mm), same locality as paratype 6,
16 lv, 1 dd.
Type Locality: Southern Chile, off Chiloe, around
42° S, upper continental slope, in 500 m.
Range and Habitat: Only known from the type mate-
rial. Detailed habitat data are not available, but two of
the sites (off Concepcion and Taitao peninsula), in which
the specimens were collected, have been identified as
methane seep areas. Live collected shells are covered
with a thick, unidentified sponge-like mass (Figures 9,
17-18).
Etymology: Eosipho zephyrus new species is named
after the Greek god for the west wind Zephyrus, a name
used as a noun in apposition. The name is an allusion to
the easternmost occurrence of this species, far from the
ranges of the other congeneric species in the Indo-West
Pacific, as if it were blown to the east by the wind.
Comparative Remarks: Eosipho zephyrus new spe-
cies is characterized by the broad shape with moderately
short spire, the convex teleoconch whorls, the weak spi-
ral sculpture consisting of flat cords and narrow inter-
spaces, the broad and blunt protoconch with rapidly
increasing whorl size, greenish and rather smooth
periostracum and small adult size.
Eosipho zephyrus new species is variable in presence
of spiral sculpture (present or absent on the adapical part
of the body whorl) and in number of axial ribs (18 to
28 ribs on penultimate whorl).
K. Fraussen et al., 2012
Page 35
Figures 1-12. Eosipho species. 1-10. Eosipho zephi/ms new species. 1-2. Holotype, 10.5 mm, Chile, Chilean upper continental
slope, in 500 m, MN1INCL 6677. 3—4. Paratype 2, 10.5 mm, same locality, KF 5442. 5-7. Paratype 5, 9.7 mm, Chile, oil Concepcion,
AIW Cruise, 36° 24. 12' S, 73°36.44' W, in 606 m, KF 5443. 8. Operculum of paratype 5, 2.2 mm, KF 5443. 9. Paratype 4, 10.7 mm,
same locality, MNPINCL 66S0. 10. Same shell, cleaned. 11-12. Eosipho aldermenensis (Powell, 1971), 16.8 mm, East China Sea, off
China, in 280-380 m, KF 5244.
The protoconch (Figure 13, 14) is quite atypical of
both Eosipho Schepman, 1911 and Manaria Smith,
1906. While species belonging to these genera have a
protoconch consisting of gradually larger, convex whorls
(Figure 15, 16), the new species differs in having a big-
ger last whorl and a flattened appearance.
The radula of Eosipho zephyrus new species is atypi-
cal of Cominellinae (containing both Eosipho Schepman,
1911 and Manaria Smith, 1906) by the presence of a
third cusp on the lateral teeth (Figures 19-21). This
additional cusp is usually small to minute and appears to
be an additional rather than a constant presence. Vari-
ability in number of rachidian cusps within a single genus
is not unusual in Bueeinidae: Neptunea lidding, 1798 with
3-7 lateral cusps (Golikov 1964: 17, 29-33; Fraussen and
Terryn, 2007: 21, 39, 64), Buccinum Linnaeus, 1758 with
2-6 lateral cusps (Golikov 1980: 54, 99-104), Prosipho
Thiele, 1912 with 2-6 lateral cusps (Powell, 1951: 193;
Oliver and Picken, 1984: 96; Dell, 1990: 187). The pres-
ence of an additional median cusp in Cominellinae is
known in Manaria lirata Kuroda and Habe in Habe,
1961 (Fraussen and Stahlschmidt, unpublished) and in
Buccinulinae in Drepanodontus Harasewych and Kantor,
2004 (Harasewych and Kantor, 2004: 7, 12, figs. 36-40).
Eosipho aldermenensis (Powell, 1971) (type locality:
“E. of the Aldermen Islands, New Zealand, 366-475 m”)
Page 36
THE NAUTILUS, Vol. 126, No. 1
Figures 13-21. Eosipho species. 13-14. Eosipho zephyrus new species, protoconch of holotype, scale bar = 300 micrometer,
MNHNCL 5866. 15-16. Eosipho aldennenensis (Powell, 1971). Vanuatu (BOA O expedition, station CP2319 ), MNIIN. Scale bar =
300 pm. 17-21. Eosipho zephyrus new species. 17-18. Paratype 3, 9.8 mm, Chile, off Concepcion, RIW Cruise, 36°24.12' S,
73°36'44 W, in 606 m, MNIINCL 6679. 19-21. Radula of holotype. Scale bar = 40 pm.
(Figures 1 1-12, 15-16), a widespread species in the Indo-
West pacific, differs in having a smaller protoconch com-
pared to shell length, broader spiral cords with broader
interspaces, a lower number of axial ribs and a larger
adult size. Some evidence may exist (Fraussen and
Stahlschmidt, unpublished) that several Indo-West Pacific
records assigned to Eosipho aldennenensis belong to dis-
tinct species. A thorough study of this complex is beyond
the scope of the present paper, and we hereby tentatively
regard “ aldennenensis ” as a group and we use this name
for the Indo-West Pacific species.
Manaria brevicaudata (Sehepman, 1911) (type
locality: Borneo, north off Kagayan de Sulu Island,
Flores Sea, SIBOGA stn. 45, 07°24' S, 118°15' E,
794 m) is similar in spiral sculpture (fine spiral
grooves), in variability (absence or presence of spiral
sculpture on the adapical part of the body whorl) and
periostracum (greenish and smooth) but differs in
having a slender shape with a high spire, a slightly
longer siphonal canal, a lower number of axial ribs
and a larger adult size. The protoconch of Manaria
brevicaudata is unknown, all specimens studied by us
have eroded protoconch.
ACKNOWLEDGMENTS
We are thankful to Philippe Bouehet and Virginie Heros
(Museum national d’Histoire naturelle, Pans, France)
K. Fraussen et al., 2012
Page 37
for the loan of Eosipho aldermenensis material, and to
David Monseeonr (Belgium) for reading and correcting
the English text. We also thank the captain and crew
of R/V Vidal Gormaz of the Chilean Navy and R/V
Melville, Scripps Institution of Oceonography. This
work was partially funded by FONDECYT projects No.
1 100166 and No. 1061217 to J.S. and COPAS center of
the University of Concepcion. FONDECYT project
No. 1061214 to Praxedes Munoz, NOAA Ocean Explo-
ration Program (via SCRIPPS Institution of Oceano-
graphy, contract NOAA NA17RJ1231) and the Office of
Naval Research of the US Navy and the Census of
Marine Life, through its field project COMARGE,
provided extra funding for logistics and ship time.
LITERATURE CITED
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nomieal notes on tropical deep water Buccinidae with
descriptions of new taxa. Resultats des Campagnes
MUSORSTOM. I & II. Philippines, tome 2. Memoires
du Museum national d'Histoire naturelle, 1985, serie A,
Zoologie 133: 457-499.
Daneri, G., V. Dellarossa, R. Quiiiones, B. Jacob, P. Montero,
and O. Ulloa. 2000. Primary production and community
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Dell, R.K. 1990. Antarctic Mollusca, with special reference to
the fauna of the Ross Sea. The Royal Society of New
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Fraussen, K. 2001. A new Eosipho (Gastropoda: Buccinidae)
from the Philippine Islands. Gloria Maris 39 (5-6): 90-97.
Fraussen, K. and R. Hadorn. 2005. A new species of Eosipho
(Gastropoda: Buccinidae) from Guadeloupe, Western
Atlantic. Novapex 6: 107-109.
Fraussen, K. and J. Sellanes. 2008. Three New Buccinid Spe-
cies (Gastropoda: Neogastropoda) from Chilean Deep-
Water, Including One from a Menthane Seep. The Veliger
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Fraussen, K. and Y. Terryn. 2007. The Family Buccinidae:
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A Conchologieal Iconography. ConchBooks, Hackenheim,
166 pp., 154 pi.
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Neptunea Bolten. Fauna of the USSR, Mollusks. USSR
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THE NAUTILUS 126(l):38-40, 2012
Page 38
Land slugs (Gastropoda: Pulmonata) on birds demonstrate
dispersal potential
Timothy A. Pearce
Carnegie M useum of Natural History
Section of Mollusks
4400 Forbes Ave.
Pittsburgh, PA 15213 USA
Robert S. Mulvihill1
Powdermill Nature Reserve
Carnegie Museum of Natural History
1847 Route 381,
Rector, PA 15677 USA
Katherine A. Porter
Carnegie Museum of Natural History
Section of Mollusks
4400 Forbes Ave.
Pittsburgh, PA 15213 USA
ABSTRACT
We describe the discovery of two slugs, Arion subfuscus and
Deroceras reticulatum, on living adult birds captured for
banding at Powdermill Nature Reserve, Rector, Pennsylvania.
The presence of slugs on birds suggests that bird transport
might play a role in slug dispersal.
Additional keywords: Gray fieldslug, Deroceras reticulatum ,
dusk-)' arion slug, Arion subfuscus, common yellowthroat,
Geothlypis trichas, northern saw-whet owl, Aegolius acadicus.
INTRODUCTION
Although land snails and slugs are quiotessentially slow-
moving creatures, their ability to colonize new areas
indicates that they do, in fact, disperse. Some terrestrial
gastropod species have evidently dispersed considerable
distances, including across inhospitable bodies of salt
water (Rees, 1965; Vagvolgyi, 1975; Wesselingh et ah,
1999). Other than direct locomotion or transport by
humans (including as eggs), possible agents of dispersal
include water (Smith and Djajasasmita, 1988), wind
(Kirchner et ah, 1997), and other organisms such as birds
(Dundee et ah, 1967; Purchon, 1977: 383; Wesselingh
et ah, 1999; Wada et ah, 2011).
Direct evidence for these dispersal mechanisms
remains limited, and we are not aware of any published
evidence for slug dispersal via wind or water. Most evi-
dence for gastropod dispersal via other organisms per-
tains to shelled snails. We are aware of only two previous
reports of slugs being carried by non-human animals:
one report of a slug in the fur of a house cat (McMillan,
1989) and one report of slugs being caught and subse-
quently dropped by bats (folmston, 1992). This paper
presents the first known report of slugs on living
adult birds.
1 Current address: National Aviary, Allegheny Commons West,
700 Arch Street, Pittsburgh, PA 15212 USA
MATERIALS AND METHODS
In 2003 and 2004, two land slugs were found on birds
trapped in mist nets at Powdermill Nature Reserve
(Rector, PA; 40.1636° N, 79.2668° W). The birds were
trapped by R.M. as part of the Powdermill Nature
Reserve (PNR) bird banding program. The nets were
located in a wet shrub-scrub habitat dominated by
willows. Cursory examination of all birds for ectopara-
sites is a routine part of the banding procedure, as is
documentation of molt, and, during examination, the
slugs were discovered on two species of birds. The two
birds upon which the slugs were found were a common
yellowthroat ( Geothhjpis trichas (Linnaeus, 1766)) and a
northern saw-whet owl ( Aegolius acadicus (Gmelin,
1789)). The common yellowthroat was captured on Sep-
tember 14, 2003, and the northern saw-whet owl was
captured on November 11, 2004 at night. The slugs were
sent for identification and deposited in the collection at
the Section of Mollusks, Carnegie Museum of Natural
History (CM).
RESULTS
The two birds upon which the slugs were lound were a
common yellowthroat and a northern saw-whet owl.
The slug on the common yellowthroat was Deroceras
reticulatum (Muller, 1774) (CM 66603) and was lound
among the bird’s body leathers. This slug was a juve-
nile, 7.4 mm long (in alcohol), and was identified as
D. reticulatum by its paler tan color, reticulated mark-
ings, and presence of a pale pneumostome border
(mucus color was not noted before preservation). The slug
on the northern saw-whet owl was Arion subfuscus
(Drapamaud, 1805) (CM 103313), and it also was found
among the bird’s body feathers. This slug was also a
juvenile, 10.4 mm long (in alcohol), and was identified
as A. subfuscus by the pale sole and presence of 15 rows
of tubercles between the lateral bands (mucus color was
not noted before preservation). The common yellow-
throat was high in the net when it was captured; the owl
T.A. Pearce et al., 2012
Page 39
was in the lower shelf of the net and in contact with
the ground.
DISCUSSION
The presence ol the slugs within the plumage of the
birds suggests that these migrants might have been
actively transporting the slugs when they were caught.
We acknowledge the possibility that the slugs crawled
onto the birds after they were trapped in the mist net;
however, two factors suggest that this is unlikely. First,
during R.M.s 25+ years of mist-net banding at PNR, he
has never observed a slug or snail on a mist net. Second,
one ot the birds, the common yellowthroat, was caught
high in the net, about 1.5 m off the ground.
While both slug species were already known in the
vicinity of PNR, the possibility of transport by migratory
birds has implications for distribution and gene flow of
slugs, as it has for other gastropods (Miura et al. 2011).
The fact that this is the first report of slugs being found
on living adult birds suggests that if dispersal by birds
does occur, it is rare (or rarely documented). Kew (1893:
175), in a chapter on dispersal of slugs, speculated that
slug dispersal by birds, if it occurred, would be rare, but
did not cite any actual examples of such dispersal. How-
ever, given the wide ecological and migratory ranges of
many bird species, even very occasional transport by
birds could allow some slugs to spread over much greater
distances than would otherwise be possible.
The ranges and habitats overlap for the slugs and the
birds. In addition, both slug species ascend trees or veg-
etation during moist weather (Jennings and Barkham,
1975; pers. obs.), which might increase chances of an
encounter with a bird. Deroceras reticulation, originally
native to Europe, is now introduced in most temper-
ate and subtropical regions throughout the world (Roth
and Sadeghian, 2003; Forsyth, 2004). It occurs in open
and modified environments including agricultural land,
roadsides, hedges, and grasslands, avoiding woodlands
(Kerney and Cameron, 1979; Wiktor, 2000; Barker, 2002:
422; Forsyth, 2004). The common yellowthroat extends
throughout North and Central America and it inhabits
thick, tangled vegetation (particularly in wet areas) (Guzy
and Ritchison, 1999; GBIF, 2011).
The ranges of Anon subfuscus and the northern
saw-whet owl also overlap (Rasmussen et ah, 2008;
NatureServe, 2011), as do their habitats. Arion
subfuscus, native to Europe where it mainly inhabits
deciduous and coniferous forests and sometimes pas-
tures (Wiktor, 1983), is introduced in northeast and
northwest North America. Although it is not com-
monly associated with humans in most of Europe,
in western Britain and in North America, in addition
to occurring in deciduous and coniferous forests, it
is frequently found in gardens and on roadsides
near human settlements (Chichester and Getz, 1973;
Kerney and Cameron, 1979; Kerney, 1999; Forsyth,
2004). Saw-whet owls are found in a variety of wood-
lands and forested habitats, with densities highest in
coniferous forests, especially along riparian corridors
(Johnson and Anderson, 2003; Rasmussen et ah, 2008).
Because the geographical ranges and habitats of the
slugs and birds overlap, the current distribution of these
slugs in North America might be partially the result of
bird dispersal. Dispersal of the slugs by birds might be
especially relevant to their distribution given that they
are not native to North America and have been present
on the continent for relatively short periods of time
(160-170 years (Pilsbry, 1948)).
Two other examples of interactions between slugs and
birds, while not involving dispersal, indicate that contact
occurs between birds and slugs, demonstrating the plau-
sibility of slugs attaching to feathers of living birds.
Livezey et al. (2008) reported a barred owl capturing
and eating slugs. Biasiolli (2009) suggested that an
arionid slug, possibly Arion subfuscus, was eating a living
nestling black-throated blue warbler while the female
bird continued to attend to the nest. During these inter-
actions, other slugs, had they been present, might have
become entangled in the feathers of the birds.
The two slugs reported here were alive when they
were discovered. However, it is unclear how long the
slugs had been on the birds prior to their discovery.
The increased humidity inside a bird’s feathers, coupled
with the ability of many slugs to survive significant
desiccation (e.g., up to 50% of their body weight
(Runham and Hunter, 1970: 72)), might allow sings to
survive on birds far longer than otherwise expected. On
the other hand, birds might be expected to remove
slugs through preening.
Relatively little is known about the distribution and
transport of gastropods (especially slugs) by other ani-
mals. It is likely that transport by birds is infrequent,
given the lack of any prior observations of slugs on living
adult birds. For example, R.M. lias banded hundreds of
thousands of birds (including >2,500 common yellow-
throats and >100 northern saw-whet owls). Each bird is
examined for ectoparasites, as well as signs ol active
molt, but these are the first times slugs have been
noticed. The presence of small slugs deep within the
plumage of other birds handled at Powdermill, however,
cannot be ruled out, because the examinations may not
be detailed enough (especially on busy banding days) to
detect slugs deep in the feathers or otherwise obscured
(indeed, the soft, feather-covered blobs removed from
the birds were not initially recognized as slugs until they
started to move). It seems reasonable that species of bird
such as the common yellowthroat and northern saw-
whet owl, which forage (or attack prey) in dense, often
damp vegetation on and near the ground, might come
into direct contact with slugs that then become attached
to the birds plumage. Especially in the case of birds that
are actively migrating, it is conceivable that the urge to
migrate when conditions are favorable might supersede
maintenance behavior activities, such as preening. We
know that at least some of the northern saw-whet owls
captured at Powdermill at night were, in fact, actively
Page 40
THE NAUTILUS, Vol. 126, No. 1
migrating when they were netted (Powdermill Nature
Reserve, 2007).
Range expansion via bird transport has been reported
for snails (YVesselingh et ah, 1999). Our observations of
slugs on living adult birds suggest that this dispersal
mechanism might also be important to the establishment
and expansion ol at least some slug populations, which,
like many snails, are hermaphroditic and many can self-
fertilize. Further investigation into the prevalence of
slug transport by birds could be useful in predicting the
possible spread ol invasive slug species.
ACKNOWLEDGMENTS
We thank Bob Leberman at Powdermill for his role in
establishing and overseeing the bird-banding program for
more than 40 years, beginning in 1961, and also Marilyn
Niedermeier at CM for her assistance in accessing
Powdermilfs computerized banding data. Comments
from two anonymous reviewers improved the paper.
LITERATURE CITED
Chichester, L. F. and L. L. Getz. 1973. The terrestrial slugs of
northeastern North America. Sterkiana (51): 11-42.
Biasiolli, T. G. 2009. Depredation of black-throated blue war-
bler nestlings by an introduced slug (Arionidae). Wilson
Journal of Ornithology 121: 422-423.
Barker, G.M. 2002. Molluscs as Crop Pests. CABI Publishing,
New York. i-xii+ 1—468 pp.
Dundee. D.S., P. H. Phillips, and J. D. Newsom. 1967. Snails
on migratory birds. The Nautilus 80: 89-91.
Forsyth, R.G. 2004. Land Snails of British Colombia. Royal BC
Museum, Victoria. 188 pp.
GBIF (Global Biodiversity Information Facility). 2011.
Geothlijpis trichas (Linnaeus, 1766). http://data.gbif.org/.
[accessed 13 November 2011].
Guzy, M.J. and G. Ritchison. 1999. Common Yellowdiroat
( Geothlijpis trichas). The Birds of North America Online
(A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology, http://
bna.birds.comell.edu/bna/species/448 [accessed 13 Novem-
ber 2011],
Jennings, T. J. and J.P. Barkham. 1975. Food of slugs in mixed
deciduous woodlands. Oikos 26: 211-221.
Johnson, A. S. and S.FI. Anderson. 2003. Conservation Assess-
ment for the Northern Saw-whet Owl in the Black Hills
National Forest, South Dakota and Wyoming. United
States Department of Agriculture, Black Hills National
Forest, Custer, South Dakota. 27p. http://www.fs.usda.gov/
Internet/FS E_DOC U M E NTS/fsm9_0 12447.pdf [accessed
13 November 2011],
Johnston, D. 1992. Indiscriminate prey-capture in an oppor-
tunistic gleaner, the pallid bat Antrozous pallidus. Bat
Research News 33: 60-61.
Kerney, M.P. and R.A. D. Cameron. 1979. A field guide to the
land snails of Britain and North-west Europe. Collins,
London, 288 pp.
Kerney, M. 1999. Atlas of the Land and Freshwater Molluscs
of Britain and Ireland. Harley Books, Colchester. 261 pp.
Kew, H.W. 1893. The dispersal ol shells: an inquiry into the
means of dispersal possessed by fresh-water and land
Mollusca. Kegan Paul, Trench, Triibner and Co., Ltd.,
London, xiv + 291 pp.
Kirchner, Ch., R. Kratzner, and F.W. Welter-Sclmltes. 1997.
Flying snails - how far can Truncatellina (Pulmonata:
Vertiginidae) be blown over the sea? Journal of Molluscan
Studies 63: 479-487.
Livezey, K.B., M.F. Elderkin, PA. Cott, J. Hobbs, and J.P.
Hudson. 2008. Barred owls eating worms and slugs: the
advantage in not being picky eaters. Northwestern Natu-
ralist 89: 185-190.
McMillan, N.F 1989. Cat carrying a slug. Conchologists News-
letter (111): 245-246.
Miura, O., M.E. Torchin, E. Bermingham, D.K. Jacobs, and
R.F. Hechinger. 2011. Flying shells: historical dispersal of
marine snails across Central America. Proceedings of the
Royal Society B, published online 14 September 2011,
doi: 10. 1098/rspb.201 1 . 1599.
NatureServe. 2011. Avion subfuscus. http://www.natureserve.org/
explorer, [accessed 13 November 2011],
Pilsbry, H.A. 1948. Land Mollusca of North America (North of
Mexico). Academy of Natural Sciences of Philadelphia,
Monograph 3, volume II, part 2. i-xlvii, 521-1113.
Powdermill Nature Reserve. 2007. Pictorial highlights, late
fall 2007. http://www.powdermillarc.org/higlilights/2007/
november.aspx [accessed 13 November 2011],
Purchon, R. D. 1977. The Biology of the Mollusca, 2nd edition.
Pergamon Press, Oxford, xxv + 560 pp.
Rasmussen, J.L., S.G. Sealy, and R.J. Cannings. 2008. Northern
saw-whet owl ( Aegolius acadicus ). The Birds of North
America Online (A. Poole, Ed.). Ithaca: Cornell Lab
ol Ornithology, http://bna.birds.comell.edu/bna/species/042
[accessed 13 November 2011],
Rees, W.J. 1965. The aerial dispersal ol Mollusca. Presidential
address. Proceedings of the Malaeologieal Society of
London 36: 269-282.
Roth, B. and PS. Sadeghian. 2003. Checklist of the land snails
and slugs of California. Santa Barbara Museum of Natural
Histoiy Contributions in Science 3: 1—81.
Runham, N.W. and PJ. Hunter. 1970. Terrestrial slugs. Hutch-
inson University Library, London. 184 pp.
Smith, B.J. and M. Djajasasmita. 1988. The land molluscs
ol the Krakatau Islands, Indonesia. Philosophical Trans-
actions of the Royal Society of London, Series B 323:
379-400.
Vagvolgyi, | 1975. Body size, aerial dispersal, and origin of
the Pacific land snail fauna. Systematic Zoology 24:
465-488.
Wada, S., K. Kawakami, and S. Chiba. 2011. Snails can survive
passage through a bird’s digestive system. Journal of Bio-
geography. doi: 10. 1111/j. 1365-2699.2011. 02559.x, 5 pp.
Wesselingh, F.P, G.C. Cadee, and W. Renema. 1999. Flying
high: on the airborne dispersal of aquatic organisms as
illustrated by the distribution histories of the gastropod
genera Tnjonia and Planorbarius. Geologie en Mijnbouw
78: 165-174.
Wiktor, A. 1983. The slugs of Bulgaria (Arionidae, Milacidae,
Limacidae, Agriolimacidae - Gastropoda Stylommatophora).
Annales Zoologici 37: 71-206.
Wiktor, A. 2000. Agriolimacidae (Gastropoda: Pulmonata) - a
systematic monograph. Annales Zoologici 49: 347-590.
THE NAUTILUS 126(l):41-42, 2012
Page 41
The case of the abducted The Nautilus papers
Jose II. Leal
The Nautilus/The Bailey- Matthews
Shell Museum
P.O. Box 1580
Sanibel, FL 33957 USA
R.E. Petit Silvio Felipe Barbosa de Lima
P.O. Box 30 Departamento de Sistematica e Ecologia
North Myrtle Beach, SC 29582 USA Universidade Federal da Paraiba
Campus I. Cidade Universitaria
Joao Pessoa, Paraiba-PB, CEP. 58059-900 BRAZIL
The purpose of this note is to make known the existence
on the Internet, for over a year, of two papers on which
authorship had been changed from previously published
originals. Although now removed from the Internet,
these were downloadable .pdf files and may have been
downloaded and/or printed from those .pdf files.
In 2005, Petit was asked for assistance in identifying
some deep-water Cancellariidae by Lima, then a Brazilian
malacology student. As a result, two papers based on that
material were published in The Nautilus in 2007. One of
them was coauthored by Lima's professor, Jose Carlos
Nascimento de Barros (hereafter Barros) and Petit. On the
other paper, Lima was die first audior with Barros and Petit.
For the 2010 Semana Nacional de Ciencia e Tec-
nologia (2010 National Week of Science and Tech-
nology) a program with abstracts, but also including
complete papers, was placed onto the Internet. Articles
in .pdf format had the heading X Jornada de Ensino,
Pesquisa e Extensdo - JEPEX 2010 - UFRPE: Recife,
18 a 22 deoutubro (X Meeting of Teaching, Research
and Continuing Education).
The second of the two papers mentioned above,
authored by Lima, Barros, and Petit, was copied in .pdf
format in those online Jornada abstracts, with no men-
tion of The Nautilus or original authorship. Seven
authors are listed: Guilherme Lima Moraes, |ose Carlos
Nascimento de Barros, Jonata de Arruda Francisco,
Silvio Felipe Barbosa de Lima, Severino Adriano de
Oliveira Lima, Paulo Henrique Gomes da Paixao, and
Nefi Medeiros Fernandes. Institutional affiliation was
denoted by superscript numbers corresponding to
addresses at the bottom of the page. By the time this
appeared, Lima was no longer a student in Recife, but
at a different university in Paraiba. It must be mentioned
that Lima was unaware that this paper had been changed
and posted online until this was called to his attention
in July 2011. Barros is listed as second author and the
other five persons listed are believed to be his students,
as was Lima before he moved.
That The Nautilus paper is not copied in its entirety, but
what is copied is almost verbatim. Copied are the Intro-
duction, Systematics, and Discussion. The Description is
only about half copied. Type Material, Type Locality, Geo-
graphical Distribution, Etymology, Remarks, and Material
Examined are not included. However, all data concern-
ing the holotype is given in the figure caption. The figure
and its caption are copied without change. References
cited are deleted in the text, with numbers substituted
that correspond to numbered References at the end.
The first paper mentioned above, by Barros and Petit,
was copied in a similar manner. Figures 9-1 1 were not
reproduced as they are of a different species. Authors
shown on this copy were: Nefi de Medeiro Fernandes,
Guilherme Lima Moraes, Jonata de Arruda Francisco,
and Jose Carlos Nascimento de Barros.
When this matter came to our attention in June 2011,
Lima was contacted by Petit. He knew nothing of the
matter but promised to investigate although he was no
longer in Recife.
In early December 2011, it was noted that these
papers were still on the Internet. Once again Lima was
contacted. Petit asked Lima to advise Barros and the
people in charge of keeping these articles on the Inter-
net of possible consequences if they were not removed
immediately. As even that had no effect, the situation
was explained in rather stronger terms, which Lima
passed on and the papers were then taken off the Internet
sometime between 2 1 December and 27 December 201 1.
One of the problems with these copied papers is the
matter of them having wrong authorship. A more serious
problem is that any hard copies printed from these .pdf files
would appear to be Code-compliant original papers. /Ml
features needed for validation of the names were copied.
Although these two transformed papers are not pub-
lished under the provisions of the Code, there is nothing
in their appearance to indicate that they were not issued
in print. Such listings on the Internet can lead to the
misattribution of authorship and incorrect dating of nomina
when picked up by sources scavenging the Internet for
data used in compilation of lists and data bases. There
are now companies publishing entire “books” consisting
entirely of material uncritically taken from the Internet
(Petit, in preparation).
The PDF papers that are the subject of this article are
not listed in the Literature Cited as to do so would give
them an undeserved legitimacy.
Page 42
THE NAUTILUS, Vol. 126, No. 1
LITERATURE CITED
Barros, J. C. N. de and R. E. Petit. 2007. A new species of Micro-
cancilla (Gastropoda: Cancellariidae) from the continental
slope off northeastern Brazil. The Nautilus 121: 9.5-98.
Lima, S. F. B. de, }. C. N. de Barros and R. E. Petit. 2007.
A new species of Gerdiella (Gastropoda: Cancel-
lariidae) from the South Atlantic Ocean off Brazil with
discussion of an undescribed species. The Nautilus
121: 99-103.
Gofas, S., D. Moreno, and C. Salas (eds). 201 1 . Moluscos
Marinhos de Andalucia. 2 Volumes. Servicio de Pub-
licaciones e Intercambio Cientffico, Universidad de
Malaga. Xvi, pp 1-342 pp; xii, pp. 343-798. Hardcover.
Price €150 from publisher. €228/$296 at www.nhbs.com.
The marine malacofauna of the European- Mediterranan
area is possibly one of the best documented on the
globe. Comprehensive text-only treatments date back
at least to Weinkauff (1867, 1868). An early illustrated
work includes Bouquoy et al.’s (1882-1886) Les Mol-
lusques du Roussillion covering the famous Banyuls-sur-
Mer, France, field station. Subsequent comprehensive
works include Nordsieck (1968, 1969, 1972, 1982) with
its unmistakable line drawings, Poppe and Goto (1991,
1992), and the fairly recent Cossignani and Ardovini
(2011). There is no shortage on specialty volumes either,
including Ardovini and Cossignani (1999), the atlas series
(e.g. Giannuzzi-Savelli et ah, 1994), or the classic
Opisthobranchia tome by Schmeckel and Portman
(1982) with its stunning illustrations. Not to mention
the more limited treatments such as D’Angelo and
Garguillo (1978).
On the face of it, what could a regional guide to
Andalusia (Spanish coast from the southeastern border
to Portugal past Gibraltar including a quarter of the
Mediterranean coast of Spain) possibly add to this
already impressive library? Far more than the title
might suggest.
The overall work is a two volume set, 11 1/3 x
9 Vz inches = 28.3 x 23.7 cm, i.e., slightly taller than
US letter-size paper and significantly wider. The larger
Volume 1 contains a 50-page introduction to the marine
environment and its various biomes, the Andalusian
region, as well as an overview on moll usean system-
atics and diversity. Groups covered are Solenogastres,
Caudofoveata, Polypi acoph ora, and Gastropoda up to
Conidae. Volume 2 contains the remaining Gastropoda,
as well as Bivalvia, Scaphopoda, and Cephalopoda.
The introduction to each class contains 5-7 pages of
text including anatomical drawings and scanning elec-
tron micrographs (SEM) of the radula. The treatments
of the families by one, two, or three Spanish specialists
are moderately detailed. Each family is given a 1-2 page
introduction, including a section with key references.
The species treatments are in a two-column format with
illustrations along the outer margin and the text toward
the center. Generic diagnoses are not provided. Species
name and authority are given, but synonyms are not
indicated. About three species are treated on each page
covering the description of the shell, taxonomic remarks,
and indications on habitat and distribution. There are
several aspects that help render this volume a veritable
treasure trove, both for workers in the Mediterranean
and for a global audience.
First, the comprehensive treatment of the micro-
mollusks. As it has become more common in recent
publications, those are shown in detail. Even most
Scissurellidae and Anatomidae are correctly identified,
with the exception of Anatoma crispata (Fleming,
1 828) on page 30, which in fact shows an Anatoma
tenuisculpta (Seguenza, 1877); see Hoisaeter and Geiger
(2011) for details.
Another aspect to be emphasized are the multiple
illustrations for many taxa, particularly for micro-
mollusks. In some cases, multiple specimens are shown,
but in many cases complementary documentation is
also presented. For instance, the general shell mor-
phology of eulimids associated with a close-up image
showing microsculpture. Or photographs of pyra-
midellids with complementary SEM of the apex
with protoconch.
East but not least, the many photographs of live
animals are worth noting. While they are becoming
more common (e.g., Okutani, 2000; Redfern, 2001),
photos of live animals are particularly abundant in the
work under consideration, and include highly obscure
and rarely shown groups ( Casiella abylensis Gofas, 1987:
Cerithiidae; Cerithiopsidae, Triphoridae, Velutinidae,
seven (!) species of Runcina , Galeommatidae). Some draw-
ings of live animals complement the overall work (e.g.,
Scissurellidae, Tomidae, Cystiseidae).
The work is rounded out with a 12-page glossary of
terms, 25 pages of locality data for the specimens shown,
and 12 pages of taxonomic index; there is no compre-
hensive listing of literature cited.
The most relevant comparison of Gofas et al. is with
Cossignani and Adrovini (2011). Both are comprehen-
sive treatments of all classes, rich in high quality illus-
trations. The text in Gofas et al. is in Spanish only but
is much more comprehensive than the Italian/English
sections in Cossignani and Adrovini. Gofas et al.
includes all mollusks, without any omissions, while
Cossignani and Adrovini is comprehensive for the
shelled mollusks (e.g., Pendromidae), only presents a
selection of cephalopods and opisthobranchs, and does
not contain any of the worm mollusks. The illustrations
in both volumes are excellent; Cossignani and Adrovini
shines in showing about twice to three times as many
specimens per species than Gofas et al. The latter,
however excel with flawless and crisp printing, comple-
mentary light and electron microscopy images, as well
as abundant photographs of live animals.
Considering the purpose of an identification manual
(as opposed to a faunal monograph or a taxonomic revi-
sion), Gofas et al. reach that goal easily. Some decisions
with respect to the classification may surprise some
Page 44
THE NAUTILUS, Vol. 126, No. I
readers. There are no taxonomic categories between
class and family, and although the recent reclassi-
fication of the turrids is discussed, all Conoidea are
treated under a single family heading “Conidae y
afines” [Conidae and similar]. Given the long history
of taxonomy in the European-Mediterranaen region
with a plethora of taxa introduced, indications of
species-level synonyms would have been highly desir-
able. The main drawback of this volume is possibly the
Spanish language. For people who know any other
Romance language, it does not pose any significant
difficulties. Lucas Cervera Currado (pers. comm.)
informed me that an English version is under consid-
eration. Although the work is listed on tire publisher’s
website, it is not available through them (P. Valentich-
Scott, pers. comm.); it is however expected to be
listed shortly by the usual outlets (e.g., ConchBooks,
Mai de Mer).
The editors and authors of the Moluscos Marinos de
Andalucia can be congratulated for a job well done. For
people interested in the European malacofauna this is a
must-have item, and is worthy of serious consideration
for everybody else.
LITERATURE CITED
Ardovini, R. and T. Cossignani. 1999. Atlante delle Conchiglie
de Profundita de Mediterraneo. L’Informatore Piceno,
Ancona, 111 pp.
Bucquoy, E., P. Dautzenberg, and G. Dollfus. 1882-1886. Les
Mollusques Marins du Roussillon. j.-B. Bailliere & Fils,
Paris, 570 pp., 66 pis.
Cossignani, T. and R. Ardovini. 2011. Malaeologia
Mediterranea. L’Informatore Piceno, Ancona. 536 pp.
D'Angelo G. and S. Garguillo. 1978. Guida Alle Conchiglie
Mediterranee. Fabbri Editori, Milano, 224 pp.
Giannuzzi-Savelli, R., F. Pusateri, A. Palmeri, and C. Ebreo.
1994. Atlas of Mediterranean Sea Shells, Volume 1. La
Conehiglia, Rome, 125 pp.
Iloisaeter, T. and D. L. Geiger. 2011. Species of Anatoma
(Gastropoda: Anatomidae) in Norwegian and adjacent
waters, with the description of two new species. The
Nautilus 125: 89-112.
Nordsieek, F. 1968. Die europaischen Meeres-Gehausesehnecken
(Prosobranehia) vom Eismeer bis Kapveden und Mittelmeer.
Gustav Fischer Verlag, Stuttgart, 273 pp.
Nordsieek, F. 1969. Die europaischen Meeresmuseheln
(Bivalvia) vom Eismeer bis Kapveden, Mittelmeer und
Schwarzes Meer. Gustav Fischer Verlag, Stuttgart, 256 pp.
Nordsieek, F. 1972. Die europaischen Meeresselmeeken
(Opisthobranehia mit Pyramidellidae, Rissoacae). Gustav
Fischer Verlag, Stuttgart, 327 pp.
Nordsieek, F. 1982. Die europaischen Meeresgehausesehneeken,
2. Auflage. Gustav Fischer, Stuttgart, 539 pp.
Okutani, T. (ed.). 2000. Marine Mollusks in Japan. Tokai Uni-
versity Press, Tokyo, 1173 pp.
Poppe, G. T. and Y. Goto. 1991. European Seashells, Vol. I.
Christa Hemmen, Wiesbaden, 352 pp.
Poppe, G.T. and Y. Goto. 1993. European Seashells, Vol. II.
Christa Hemmen, Wiesbaden, 221 pp.
Redfern, C. 2001. Bahamian Seashells. Bahamianseashells
.com, Boca Raton, Florida, 280 pp.
Sehmekel, L. and A. Portman. 1982. Opisthobranehia des
Mittelmeeres. Nudibranchia und Saccoglossa. Springer,
Berlin, 410 pp.
Book Review, 2012
Page 45
Weinkauff, H.C. 1867. Die Conchylien des Mittelmeeres,
ihre geographische und geologische Verbreitung. Band I
Mollusea acephala. Theodor Fischer, Cassel, 301 pp.
Weinkauff, H.C. 1868. Die Conchylien des Mittelmeeres, ihre
geographische und geologische Verbreitung. Band II
Mollusea cephala. Theodor Fischer, Cassel, 512 pp.
Daniel L. Geiger
Santa Barbara Museum of Natural History
2559 Puesta del Sol Road
Santa Barbara, CA 93105 USA
[email protected]
THE 2012 R. T. ABBOTT VISITING CURATORSHIP
The Bailey-Matthews Shell Museum is pleased to invite applications for the 2012 R. T. Abbott Visiting Curatorship.
The Curatorship, established originally in accordance with the wishes of the late Dr. R. Tucker Abbott, Founding Director of the
Shell Museum, is awarded annually to enable malacologists to visit the museum for a period of one week. Abbott Fellows are
expected, by performing collection-based research, to assist with the curation of portions of the Museum’s collection and to provide
one talk for the general public. The Museum collection consists of marine, freshwater, and terrestrial specimens. A large percentage
of our holdings have been catalogued through a computerized database management system; part of the catalogue is already
available for searches online at: www.shellmuseum.org/collection.html. A substantial portion of the time will be available for
research in the collection, but field work in Southwest Florida can be arranged. The R. T. Abbott Visiting Curatorship is
accompanied by a stipend of $1,500.
Interested malacologists are invited to send a copy of their curriculum vitae, a letter detailing their areas of taxonomic expertise and
research objectives, and to provide a tentative subject for their talk. Send materials to:
Dr. Jose II. Leal, Di rector
The Bailey-Matthews Shell Museum
P.O. Box 1580
Sanibel, FL 33957
[email protected]
Applications for the 2012 Visiting Curatorship should be sent electronically to the above e-mail address no later than May 15, 2012,
or postmarked by that date if sent by regular mail. The award will be announced by mid-June 2012. Questions about the Visiting
Curatorship should be sent to the e-mail address above, or by phone at:
(239) 395-2233; fax (239) 395-6706
Sponsored in part by the State of
Florida, Department of State,
Division of Cultural Affairs and the
Florida Council on Arts and Culture
INSTRUCTIONS TO AUTHORS
THE NAUTILUS publishes articles on all aspects of the
biology, paleontology, and systematics of mollusks.
Manuscripts describing original, unpublished research
and review articles will be considered. Brief articles, not
exceeding 1000 words, will be published as notes and do
not require an abstract. Notices of interest to the mala-
cological community will appear in a notices section.
Manuscripts: Each original manuscript and accompa-
nying illustrations should be submitted to the editor pref-
erably via e-mail or as hardcopy in triplicate.
Text must conform to the dimensions of 8V2 x 11 -inch
paper, double-spaced, and single-column throughout (in-
cluding literature cited, tables, and figure captions). Au-
thors should follow the general recommendations of Sci-
entific Style and Fo rmat — -The CSE Manual for Authors,
Editors, and Publishers, available from the Council of
Science Editors at www.councilscienceeditors.org. The
first mention of a scientific name in the text should be
accompanied by the taxonomic authority, including year.
Latinized names and other words to be printed in italics
must be underlined; leave other formatting indications to
the editor. Metric, not English, units are to be used. Geo-
chronologic modifiers should be capitalized only when
units are formally recognized: for instance, use Late Cre-
taceous but early Miocene. Likewise, only modifiers of
formally recognized chronostratigraphic units are capi-
talized: use Lower Jurassic but upper Oligocene.
The sequence of sections should be title page, ab-
stract, introduction, materials and methods, results, dis-
cussion, acknowledgments, literature cited, tables, figure
captions, figures. The title page should include the title,
authors name(s) and address(es). If corresponding au-
thor is not the senior author, please indicate. The ab-
stract should summarize in 250 words or less the scope,
main results, and conclusions of the article. Abstracts
should be followed by a list of additional key words. All
references cited in the text must appear in the Literature
Cited section and vice-versa. Please follow a recent issue
of THE NAUTILUS for bibliographic style, noting that
journal titles must be unabbreviated. Information on
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please follow a recent issue of the journal for sequence of
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Illustrations: Illustrations are rendered either at full-
page width (maximum width 17 cm) or column width
(maximum width 8.2 cm). Please take these dimensions
into consideration when preparing illustrations. Page-
width illustrations ideally should span the entire width of
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should be avoided, square or “landscape” formats work
better. Please design plates accordingly, such that there
will be enough space left at the bottom of printed page
for plate caption. (Digital technology has made this task
much easier.)
All line drawings must be in black, clearly detailed,
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Photographs may be submitted in black-and-white or
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Photographs must be high resolution files at least 300 dpi
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If more than one figure is included in an illustration,
all figures are to be consecutively numbered (Ligures 1,
2, 3, . . . , NOT Ligures 1A, IB, 1C, ... , NOR Plate 1,
Ligure 1, . . .). In illustrations with more than one figure,
make sure that blank areas between figures is kept to a
minimum, thereby allowing for more area for each indi-
vidual figure.
Compressed files (e.g., .jpg) may be used to facilitate
transmission of files during original submission, but may
not be acceptable at final submission (see below).
Voucher Specimens: Deposition of the holotype in a
recognized institutional, public collection is a require-
ment for publication of articles in which new species-
level t&xa are described. Deposition of paratypes in in-
stitutional collections is strongly encouraged, as is the
deposition of representative voucher specimens for all
other types of research work.
The Editorial Process: Upon receipt, all manuscripts are
assigned a number and acknowledged. The editor reserves
the right to return manuscripts that are substandard or
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THEt'NAUTILU S
Volume 126 , Number 2
July 6 , 2012
ISSN 0028-1344
CONTENTS
Markus Muttenthaler Abundance and diversity of Conus species (Gastropoda: Conidae) at the
Sebastien Dutertre northern tip of New Ireland province of Papua New Guinea 47
Joshua S. Wingerd
John W. Aini
Hugh Walton
Paul F. Alevvood
Richard J. Lewis
Carole S. Hickman A new genus and two new species of deep-sea gastropods (Gastropoda:
Vetigastropoda: Gazidae) 57
Guido Pastorino A taxonomic revision of the genus Trochita Schumacher, 1817 (Gastropoda:
Diego Urteaga Calyptraeidae) from the southwestern Atlantic 68
Kazutaka Amano Two Neogene vesicomyid species (Bivalvia) from Japan and their
Steffen Kiel biogeographic implications 79
Research Note
Michael A. Rex
Jason D. Chaytor
Carol T. Stuart
Late Pleistocene Coins stimpsoni (Morch, 1S68) (Gastropoda: Buccinidae)
from a seabed core (2520 m) in the western North Atlantic
86
THE NAUTILUS 126(2):47-56, 2012
Page 47
Abundance and diversity of Conus species (Gastropoda: Conidae)
at the northern tip of New Ireland province of Papua New Guinea
Markus Muttenthaler
Departments of Chemistry and Cell Biology
The Scripps Research Institute
La Jolla, CA 92037 USA
[email protected]
Sebastien Dutertre
Joshua S. Wingerd
Institute for Molecular Bioscience
The University of Queensland
Brisbane, Queensland 4072, AUSTRALIA
Job u W. Aiui
Ailan Awareness, Inc.
P.O. Box 337
Kavieng, New Ireland Province,
PAPUA NEW GUINEA
Hugh Walton
Pacific Islands Forum Fisheries Agency (FFA)
P.O. Box 629
Honiara, SOLOMON ISLANDS
Paul F. Alewood
Richard J. Lewis
Institute for Molecular Bioscience
The University of Queensland
Brisbane, Queensland 4072, AUSTRALIA
ABSTRACT
In this study we conducted a survey of the diversity and abun-
dance of species of the genus Conus over two macrohabitats in
the eastern rim of the Bismarck Archipelago at the northern
tip of New Ireland province of Papua New Guinea. The main
aim was to set a baseline for the future use of Conus species
as a biodiversity indicator to monitor human impact and con-
servation efforts in this region. We observed 422 live cone
snails on 10 reefs covering 30 different species, with an over-
all diversity index of 0.924. The two habitats displayed a 74%
proportional similarity with an average Conus density ol
24 ± 9/ha (x ± SD, total sampled surface area: ~0.652 km2).
Conus flavidus, C. miles , C. sanguinolentus , C. distans and
C. litteratus were the five most abundant species. Night sam-
pling at marine-protected Lissenung Island highlighted sub-
stantial differences between day and night observations.
Sampling at 30 m depth confirmed the presence of the
recently identified C. moncuri in this area. Overall, genus
Conus was strongly present in all of the sampled sites and
represents a good low-cost indicator for long-term studies of
human impact and facile comparison of the health of similar,
but geographically distant ecosystems.
Additional keywords: Conus , cone snail, biodiversity indicator,
Papua New Guinea, New Ireland, abundance
INTRODUCTION
Conus is the largest known genus of marine inverte-
brates, with over 500 extant species, although the total
described number of species may be much higher with
more than 2,000 species catalogued between 1758-1998
(Filmer, 2001). Conus belongs to the superfamily
Conacea (suborder Toxoglossa), collectively referred to
as the toxoglossate gastropods (Taylor et ah, 1993;
Olivera, 2002). Cone snails are found in tropical marine
environments and are particularly prominent around
coral reefs and other shallow-water tropical marine hab-
itats (Kohn, 1967; Kohn, 1968; Kohn, 1971; Kohn and
Nybakken, 1975; Heck and Wetstone, 1977). In particu-
lar, the “Coral Triangle” in the central Indo-Pacific
region, encompassing parts of Indonesia, Papua New
Guinea (PNG), Malaysia and the Philippines, is well
known for its rich marine biodiversity and considered a
hot spot for Conus with more than 30 different species
observed to co-occur on a single reef (Kohn, 2001). At
the same time, it is an area that is severely threatened
by human impact due to over-fishing, pollution and
expanding populations (Allen and Werner, 2002). Highly
prized by collectors, the shell of cone snails comes in a
variety of patterns and colors, making them attractive
and valuable items. Yet, the decorative shell of the cone
snail provides no clue to its reputation as one of the most
venomous creatures on the planet (Nelson, 2004).
Indeed, all Conus species possess an efficient venom
delivery system used for prey capture and defense
(Olivera, 2002). The venom is a complex mixture
containing hundreds of highly structured biologically
active peptides (Davis et al., 2009), which functionally
modulate various ion channels to induce paralysis (Han
et al., 2008; Lewis, 2009). Only a small number (< 100)
of these venom peptides have been characterized so far,
still these peptides already afforded a drug of proven
clinical utility (Prialt, isolated from C. magus (Miljanich,
2004)), several pre-clinical leads for CNS disorders and
many valuable tools for neuroscientists, in particular for
the treatment and understanding of pain. It should
therefore be of high interest to increase conservation
efforts to preserve this remarkable pharmacopeia.
Hence, we set out to investigate the diversity and
abundance of the genus Conus at the northern tip of
New Ireland province of Papua New Guinea (Figure 1),
Page 48
THE NAUTILUS, Vo!, 126, No. 2
to increase local understanding of the benefits of marine
protected zones, to train local research staff, and to
assess if cone snails can be used as a local biodiversity
indicator for long-term monitoring of human impact and
conservation efforts in this region.
Mollusks in general are one of few phyla routinely
used as biomarkers in marine biodiversity surveys and
are considered an appropriate indicator group for the
rapid assessment of biodiversity of organisms inhabit-
ing coral reefs (Wells, 1998; Wells and Kinch, 2003).
Although Conus abundance and diversity are inversely
proportional to living coral cover on Indo-Pacific reefs
(Kohn, 1983), its presence and diversity adjacent to reefs
relates to a healthy ecosystem since they are predators of
many other animals (worms, mollusks, and fish). Addi-
tionally, species identification is unproblematic due to
the long interest of numerous malacologists and shell
collectors providing quality information on mollusean
shell patterns surpassed probably only by that on fish.
Three earlier studies conducted in PNG (Figure 1), two
in 1997 and 2000 in Milne Bay province (Wells, 1998;
Wells and Kinch, 2003), and another one in 1997 in the
Madang area (Kohn, 2001) already reported high diver-
sity of Conus, observing 66 different species in the Milne
Bay area and 45 in the Madang area. The survey
described in this publication observed 422 live Conus
specimens of 30 different species in the Kavieng area at
the northern tip of New Ireland province of PNG
(Figures 1-3). This strong presence and diversity of
Conus in this region in combination with its correlation
to ecosystem health (Veron, 2000; Roberts et ah, 2002)
warrants its use as an indicator for long-term monitoring
ol human impact in this region.
MATERIALS AND METHODS
Study Sites: Ten different reefs were investigated
between the 8-14 July 2010 for cone snail abundance
and diversity in Papua New Guinea (Figure 3). The sites
selected for this study were located at the northern tip of
New Ireland (Tok Pisin: Niu Allan, approximately 2°37' S,
150°45/ E) province, a long and narrow island (approxi-
mately 8650 km-) situated on the eastern rim of the
Bismarck Archipelago. Two seasons are experienced
annually in this area: diy southeasterly trade winds blow
almost continuously between May and August, and the
northwesterly, rain-bearing monsoon prevails from
December to March. The months between April and Sep-
tember are generally recommended for such expeditions
due to aerial reef exposure at low tides (0. 2-0.6 m), which
significantly facilitates sampling. The water temperature
does not vaiy much over the year, ranging from 28-30° C.
Different marine environments in the area surround-
ing Kavieng were investigated at depths to 30 m. Small
islands on barrier reef and the rocky island of New
Ireland were surrounded by multiple reef types, includ-
ing coral-rich fringing reefs, reef shelves, patch reefs,
sea grass and sandy-bottom reef flats (Table 1 and
Figures 1-3. Location of study sites in the Bismarck Archi-
pelago / Papua New Guinea. 1. Geographical overview of the
three study sites: Milne Bay area (1997, 2000), the Laing Island
and Madang area (1997), and the Kavieng area (2010, this
paper). 2. Close-up on New Ireland, showing Kavieng area,
which is situated at the northern tip of New Ireland Province.
3. Close-up on the Kavieng area, displaying the reef locations
that were studied. The lagoon-like sites were labeled N1-N7
and the outer reef sites on the East Coast of New Ireland were
labeled E8-E10.
M. Muttenthaler et al., 2012
Page 49
Table 1. Description of the individual study sites including GPS coordinates, measured sample area, description ol habitat,
sampling time, high and low tides, number of people collecting per site and calculated sampling efficiency based on surface area and
sampling effort.
# Site name GPS Area m2 Habitat Time Tides Depth People Efficiency
A Northern Island platform reefs
canyons
Figures 4-9 Figure 3). The inner reefs and reef crests
were exposed during low tides and explored on foot
during these times. The outer reefs and lagoons were
explored through a combination of free-diving and
SCUBA. The explored sites were grouped into two gen-
eral zones, each incorporating multiple reef environ-
ments and microhabitats. The Northern Islands (NI)
group consisted of an island chain on the western side
ol New Ireland connected by a barrier reef, extending
south from Kavieng to the western coast of Manus Island
(Figure 3). This area displayed a rich environmental het-
erogeneity including barrier reel, fringing reel, reef
shelves, patch reefs, lagoons and subtidal reef flats.
The outer islands, in particular Nusa Island (site Nl in
Figure 3), contained many boulders, sand-filled depres-
sions and few corals, a preferred habitat for many cone
snail species. A few islands on the inner (eastern side)
barrier reef were also explored including a marine
protected zone at Lissenung Island (Site N7 in Figure 3).
This inner region was made up of an intra-lagoon patch
reef and reef flat environments. The southern end of a
barrier reef terminated in a reef wall between two small
islands in an area called Albatross Passage (N6), which
was also explored. The NI region generally exhibited slow
Page 50
THE NAUTILUS, Vol. 126, No. 2
Figures 4-9. Habitats and reef morphologies of the study sites. 4-7. Northern Islands Region. 4. Large subtidal reef flat of Nusa
Island (Nl). 5. Rocky and coral-rich patch reef of Edmagon Island (N2). 6. Rocky limestone reef flat of Limus Island (N4). 7. Shallow
sandy lagoon of the southern part of Nusa Island (Nl). 8-9. East Coast Region. Typical fringing reef environment found in the EC
region consisting of boulder and coral-rubble rich inner reef flats, limestone benches and coral-rich canyons in the outer reef.
M. Muttenthaler et al., 2012
Page 51
surface currents with low wave energy on reef edges, as it
was located on or inside of a protective barrier reef.
The second zone incorporated the northern tip of
the East Coast (EC) of New Ireland and exhibited less
environmental heterogeneity than the NI region. The
EC region was composed of fringing reef extending to a
reef shelf into deep, open water and was explored from
Kaselok to Fangalava (Figure 3). This region experiences
strong surface currents at the reef shelf as well as
increased wave energy where the fringing reef drops off
as a reef shelf, due to the exposed and unprotected
nature of the reef facing the open ocean.
Sampling Methods: Sampling was conducted at low
tides by a combination of walking on aerially exposed
reefs and snorkeling at the edges of the exposed reef
platforms and lagoons at depths of 0-5 m. Boulders were
overturned and examined for live snails, trails in sand
were followed up and coral rubble was examined. Site
N6 was sampled at a depth of 5-30 m and site N7 at a
depth of 5-15 m by SCUBA. Identification of the species
was carried out using the shell book ‘Manual of the living
Conidae’ (Rockel et ah, 1995) either on site or at a later
stage, when the periostracum was removed from col-
lected shellswith bleach. Conus moncuri was identified
via correspondence with the cone snail expert R. M.
Filmer. Only live cone snails were recorded. Most obser-
vations had to be conducted during the daytime (Table 1 )
since nocturnal observations were not feasible, with the
exception of N7, where necessary facilities were in place.
The number of people sampling varied at individual
sites (Table 1) due to the educational training program
requirements between the University of Queensland
and the National Fisheries Authority. Time sampling
was preferred, as initial attempts to use spatial transects
indicated a low Conus density, which would result in an
inefficient use of time within the intertidal zones. Hence
only time-relative densities were used for analysis. Sam-
pling effort at the individual sites correlated well with the
estimated surface area (Figure 10, r“ = 0.9389, using
GPS coordinates and a surface measurement software
www.freemaptools.com/area-calculator.htm). The method
had the distinct advantage of time efficiency in the inter-
tidal zone, which allowed observers to cover large reef
areas within reasonable time limits.
Accumulating effort
(# of people x sampling time in h)
Figures 10-13. 10 Linear correlation of measured surface area and sampling effort (r2 = 0.9389). 11 and 12 Calculated
macrohabitat-specific (11) and site-specific (12) sampling efficiencies.. 13 Correlation of live Conus species observed during the
day with accumulation of effort.
Page 52
THE NAUTILUS, Vol. 126, No. 2
In total, 16 hours were spent sampling over an esti-
mated total area of 0.652 km“. Considering an average
of 1000 m that a person could sample per hour (based
on the first three sites N1-N3), an overall sampling
efficiency of 26% was observed, reflecting factors such
as reef moiphologies, accessibility. Conus habitat (boul-
ders, coral rubble, etc.) and type of sampling (reef walk-
ing, snorkeling, diving). The sampling efficiency was
calculated using the following formula: E [%] = {sam-
pling time [h] * number of people) * 1000 [in2 IT1]
*100 [%] / surface area [m-]. The EC sites were gener-
ally more accessible than the NI sites, mainly due to reef
morphology, which correlated well with the individual
sampling efficiencies observed per site (Figure 11 and 12).
Increasing collecting experience might also have contrib-
uted to the higher efficiencies at the EC sites, which
were sampled last.
Data Analyses: The study sites were marked using
a handheld Garmin 60CXS GPS system and the sam-
pled surface areas were measured using GPS guided
surface area calculating software available online (www
.freemaptools.com/area-calculator.htm). All GPS coordi-
nates are in the world geodetic system 1984 format.
Conus species abundance was determined by measure
ol elfort, based on the absolute numbers of species
observed divided by the number of people sampling and
by the sampling time in hours. Conus species diversity was
determined using the Simpsons Index D = l-^)(n/N r,
where n is the total number of organisms of a particular
species and N is the total number of organisms of all
species. The Simpson’s Index was used instead of the
Shannon-Weaver Index because it is less sensitive to
sample size and therefore more representative for diver-
sity when densities are low (Kohn and Nybakken, 1975;
Routledge, 1979). The proportional similarity of the two
habitats was carried out using the Sorensen Index
QS=2C/(A+B), where A and B are the number of spe-
cies in NI and EC, respectively, and C is the number of
species shared by both habitats. The time-related Conus
species densities were calculated based on the number
Table 2. Absolute numbers of live cone snails observed per site.
M. Muttenthaler et al., 2012
Page 53
of Conus species individuals found per person per hour
at each site. The space-related densities were derived
from the time-related densities considering ~1000 nr
that a person could sample per hour (based on the first
three sites N1-N3) including the sampling efficiency
(% of actual sampled surface area) (Figure 17).
RESULTS
In total, 422 specimens of 30 different species were
observed, out of which 26 species were found in the
NI region and 17 in the EC region (Table 2). Thirteen
Conus species ( canonicus , chaldaeus , distans, ebraeus,
flavidus, frigidus, imperialis, leopardus, lividus , miles,
rattus, sanguinolentus, and virgo) were common in both
habitats, five Conus species (catus, emaciatus, musicus,
litteratus, and textile) were only found in the NI region,
four Conus species ( coronatus , miliaris, striolatus, and
tulipu ) only in the EC region and seven Conus species
( arenatus , aulicus, geographus, marmoreus, striatus,
terebra, and vexillum) were only found at night at the
protected marine zone Lissenung (N7) (Figure 14).
Conus moncuri, which has so far not been observed in
the PNG area, was found at a divide in the barrier reef,
called the Albatross Passage (N6) at a depth of 30 m
(Figure 3). The majority of species found was worm
hunters (21), followed by fish hunters (5) and mollusk
hunters (4) (Figure 15).
The Simpsons Diversity Index for the whole region
was 0.924, for the NI region (N1-N5) 0.858 and for the
EC region 0.882. N6 and N7 were analyzed separately
due to site and sampling differences (Table 1). However,
moncuri
geographus
aulicus
arenatus
vexillum
striatus
textile
terebra
virgo
marmoreus
- East Coast (EC8-EC10)
Northern Islands (N1-N5)
Lissenung Island (N7)
» Albatross Passage (N6)
leopardus
musicus
catus
emaciatus
litteratus
coronatus
strialatus
chaldaeus
canonicus
imperialis
ebraeus ,««* .
tulipa
distans
rattus
frigidus
miliaris
lividus
flavidus
sanguinolentus
miles
0 0.5 1 1.5 2 2.5 3 3.5 4
Conus observed per person per hour
Figure 14. Conus species abundance of Northern Islands
(blue). East Coast (red), Lissenung Island (green) and Albatross
Passage (purple).
both sites were located within the NI area and the diver-
sity index including N6 and N7 was calculated to be
0.918. Tire Simpsons Diversity Indices for each individ-
ual site are listed in Figure 16 with site EC 10 having the
highest diversity with an index of 0.888. The Diversity
Index of the protected marine zone Lissenung (N7)
was 0.857. In comparison, the Serensen Index showed a
74% species similarity between the two macrohabitats
(N1-N5 vs. EC8-EC10), with a 62% species similarity
observed when the sites N6 and N7 were included.
The total number of each species found in each
macrohabitat is listed in Figure 15. The five most abun-
dant species across the regions studied were C. miles, C.
flavidus, C. sanguinolentus, C. distans , and C. litteratus,
in the EC region C. miles, C. sanguinolentus, C. flavidus,
C. lividus, and C. miliaris , and in the NI region C.
flavidus, C. miles, C. litteratus, C. distans, C. leopardus
and C. marmoreus, ranked by their abundance respec-
tively (Figure 14). Tire EC area showed slightly higher
abundance with 3.0 ± 0.3 (x ± SD) cone snails per
1000 m (or per hour per person) than the NI area (Nl-
N5) with 2.0 ± 0.9 (x ± SD) (Figure 17). This relates to
an overall mean density of 2.4 ± 0.9 (x ± SD) Conus per
1000 m . The curve of newly identified species observed
with increasing sampling effort tends to level off indicat-
ing that the species found in the Kavieng area represent
an adequate reflection of the entire community, exclud-
ing diurnally buried species as well as cone snails found
at depths > 5 nr (Figure 13).
DISCUSSION
Many factors make the genus Conus an important biodi-
versity indicator, particularly for ecosystems adjacent to
reefs where corals are generally absent or scarce (e.g.:
mud, sand, and rubble bottoms). These factors include
high species diversity, its global presence in tropical
waters, well-established taxonomy, ecological and eco-
nomic importance, its survival over millions of years and
its strong correlation to local fish and coral biodiversity
(Veron, 2000; Roberts et ah, 2002). 422 live cone snails
encompassing 30 different species were observed in the
two main macrohabitats, the Northern Island (NI, includ-
ing fringing reef, barrier reef, platform reefs and lagoons)
and the East Coast (EC, fringing reefs), over a total area
of ~0.652 km2. These numbers warrant enough abun-
dance and diversity to measure ecosystem health and
address human impact if assessed on a regular basis.
26 species were found in the NI region and 17 species on
the fringing reefs of the EC area. The two macrohabitats
displayed a 74% proportional species similarity with 1 3 spe-
cies present in both regions (Figure 14). Of these 13,
C. flavidus, C. miles, C. sanguinolentus, C. distans, and
C. frigidus were the five most abundant. By contrast,
C. miliaris, C. tulipa, C. striolatus, and C. coronatus were
confined to the EC habitat, while C. litteratus, C.
emaciatus, C. catus, C. musicus, and C. textile seemed to
prefer the NI region. The NI region had a richer display
Page 54
THE NAUTILUS, Vol. 126, No. 2
Figure 15. Number of Conus species and individuals observed including their type of prey.
of environmental heterogeneity compared to the EC
region including multiple reef types such as barrier reef,
patch reef, fringing reef and sandy-bottom reef flats.
These different microhabitats in turn influence Conus
presence at the studied sites: Conus litteratus and
marmoreus, for example, were not only confined to the
NI region, but more particularly to sandy bottoms devoid
of corals and strong currents. Conus flavidus, sanguinolentus,
and ebraeus were strongly present at site Nl, which
contained a large subtidal reef flat with many boulders
and sand-filled depression (Figure 4), yet both species
were absent at neighboring site N2, which was a coral-
rich rocky limestone reef (Figure 5).
Many cone snails are nocturnal predators and are hid-
ing under rocks/coral rubble or are buried in sand during
the day (Terlau and Olivera, 2004). Lissenung Island
(N7) had the facilities necessary (dive resort, high-
energy light sources, trained dive staff and local guides)
to conduct night sampling, which allowed us to observe
some of the nocturnal species. Night sampling resulted
in observation of 7 out of 10 Conus species ( marmoreus ,
terebra, st rial us, vexillum, arenatus, milieus, and geographus)
that were not found at any of the other sites (Figure 14)
strongly suggesting that night sampling should lie incor-
porated in future surveys. Lissenung Island was also tire
only protected marine zone in this area (no fishing or shell
collecting), which could have been another factor contrib-
uting to the high abundance and diversity observed at this
site (Figure 16).
Two earlier studies reporting on Conus species diver-
sity in PNG were part of the Rapid Marine Biodiversity
Assessment conducted in 1997 and 2000 that covered
the area of the Milne Bay Province, which encompasses
the extreme southeastern tip of mainland Papua New
Guinea and an extensive offshore area immediately east-
ward (Figure 1) (Wells, 1998; Wells and Kineh, 2003).
They covered an area heavily dotted with islands and
shoals in the Solomon Sea separating PNG from the
neighboring Solomon Islands. The mollusk survey in
2000 included 28 different sites over a period of 1 1 days
and to date represents the most thoroughly conducted
assessment of Conus species diversity in this region.
66 different Conus species were observed with C. miles
being the most abundant and widely distributed species
in Milne Bay (15 out of 28 sites). From the 30 species
observed in the Kavieng area, all but two Conus spe-
cies (C. striolatus, C. moncuri) were also found in the
Milne Bay survey. Conus moncuri is a relatively newly
discovered species that has so far not been reported in
PNG, although it has been observed in the Solomon
Islands and the Philippines. Its described habitat is
between 30 and 50 m depth, which might be the main
reason for its rare spotting, as diving at these depths is
still not very common for shell collectors.
Nl N2 N3 N4 N7 E8 E9 E10
Figures 16-17. 16. Comparison of Conus species abundance and diversity per study site. The sites N5-6 were excluded because
only a single species was observed at these sites. 17. Cone snail density of the two macrohabitats. Northern Islands area (N1-N5) and
East Coast area (E8-E10, mean ± S.E.).
per m
M. Muttenthaler et al., 2012
Page 55
The third study conducted in PNG in 1997 covered
Laing Island and the Madang area (200 km apart.
Figure 1) and observed 45 species, which correlated
well with the species observed in this survey. Only
C. sanguinolentus and C. moncuri were not observed,
which is surprising, as C. sanguinolentus was one of the
most abundant species in our study. Conus miliaris was
the most abundant species in the Madang area, which
was also highly abundant in the Kavieng area, although
only confined to the EC fringing reef habitat. The pre-
ferred microhabitats for C. miliaris on Laing Island were
limestone bench and coral rock, both of which were
also found in abundance at all three EC sites. Differ-
ences in site selection (non-random selection to maxi-
mize Conus species diversity and density), sampled
surface area (386 m" Laing Island and 3511 irT Madang),
observation period (5 weeks), sampling methods (tran-
sects, quadrants, inclusion of dead shells) and data anal-
ysis do not allow quantitative comparison with our study,
however observations such as 36 different Conus species
co-occurring on single reef platforms and maximal
densities of 3/mf were not observed in our survey. The
studies were conducted more than a decade ago and it
certainly would be of interest to see if this rich diversity
and abundance has persisted in these areas.
Overall, Conus species are strongly present in the
northern part of the province of New Ireland of Papua
New Guinea, with 422 individuals of 30 different species
observed. The local residents may easily report their
observations, making Conus an attractive low cost bio-
diversity indicator for long-term monitoring of a wide
region. Additionally, the global presence ol the species ol
the genus enables facile comparison of similar, but geo-
graphically distant ecosystems. As expected, significant
species differences were observed between day and night
sampling recommending the inclusion of night sampling
in future surveys. It is important that the staff conducting
these surveys will receive appropriate training and gain
experience, so that the sampling efficiency only reflects
reef accessibility and morphology rather than sampling
experience. The high abundance and diversity observed
at the marine-protected zone at Lissenung Island shows
promise that Conus species can be used to monitor and
validate human conservation efforts in this region. This
project should lead to regular assessments in this region,
stimulate further studies in this field and initiate local
involvement to preserve regional biodiversity and marine
richness in Papua New Guinea.
ACKNOWLEDGMENTS
We would like to thank the people from the various
communities in the Tigak islands and East Coast of
mainland New Ireland for their understanding and
allowing us to access their reef areas. Additionally, we
thank the National Fishery Authority of PNG and Ailan
Awareness Inc for the collaborative effort and in
particular we thank all the individual people involved
in this project. Dr. Thomas Durek, Dr. Lachlan Rash,
Mr. Peter Minimulu, Mr. Casper Dako, Mr. Jacob Wani,
Mr. Shaun Keane, Mr. Dietmar Anion, John and all the
students that helped with the collection. The research
leading to these results has received funding from the
NHMRC Program Grant (569927) and from the
European Union Seventh Framework Programme
( FP7 12007 -2013 ) under grant agreement n° [254897],
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THE NAUTILUS 126(2):57-67, 2012
Page 57
A new genus and two new species of deep-sea gastropods
(Gastropoda: Vetigastropoda: Gazidae)
Carole S. Hickman
University of California
Department oi Integrative Biology
and
Museum of Paleontology
Berkeley, CA 94720-3140 USA
[email protected]
ABSTRACT
A new genus and two new species of deep-sea veti gastropod gastro-
pods extend knowledge ol the unique shell morphology, radula, anat-
omy, ecology, and distribution ol a family group of undetermined
relationship to other vetigastropods. Anomphalogaz-a moluccensis
new genus and new species is the first record of die family in die
Indonesian biogeographie realm of Wallacea. Ccdbgaza cohnani new
species is die first record of die family group from the Australian
Plate. Determinate growdi in all known gazid species is expressed in
a pronounced thickening and reflection of the final lip along widi a
descending suture. In die umbilicate species, determinate growdi is
also marked by formation of a thin reflected callus that partially or
fully isolates a hollow umbilical chamber. Unique features of die
Gazid radula include extremely elongated marginal teedi with
strongly serrate shafts and an unusual pattern of integration in which
rows of marginal tootii bases and cusps are aligned in separate rows of
rachidian and lateral teetii. Distinctive anatomical features include
expanded oral surface of the snout, antero-lateral extensions of die
foot, and an enlarged liindgut. Numerous repaired breakages of
delicate gazid shells prior to formation of die terminal lip suggest high
predation pressure. Occurrences in regions of mediane and sulfide
seepage suggest diat diese relatively large deposit-feeding gastro-
pods are nutritionally linked with unconventional carbon sources.
Original description of the family group as a tribe within
Margaritinae Thiele, 1924 was based on analysis of morpholog-
ical features. Subsequent molecular sequence data have shown
that Margaritinae is not monophyletie, and some deep-water
species described under Margarites Gray, 1847 may be more
closely related to the gazid species described herein. Because
classification of vetigastropod family group taxa is currently in a
state of flux, this treatment abstains from traditional ranked
classification and uses the family-group ending “-idae" in a
provisional sense until complete phylogenies are available.
Additional keywords: bathyal, chemosynthesis, methane, sul-
fide, cold seep. Molucca Sea, Australia, deposit feeding
INTRODUCTION
Morphological support for monophyletie vetigastropod
groups, including monophyletie Turbinidae and Trochidae
(Hickman and McLean, 1990; Hickman, 1996, 1998) is in
increasing conflict with the results oi more recent molecu-
lar analyses (e.g., Geiger and Thacker, 2005; Yoon and
Kim, 2005; Williams and Ozawa, 2006; Kano, 2008;
Williams et ah, 2008, 2010). Although monophyly of some
ol the morphologically based subfamilies and tribes is well
supported by uew genetic data, there have been some
major realignments of their relationships to one another.
New morphological and/or molecular data have led to a
revised working classification (Bouchet and Rocroi, 2005)
and redefinitions of Trochidae, Turbinidae, and Trochoidea
(Williams et ah, 2008).
Some of the most interesting realignments are
among higher taxa restricted to habitats in the deep
sea (>200 meters). Recurring features that appear to be
convergent in deep-water vetigastropods include minute
shell size (<5 mm), reduced shell thickness and visibility
of nacre (tabular aragonite) through a thin exterior layer,
extreme thinning or putative loss ol nacre, secondary loss of
coiling resulting in a limpet shell form, enlargement oi the
hindgut associated with deposit feeding, elaboration of
structures for sperm transfer and sperm storage, and, in
some cases, hermaphroditism.
Examples of hypothesized molecular unmasking of con-
vergence include the recognition of two discrete clades of
minute slit-shells formerly united under Seissurellidae
(Geiger and Thacker, 2005) and recognition of a close
relationship among minute shells of seguenziiform taxa,
large- and small-shelled deep-sea trochiform taxa, and a
number of minute skeneiform taxa (Kano, 2008). Taxon
sampling for molecular phylogenetic analysis is still prob-
lematic and dif ficult for the many deep-sea taxa that are
known only from shells and occasional formalin fixed
specimens in wet collections.
A major gap in sampling is that of the deep-sea family-
group Gazidae Hickman and Mclean, 1990. Eight spe-
cies in two genera, Gaza Watson, 1879, and Callogaza
Dali, 1891, have many shared features (Hickman and
McLean, 1990) that unite them phylogenetically (Hickman,
1996, 1998). These include shared character states for
Page 58
THE NAUTILUS, Vol. 126, No. 2
38 of 40 radular characters (Hickman, 1996). However,
the original placement of gazids as a sister group to
margaritids (Hickman and McLean, 1990: Hickman,
1996) has not been tested by molecular sequence data.
A recent review of Gaza and Callogaza and description
of a ninth species from Brazil (Simone and Cunha,
2006) did not address relationships, but it underscores
the global distribution of the group and the potential
for undocumented species.
Molecular sequence data for margaritid species resulted
in a new phylogenetic hypothesis ol the relationship of
the family group to other vetigastropods (Williams et al.,
2008). However, subsequent analyses have demonstrated
that the group is not monophyletic (Williams et. al.,
2010). Further molecular genetic analyses are likely to
alter the relationship of gazids and margaritids to each
other as well as to other vetigastropods (Williams, pers.
comm., 2011).
The objectives of this paper are (1) to describe a new
gazid genus and two new gazid species, and (2) to pro-
vide new data highlighting the unusual features of the
group, its biogeography, its ecology, and its role in the
deep sea.
The family group ending -idae is used provisionally and
without placing Gazid ae into a ranked classification of
higher taxa of vetigastropods. It does not alter its currently
unresolved relationship to the family group Margaritidae,
although morphological evidence is presented to sug-
gest that some deep-water species currently assigned to
Margaritidae may require re-allocation to Gazidae.
Abbreviations for specimen repositories are: AMS,
Australian Museum, Sydney; LACM, Los Angeles County
Museum of Natural History, Los Angeles; UCMP,
University of California, Berkeley; USNM, National
Museum of Natural History, Smithsonian Institution,
Washington, DC.
SYSTEM ATICS
Vetigastropoda (unranked)
Family Group Gazidae Hickman and McLean, 1990
(unclassified)
Genus Callogaza Dali, 1881
Type Species: By original designation, Callogaza
watsoni Dali, 1881. Recent, off Havana, Cuba, 24°34' N,
83° 16' W, 177 fathoms. Distribution: Northern Cuba
south to Brazil, 117-500 fathoms.
Description: Shell of 1.5 smooth protoconch whorls
and 6 teleoconeh whorls with distinct fine spiral and
axial sculpture. Smaller than Gaza Watson, 1879 (width <
25 nnn), relatively broader, and prominently pigmented
with alternating white and brown splotches or fine spiral
lines on a beige background. Teleoconeh whorls shoul-
dered and finely carinate. Umbilical callus thinner than
in Gaza and convex when completely covering umbilicus.
Callogaza colmani new species
(Figures 1-10, 12, 14-15)
Gaza (Callogaza) sp. — Hickman and McLean, 1990,
figs. 53b, c, e, f. (originally mislabeled; reported cor-
rectly here)
D escription (Figures 1-8): Shell width (up to
23.3 mm) exceeds shell height (up to 17.4 mm), and final
whorl height exceeds that of spire. Aperture is strongly
prosocline, inclined to axis of coiling at an angle of 50°.
Terminal lip both thickened and reflected and marked
by a descending suture. Thin, transparent parietal Callus
continuous from suture to umbilicus, where it is reflexed
as a thin, slightly convex covering, creating a closed
umbilical chamber. Most abapical of four spiral cords
marks periphery of shell, and suture follows fourth spi-
ral. Most apical spiral cord separates a slight shoulder
from rest of whorl. On shoulder, 15 very fine spiral lines
crossed by slightly stronger, finely spaced axial lines to
produce a very finely cancellate surface (Figure 7). A
similarly fine cancellate surface occurs between each of
pairs of spirals between shoulder and periphery. There
are 27 very finely incised spiral lines on base (Figure 8).
Shell pigmentation consists of alternating tan and white
patches on spiral ribs, with two spiral rows of prominent
white blotches on base. Nacre visible only through small
windows of transparent shell material and only when
shell is moved, generating lustrous flashes. Protoconch
is small, glossy, and of 1.5 whorls.
Radula (Figures 9-10, 12): Rachidian is largest tooth
in central field. Rachidian tooth with broad, ovate base
with triangular over-hanging cusp that emerges directly
from base and is finely denticulate along both margins
(Figures 9, 12). Seven pairs of lateral teeth present,
also with broad bases and triangular, denticulate cusps
(Figures 9, 12). Inner portion of each lateral base is
obscured by overlapping base of adjacent tooth. Cusp of
each lateral tooth directed toward midline, overhanging
outer portion of base of adjacent lateral tooth (Figure 9).
Outermost lateral teeth with elongate shafts and nar-
rower, longer cusps increasingly curved inward toward
rachidian. Between outermost lateral and marginal teeth,
in position occupied by lateronrarginal plate in general-
ized troehoidean radula, there are several irregularly-
developed tooth bases that have failed to develop shafts
and cusps (Hickman and McLean, 1990, Figure 53E,
p. 92). Marginal teeth (Figure 10) with extremely long,
well-developed shafts and long, narrow, overhanging den-
ticulate cusps. Marginal cusp rows and base rows do not
correspond (. sensu Hickman, 1984a). Prominent serration
of the marginal shafts (Figure 10) is a unique feature.
Outermost marginal teeth have unusually broad shafts
and cusps and may be partially or fully fused to one
another (Figure 10).
Exterior Anatomy (Figures 14—15): Deep transverse
groove (inferred pedal gland) present along anterior
margin of the propodium. In retracted and preserved
animal, anterior propodium and mesopodium folded
under and against sole of foot, with oral disk of snout
C. S. Hickman, 2012
Page 59
Figures 1-8. Callogaza colmani new species. 1. Apertural, 2. Basal, and 3. Apical views of the holotype, AMS 115689, height =
17.4 mm. 4. Apertural, 5. Basal, and 6. Apical views of paratype 1, AMS 115689a, height = 17.9 mm. 7. Details of sculpture on apical
whorls of holotype. 8. Details of sculpture on base of holotype.
pressed against fold. Large snout cylindrical, expanding
distally to a tentacular margin surrounding broad oral disc
densely covered with shorter tentacles. Mouth lies within
longitudinal groove on oral disc. Large, black-pigmented
eyes lie at distal ends of short, thick, dorsoventrally flat-
tened eyestalks. Inner basal margins of eyestalks continue
over bases of cephalic tentacles, joining small crescent-
shaped cephalic lappets. Lappets have entire margins
and do not reach midline of snout. Cephalic tentacles
relatively long even in their contracted state. No evidence
of micropapillae on either cephalic or epipodial tentacles,
but this may be a preservation artifact. Left (inhalant) and
right (exhalant) neck lobes large and well-developed flaps
of tissue with simple margins. Beneath each necklobe and
continuing posteriorly along side of foot is a row of prom-
inent upraised papillae, six on right side and seven on left.
(It is not known whether these papillae are innervated and
sensory in function or glandular and secretoiy, although
they are referred to here as epipodial sense organs. They
are similar in appearance to the epipodial sense organs
that are paired posteriorly with each of the epipodial ten-
tacles.) Seven epipodial tentacles present on right side of
animal and six on left. Each tentacle and its basal sense
organ arise beneath a separate, thin, crescent-shaped flap
of epipodial tissue. Epipodial flaps partially overlap one
another in imbricate fashion. Epipodial tentacle length
decreases posteriorly on both sides. On left side, ante-
riormost epipodial tentacle lacks a flap and basal sense
organ and is longer and set apart from other five tentacles.
Mantle cavity of holotype not dissected, but long free tip
Page 60
THE NAUTILUS, Vol. 126, No. 2
Figures 9-13. Radulae. 9-10. 12. Callogaza colmani new species. 9. Raehidian and left lateral teeth of holotype, AMS 115689,
scale bar = 100 pm. 10. Right marginal teeth of holotype, bar = 100 pm. 12. Detail of raehidian base and serration of raehidian and
inner lateral cusps of holotype, scale bar = 40 pm. 11. Raehidian and left lateral teeth of Gaza superba Dali, 1881, UCMP D-3763.
13. Raehidian and inner lateral teeth of Margarites simbla Dali, 1913, LACM 71-374.
of bipectinate ctenidium projects from beneath mantle
margin of preserved specimen.
Remarks: The new species is distinguished from
Callogaza watsoni, by complete nacreous terminal clo-
sure of the umbilicus and relatively higher spire. The
spire is relatively lower than that of Callogaza sericata
(Kira, 1959) from Japan, which also has a smaller shell,
forming its terminal aperture at half the size of adult
C. colmani. The Japanese species also differs in details
of spiral and axial sculpture and pigmentation pattern.
Although the pigments are similar, they are distributed
differently, as in the arrangement ol white splotches on
the base of the shell. The radula and soft anatomy are
unknown for both C. watsoni and C. sericata , but the
shells have been adequately figured by previous authors
(see Simone and Cunha, 2006, Figures 41-48).
The shell has a relatively higher spire than the type
species and a relatively lower spire than Callogaza sericata
(Kara, 1959) from Japan. Although the Japanese species is
C. S. Hickman, 2012
Page 61
cm
Figures 14-15. External anatomy of Call ogaza colmani new species, holotype (AMS 115689) and paratype 3 (AMS 115689b).
14. Right lateral view, 15. Oblique view ot head/foot. Abbreviations: apg, anterior pedal gland; el, cephalic lappet; cm, columellar
muscle; ct, ctenidium; e, eye and eyestalk; ef, epipodial flap; esu, epipodial sense organ; et, epipodial tentacle; f, foot (sole of
mesopodium); h, head; lul, left (inhalant) neck lobe; op, operculum; me, mantle edge; p, propodium; ml, right (exhalant) neck lobe;
sn, snout; st, snout tentacles; t, cephalic tentacle.
more similar in the pigments present in the shell, they
are patterned differently, as in the arrangement of white
splotches on the base of C. sericata. The Japanese species
has a smaller shell, forming a terminal aperture at half the
size of adult C. colmani.
This is the first account of the soft parts of Callogaza ,
and the major features link it clearly with Gaza. The
large number of epipodial tentacles is a feature shared
with some species of Margarites Gray, 1847, and was a
significant factor in classifying Margaritini and Gazini as
sister tribes by Hickman and McLean (1990). Isolated
epipodial sense organs anterior to the epipodial tentacles
do occur in some species of Margarites , although iso-
lated papillae also occur beneath the neck lobes in a
number of cantharidine gastropods (Hickman personal
observation). Series of “holes” or anterior “foot orifices”
illustrated in several species of Gaza (Simone and
Cunha, 2006) are identical in placement on the foot, but
there is no evidence of holes or openings in the upraised,
papillae on the foot ol Callogaza colmani.
There is no information on the radula of other species
of Callogaza. The radula of C. colmani is similar to that
of Gaza superba (Dali, 1881) (Figure 11), although the
tooth bases in the central field are thinner and the cusps
are thinner and more denticulate. The large number of
lateral teeth (>5 per half row) was a shared feature
originally linking Gazini and Margaritini (Hickman and
McLean, 1990; Hickman, 1996).
The rachidian and lateral teeth of a thin-shelled deep-
water species originally described as Margarites simbla
Dali, 1913 have many features in common with the new
species of Callogaza and are illustrated here (Figure 13)
to call attention to two eastern Pacific species that cannot
be allocated at this time. The other species, Margarites
huloti Vilvens and Sellanes, 2006, was described as
part of a Chilean methane seep biota. Both species are
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THE NAUTILUS, Vol. 126, No. 2
thin-shelled but lack gazid determinate growth features.
Vilvens and Sellanes (2006) and Waren et al. (2011)
compared M. kuloti to two species described from meth-
ane seeps off Japan: Margarites njukyuensis Okutani,
Sasaki, and Tsuehida, 2000 and Margarites shinkai
Okutani, Tsuehida, and Fukikura, 1992. However, both of
the Japanese species have radulae with margaritid fea-
tures that include the shapes of the rachidian and lateral
teeth, the presence of a lateromarginal plate, and marginal
teeth with base rows and cusp rows that correspond with
the base and cusp rows oi the central tooth complex.
Type Material: Holotype: Australian Museum, Sydney
115689 (shell and figured animal). Figured paratype 1:
AMS 1 15689a (shell and radula). Figured paratype 2:
AMS 115689b (animal). Additional paratypes: AMS
115689c-f (shells with preserved animals), Fisheries RA7
Kara la Stn. K78-23-09, 6 November 1978.
Dimensions: Holotype: height 17.4 mm, maximum
width 23.3 mm. Figured paratype: height 17.9 mm, max-
imum width 22.9 mm.
Type Locality: Off Point Danger, New South Wales,
Australia, 27° 55-57' S, 154° 03' E, 550 m. Point Danger
is not recognized as a place name by the Australian
Department ol Natural Resources, Division of National
Mapping. It is, however, on Admiralty Charts used by the
Kapala, and P. H. Colman (pers. comm., 2008) traces its
origin to 1770 when Captain James Cook gave the name
of “Mount Warning” to a prominent feature onshore and
“Point Danger” to the corresponding offshore point where
shoals lay. Cook further noted: “Point Danger is the
boundary point on the coast between NSW and Qld.”
Distribution: The bathymetric and geographic distribu-
tions of this species are yet to be determined, but the recov-
ery ol 7 live specimens in a single dredge haul suggests that
it may be locally abundant. The same haul contained Uiree
live specimens of another large-shelled vetigastropod, the
calliotropid Calliotropis glypta (Watson, 1879).
Etymology: Named in honor ol Philip H. Colman,
formerly of the Australian Museum, Sydney, who was on
the dredging expedition and called the material to the
attention ol the author.
Genus A nomphalogaza new genus
Type Species: Anomphalogaza moluccensis new spe-
cies, by monotypy.
Description: Shell lacking umbilicus, shell height (up
to 29.2 mm) exceeding shell width (up to 22.1 mm).
Interior nacreous layer thickest, covered by two distinct,
unpigmented outer layers, No evidence of opalescent
sheen from underlying nacreous layer. Nacreous layer,
as well as outer layers, thickened and re Hexed at terminal
aperture. Terminal aperture descending. Nacreous pari-
etal callus divided into two portions, one on upper pari-
etal wall immediately below upper termination of outer
lip and other on lowermost parietal wall and connected
to eolumellar callus. Thin, transparent shell layer of
unidentified composition superimposed as secondary
callus on underlying eolumellar and parietal callus and
continuous across parietal wall. Final whorl sculpture
consisting of 130 closely-spaced, very finely-incised spi-
ral lines crossed by fine closely-spaced eollabral growth
lines, visible only with magnification. Suture slightly
adpressed to preceding whorl throughout coiling.
Remarks: The thickened and reflexed outer lip is a
unique shared feature of all species of Gazidae. It is
accompanied by a descending suture immediately prior
to formation of the terminal lip. Both features are indica-
tive of determinate growth. The thin parietal and columel-
lar callus may also be terminal features, but younger
individuals will be required to determine if this is the case.
The genus is distinguished from Gaza and Callogaza
by the absence of an umbilicus by and relatively greater
shell height. Height exceeds width, and the generating
curve is tangent to the axis of coiling, precluding the
presence of umbilical space. It is further distinguished
from Callogaza by larger adult size ( >20 mm), lack of
pigmentation, and the absence both of axial sculpture
and any strong spiral elements or peripheral demarca-
tion. Shells lack the lustrous sheen typical of both Gaza
and Callogaza. The suture is slightly adpressed to the
preceding whorl in contrast to the impressed suture in
species of Gaza, and the aperture is less prosoeline.
The shell apex is worn on both holotype and paratype,
obscuring the details of protoconch size and early
teleoconch sculpture.
Etymology: an (Gr. without) + omphalos (Gr. umbili-
cus), recognizing that there is neither an open umbilicus
or an umbilicus that has been sealed over by secretion of
a terminal callus deposit.
Anomphalogaza moluccensis new species
(Figures 16-21)
Description: Same as for genus (monotypic genus).
Remarks: The holotype and paratype were both col-
lected as empty shells. Unlike many gazid species, the
apex of the shells is not perforated and the shells are
free of epi- and endobionts. However, the shell of the
paratype is corroded; and the exfoliation of outer shell
has exposed the evidence of two distinct layers. Both
shells sustained and repaired episodes of sublethal dam-
age to the growing margin of the aperture. On the holo-
type there are six repaired breakages on the body whorl
and eight on the exposed portion of spire whorls. Both
specimens repaired an irregular breakage along the
entire apertural margin immediately prior to secretion
of the terminal apertural rim.
Type Material: Holotype USNM 311367. Paratype
USNM 311367b. Both empty shells.
C. S. Hickman, 2012
Page 63
Figures 16-21. Anomphalogaza moliiccensis new species. 16. Apertural, 17. Basal, and 18. Apical views of the holotype, USNM
311367, height = 29.2 mm. 19. Apertural, 20. Basal, and 21. Apical views of paratype, USNM 311367a, height = 28.0 mm.
Dimensions: Holotype: Height 29.2 mm, maximum
width 22. 1 mm. Paratype: Height 28.0 mm, maximum
width 21 .9 mm.
Type Locality: U.S. Fish Commission, R/V Albatross, Sta-
tion 5601. 13 November 1909. 01° 13' 10" N, 125°17'05" E.
Molucca Sea Indonesia Celebes (Sulawesi). 765 Fathoms
(=1,399 m). Sand, Globigerina and pteropods.
Distribution: The genus and species are thus far
known from only a single locality at greater depth than
any of the gazid taxa described to date and in a setting
that is more geologically remote and isolated. It is iso-
lated by its tectonic setting at the western edge the
Molucca Sea Microplate, the only known site of active
collision between two facing Island Arcs. The geologic
complexity of this region is treated further in the discus-
sion because it is pertinent to understanding the geologic
and evolutionary history of the deep-sea fauna.
DISCUSSION
Biological And Paleobiological Reassessments
Gaza and Callogaza have been recognized for many years
as conspicuous elements of the tropical Western Atlantic
and Caribbean fauna (Clench and Abbott, 1943). Their
relatively large shell sizes and local abundance (Quinn,
1979) have facilitated their use as models for understand-
ing taphonomie and paleoecologieal phenomena in deep-
sea gastropods (Walker and Voight, 1994; Voight and
Walker, 1995). The Gaza Community was designated as
an example of a deep-water community type with origins
in the Eocene (Hickman, 1984b). Discoveries of chemo-
synthetically nourished communities in the northern Gulf
of Mexico (Kennicutt et al., 1985) suggest a potentially
unconventional carbon source lor these large gastropods.
Four aspects ol the morphology and biology of these
gastropods are reconsidered here: (1) the phenomenon
of determinate growth, (2) the high incidence of repair
of sublethal shell damage, (3) the umbilicate condition,
and (4) the nutritional carbon source.
Determinate Growth in Gazid Gastropods: Determi-
nate growth is manifest in three morphological features of
gazid gastropods: (1) reflection and thickening of the
outer lip, (2) reflection of the columellar lip to form a thin
callus completely or partially covering the umbilicus, and
(3) a descending suture. It is the only family-group
vetigastropod taxon in which all known species develop
these three features. The thickened and reflected lip was
Page 64
THE NAUTILUS, Vol. 126, No. 2
used as a diagnostic feature in proposal of the family-
group name (Hickman and McLean, 1990). The thin,
translucent, bubble-like callus is, likewise, a unique
derived feature of the genus Callogaza (Hickman, 1998).
In marine gastropods, determinate growth occurs pri-
marily in siphonate caenogastropods and in shallow-water,
tropical habitats (Vermeij and Signor, 1992). It is recog-
nized most often by a terminal elaboration of the aper-
ture, which may be thickened, reflected, or flaired. Other
signatures of cessation of spiral growth in caenogastropods
include the formation of a distinctive parietal callus that
may extend onto the spire and a change in the direction of
spiral growth, where it is referred to as an “ascending
suture” (Vermeij and Signor, 1992).
Although determinate growth has been considered
rare in the basal “archaeogastropod” groups (Vermeij
and Signor, 1992), there is increasing evidence of termi-
nal features that do not occur earlier in ontogeny. In
contrast to siphonate caenogatropods, changes in direc-
tion of spiral growth are typically recorded as a descend-
ing suture between the body whorl and penultimate
whorl. A descending suture immediately precedes the
formation of the thickened terminal lip can be seen in
both the new species of Callogaza and the new genus
Anomphalogaza. In other basal gastropod clades a descend-
ing suture occurs in some, but uot all, species. If there is
no terminal modification of the lip, a descending suture
may accompany a slowing of growth in the largest individ-
uals. Lor example, some species of Clanculus Montfort,
1810 have a descending suture without any terminal mod-
ification of the aperture (Hickman, personal observation).
In contrast, a descending suture is paired with a flaired
and reflexed aperture in some species of deep-water
gastropods in the genus Calliotropis Seguenza, 1903
(Hickman, personal observation).
The descending suture forces the spire of the active
crawling animal upward, increasing the angle between
the axis of coiling and the substrate. Lunctional signifi-
cance of the change in coiling geometry is not known,
nor is the function of terminal modification of the aper-
ture. Walker and Voight ( 1994) reported high shell repair
frequency in two species of Gaza in a study showing
that attempted predation is common in large-shelled
(>25 mm height) gastropods, although rare in small-
shelled species. If the ability to repair apertural damage
decreases with age, formation of a thickened terminal
aperture may provide an adaptive advantage to repro-
ductively mature adults. Irregular breakages on the
shells of Gaza superba indicate that there was no termi-
nal thickening at the time the damage occurred.
Shell Repair in Gazid Gastropods: High frequency of
shell repair in living and fossil marine gastropods serves
as in indicator of predation pressure and of predator
ability to survive attempted predation (Vermeij, 1982).
Studies of shell repair frequencies on small-shelled gas-
tropods concluded that predation pressure was low in
the deep sea (Vale and Rex, 1988, 1989). It is therefore
surprising that later studies of large-shelled deep-sea
gastropods not only show high repair frequencies but
also that the shells of these species lack the typical
anti-predator adaptations (Walker and Voight, 1994;
Harasewych, 2002). A study of repaired breakage in
pleurotomariid gastropods reported “extraordinarily high
level of unsuccessful predation for all species examined”
(Harasewych, 2002). This is surprising because these
species have an uncalcified operculum, relatively thin
shell, and open umbilicus that should render them
mechanically more vulnerable than taxa with robust
shells, no umbilicus, and a heavy calcareous operculum.
However, living pleurotomariids also appear to be well
defended chemically by a hypobranchial gland secretion
that is released in response to disturbance and shell
damage (Harasewych, 2002). High shell repair frequen-
cies were also reported in a study of species of Gaza
and large-shelled species of the calliotropid genus
Bathybembix Crosse, 1893 (Walker and Voight, 1994).
The Umbilicate Condition in Gazid Gastropods: The
terms used to describe the umbilicate condition in gas-
tropods are difficult to apply to gazids because closure of
an umbilical opening, il it occurs, is a thin reflected
nacreous layer secreted at the cessation of growth. The
shell is phaneromphalus until growth ceases and never
becomes partially or completely plugged in the sense of
hemiomphalous or cryptomphalous. The terminal partial
or complete sealing off of the umbilicus partially or
completely hides its presence as an empty chamber.
In tlie new genus Anomphalogaza, there simply is no
umbilicus, and the terminal columellar and parietal calluses
are reflected over shell only: there is no opening to seal off.
The lack of an umbilical plug contributes to the light-
ness of the shell and low investment in calcium carbonate.
The perforation of the apex in some species, rendering
the umbilicus open at both ends, is permitted by the lack
of filling. The lack of umbilical filling has permitted use of
the umbilicus of Gaza superba and Gaza olivacea Quinn,
1981 by a polychaete that is alleged to create the apical
perforation (Quinn, 1991; Voight and Walker, 1995).
Nutritional Carbon Source of Gazid Gastropods:
Gaza and Callogaza have not been identified conclusively
as elements of chemosynthetieally nourished communi-
ties. However, Gaza superba and Gaza fischeri Dali,
1889 are locally abundant on the upper slope in the Gulf
of Mexico in regions of confirmed methane and sulfide
seepage (Kennicutt et al., 1985; Carney, 1994). Live new
records of Gaza fischeri from the Louisiana slope are
included in an account of seep and vent gastropods
(Waren and Rouchet, 1993). There is no evidence of an
enriched photosynthetically derived (detrital) carbon
source to support communities dominated by gazid spe-
cies. However, the alternative possibility of diffuse seep-
age ( sensu Nesbitt and Campbell, 2004) supporting
chemoautotrophic bacterial synthesis of organic carbon
in and on the sediments is worth investigating. Possible
sources of enriched dietary carbon also include bacterial
degradation of hydrocarbons in sediments (Brooks et al.,
C. S. Hickman, 2012
Page 65
1987). In this instance, the concept of a seep community
is not synonymous with chemosymbiosis. It is, however,
indicative of an unconventional carbon source.
The enlarged hindguts of these gastropods are packed
with sediment, and sedimentary grain coatings have
been implicated as a significant source oi nutrition for
large deposit-feeding gastropods that selectively ingest
grains with the greatest surface area to volume ratio
(Hickman, 1981). The earliest occurrences (late Eocene)
of large-shelled calliotropid gastropods of the genus
Bathybembix are in a geological setting of both discrete
and diffuse methane seepage on an active continental
margin (Hickman, 2003).
Biogeographic Reassessment
The new taxa described here call attention to a more
global biogeograpliic pattern and to extra-tropical occur-
rences in the Western Pacific. Callogaza colmani is the
first representative of the family recognized from the
Australian plate with its Gondwanan origin and history.
Anomphalogaza moluccensis is the first gazid described
from the biogeographic region of Wallacea. Indonesia as
political and modern geographic unit has been assembled
over the past 55 million years from complex tectonic evo-
lution of major plate boundaries and microplates. In the
Eocene, present day Sulawesi (although not emergent as
land) was situated south of Borneo and west of its present
location in eastern Indonesia (Hall, 2001). The type local-
ity for the new gazid genus and species is on the dis-
appearing western boundary of what was once a large
sea. The ancient Molucca Sea has been squeezed
and subducted by convergence of the Philippine Sea
Plate, the Australian Plate, and the Eurasian Plate
(Widiwijayanti et ah, 2003). The disappearance of the
Molucca Sea has received considerable attention because
it is the only present-day example of active collision
between two facing volcanic arc -trench systems. Subduc-
tion beneath Halmahera to the east and beneath Sulawesi
to the west is consuming the last of the microplate
(McCaffery et ah, 1980; Hall et ah, 1995; Hall, 2000).
The thick, deformed seafloor collision complex trapped
between north arm of Sulawesi and Halmahera is, at min-
imum, only 250 km in width (McCaffrey et ah, 1980). The
potential marine biogeographic importance of this com-
plex history is that the shrinking ocean basin in which
Anomphalogaza occurs will have disappeared in another
three million years (Hickman, 2009).
The northern Arm of Sulawesi, the Molucca Sea, and
Halmahera are situated at the northern edge of the bio-
geographic region of Wallacea in the region of least
distance between Wallaces Line and Lydekkers Line.
The sharp terrestrial biogeographic breaks that define
Wallacea have been considered invisible to shallow
marine taxa, although there are recent challenges that
suggest a “marine Wallaces line” (Barber et ah, 2000).
For the deep-water fauna, the concept of Wallacea
requires greater attention to structural features, tectonics.
and longer geologic history (see Lee and McCabe, 1986).
With the knowledge that active plate margins are sites ol
venting and seepage fueling chemosynthetically based
communities, it is now possible to target sites that have
never been sampled for ehemosymbiotie taxa in the
waters of Southeast Asia and the SW Pacific.
FUTURE DIRECTIONS
New morphological and anatomical data from this
report, combined with detailed anatomical data from a
new Brazilian species (Simone and Cunha, 2006) are
consistent with previous morphological inference of
gazid monophyly, regardless of rank and classification.
The novel combination of features of shell, radula, anat-
omy, nutrition, and ecology distinguish it from other
vetigastropod family groups. However, the relation-
ships of family groups of vetigastropods are still poorly
resolved, and morphological data presented here do not
address phylogenetic relationships. For the morpholo-
gist, the challenge lies in unmasking convergence.
Strong morphological convergence in deep-sea veti-
gastropods is clearly related to deep-sea benthic ecology
and deposit-feeding (Hickman, 1981, 1984b, 2003). The
enlarged hindgut, expanded and tentacular oral disk
margin, and anteriolateral expansion of the foot also
occur in large, deposit-feeding seguenzioid gastropods
such as species of Bathybembix Crosse, 1893 and
Calliotropis Seguenza, 1903, a genus that is unusually
abundant and speciose at bathyal depths throughout the
world and especially in the Indo-Pacific (Vilvens, 2007).
Shells also have converged on visibility ol nacre through
an extremely thin outer shell layer and terminal growth
features that include flaring or thickening ol the terminal
aperture, a descending suture, and reflection of the col-
umellar callus to partially or fully cover (but not fill) the
umbilicus (Hickman, personal observation).
Thinning of the shell and terminal growth features are
also common convergent shell features in several deep-
water skeneilorm gastropod groups (Hickman, personal
observation). In describing the enigmatic Australasian
family group Kaiparathini (under Margaritinae), Marshall
(1993), noted several anatomical similarities to Gaza
superba. Kaiparathinids do not fit comfortably into exist-
ing vetigastropod classification.
The genus Margarites Gray, 1847 needs both morpho-
logical and molecular work to identify taxa that belong
elsewhere. More than 50 species have been assigned to
the genus. In addition to the deep-water species noted
above as candidates for transfer to Gazidae, there are
extinct deep-water species dating back to the Cretaceous.
Kaim et al. (2009) described Margarites sasakii from a
Campanian cold seep site in northern Hokkaido, Japan,
indicating deep origins of vetigastropods in chemosyn-
thetically based communities. Deep-water fossil species
are also described from Eocene cold-seep carbonates in
Paleogene rocks in the Pacific Northwest (e.g. Squires
and Goedert, 1991).
Page 66
THE NAUTILUS, Vol. 126, No. 2
ACKNOWLEDGMENTS
This paper is part of a project begun in 1975 to document
new taxa of deep-water vetigastropods. I am grateful to
jerry Harasewych and the late joe Rosewater for their
assistance and encouragement in the use of material from
the Smithsonian Institution and to Winston Ponder, Phil
Colman, and Ian Loch of the Australian Museum, Sydney
for assistance in the study of Australian material. I am
grateful to Jim Quinn, Bruce Marshall, and Dai Herbert,
and Jim McLean for earlier discussions of trochoidean
morphology and relationships. Sally Walker and Geerat
Vermeij have contributed to my interest in Gaza and its
allies through their fascination with unsuccessful preda-
tion, shell repair, and the phenomenon of determinate
growth in gastropods. Discussions with Suzanne Williams
have been especially helpful in considering the alterna-
tive approaches to vetigastropod taxonomy, phylogeny,
and achievement of stability in classification. I thank Jerry
Harasewych and an anonymous reviewer for helpful
suggestions for improvement of the manuscript. Marla
Coppolino rendered the anatomical drawings.
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THE NAUTILUS 126(2):68-78, 2012
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A taxonomic revision of the genus Trochita Schumacher, 1817
(Gastropoda: Calyptraeidae) from the southwestern Atlantic
Guido Pastorino
Diego Urteaga
Museo Argentine) de Cieneias Naturales
Av. Angel Gallardo 470, 3° piso lab. 80
C1405DJR Buenos Aires, ARGENTINA
[email protected]
ABSTRACT
The authors revised the Atlantic species of the genus Trochita.
Two valid species are recognized. Trochita pileus (Lamarck,
1822) and T. pileolus (d'Orbigny, 1841) are redescribed and
the type material of each species is illustrated. Both species
range from the coast of Buenos Aires province to Tierra del
Fuego. Trochita pileus is a common subtidal Patagonian species
and T. pileolus is rare and usually lives in deeper waters.
Trochita trochiformis a Pacific only species is compare with the
Atlantic representatives. Protoconch ornamentation and septum
margin are the main differences between both Atlantic species.
Trochita georgiana Powell is a junior synonym of T. pileolus.
Additional keywords: Argentina, Patagonia, geographic
distribution
INTRODUCTION
The South American species belonging in the genus
Trochita Schumacher, 1817 were known since early
times in historical malacological literature (e.g. Born,
1778; Gmelin 1791; Lamarck, 1822). Many names were
proposed from different regions, perhaps, because of
the large range of some of the species and its significant
conchological variation. Rehder (1943), in a short paper,
provided an initial account of the actual type species of
the genus Trochita and discussed some of the names in
use. Nevertheless, he did not illustrate shells, and some
older names were not included.
In this revision, we focused on the southwestern
Atlantic species of Trochita , however, because of proxim-
ity and similarity one Pacific form is also included. Type
specimens, protoconchs, and radulae of all species are
illustrated and redescribed.
MATERIALS AND METHODS
All the material considered is essentially housed at the
Invertebrate collection of the Museo Argentino de
Cieneias Naturales “Bernardino Rivadavia”, Buenos
Aires (MACN-In). Some other specimens were studied
at the National Museum of Natural History, Smithsonian
Institution, Washington, DC (USNM) and Museo de
La Plata, (MLP). Type material is deposited at Natural
History Museum, London, (NHMUK), Museo Nacional
de Historia Natural, Santiago, Chile (MNHNS), and
Museum d’Histoire naturelle de Geneve, Switzerland
(MHNG). Material from the USNM was originally col-
lected by the United States Antarctic Program (USAP)
and was collected by three ships: R/V Hero, R/V Eltanin
and R/V Professor Siedlecki. Several lots were also
collected during two oceanographic cruises aboard of
the argentine B/I Puerto Deseado.
Radulae were prepared according to the method
described in Pastorino (2005) and observed using a
Philips XL 30 scanning electron microscope (SEM) at
the MACN.
Photographs were taken using a digital camera Nikon
D100 with a 60 mm Micro lens. All images were digi-
tally processed.
SYSTEMATICS
Family Calyptraeidae Lamarck, 1809
Genus Trochita Schumacher, 1817
Type Species: Trochus spiralis Schumacher, 1817
( =Trochus radians Lamarck, 1816) by SD of Rehder,
1943 (= Trochita trochiformis (Born, 1778)).
Trochita trochiformis (Born, 1778)
(Figures 1-3)
Unnamed Knorr, 1768: pi. 29, fig. 1, 2.
Turbo trochiformis Born, 1778: 355.
Patella trochiformis varietas ((5) Gmelin, 1791: 3694.
Trochus radians Lamarck, 1816: 445, figs. 3a, b; 1822: 11.
Infundibulum radians. — Sowerby in Gray, 1839: 148,
' pi. 39, fig. 10.
G. Pastorino and D. Urteaga, 2012
Page 69
Figures 1-3. Trochita trochiformis (Born, 1778), holotype of Trochus radians Lamarck, 1816, MHNG1095/53. Scale bar = 1 cm.
Infundibulum radians. Var. — Sowerby in Gray, 1839: 148,
pi. 39, fig. 1 1 .
Trochita radians Lamarck. — Rehder, 1943: 42-43, fig. 1.
?Trochita spirata Forbes, 1852: 271, pi. 1 1, figs. la,b.
? Trochita spirata Forbes, 1850. — Reeve, 1859: pi. 2 fig. 8.
Calyptraea (Trochita) trochiformis (Rorn, 1778). — Keen,
1971: 456, fig. 804; Marincovich, 1973: 31-32, fig. 65;
Canete & Ambler, 1992 (development).
Trochita trochiformis (Rorn, 1778). — Reid & Osorio,
2000: 124, figs. 3H,I.
Trochita calyptraeformis (Born, 1778). — Collin, 2003a:
550, fig. 21, 9C; Collin, 2003b: in table 1 and 2; Collin,
2003c: 618-640; Paredes & Cardoso, 2007: 182, fig. 2g;
Canete et al., 2007: 1-7.
Type Material: [Turbo trochiformis ] not found;
[Trochus radians ] is housed at MHNG1095/53.
Type Locality: [Turbo trochiformis ] unknown; [Trochus
radians] “ merdes Antilles, proche la Guadeloupe” .
Other Material Examined: Chile: La Cuchara beach.
South of Tocopilla, 4 sp., MLP12091; Coquimbo, 8 sp.,
MACN-In24810; Antofagasta, 4 sp., MACN-Inl9580;
Antofagasta, 5 sp., MACN-Inl5090; Antofagasta, several
sp. MACN-In24745; Valparaiso, 1 sp, MACN-In29685;
Valparaiso, 4 sp., MACN-In25374; Caldera, 3 sp., MACN-
In9478;Caldera, 1 sp., MACN-In7321; Caldera, 1 sp.
MACN-In 11597-L Bahia Tangoi, 4 sp., MACN-In24795;
Caleta Horcon, 2 sp., MACN-In26030; Talcahuano,
1 sp., MACN-Inl3815; Puerto San Antonio, Provincia
de Santiago, 6 sp., MACN-Inl2866; Costas de Chile,
4 sp., MACN-Inl2164.
Distribution: According to Rehder (1943) after mate-
rial housed at the USNM, from Manta, Ecuador (1° S)
to Valparaiso (33° S), Chile. Later Keen (1971), Taylor
and Smythe (1985), and, Canete and Ambler (1992) copy
this distribution. Reid and Osorio (2000) collected mate-
rial of this species in Estero Elefantes, Chile (~45° S).
This is the southernmost locality. Paredes and Cardoso
(2007) cited the whole peruvian littoral and put some
doubt on northern citations.
Vokes (1975) cited Nickles (1950) as the source of
the presence of T. trochiformis in Cape Verde Is. and
Angola, Africa. Rehder (1943) reported Trochita sp. after
one worn specimen that could be identified also as
T. trochiformis. Finally, Taylor and Smythe (1985) men-
tioned “specimens very similar and probably conspecific
with T. trochiformis” from the same African region in the
collection of the NHMUK. Vokes (1975) discussed the
possibility that the same species could be distributed in
these two really far away regions travelling by ocean
currents. There is no modern study that assures this
hypothesis. So far, there is no record ol this species, adults
or juveniles, in the Argentine littoral. It is difficult to
accept that a non-free larvae species travels all the way
through the whole Atlantic Ocean. However, despite the
direct development ol T. trochiformis (see Canete &
Ambler, 1992), according to Canete et al. (2007) this spe-
cies performs a very particular way of moving during the
juvenile stage through planktonic drifting. Nevertheless,
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THE NAUTILUS, Vol. 126, No. 2
the African distribution of this Pacific only species should
be at least treated with care.
Remarks: Trochita trochiformis is probably one of the
commonest species of Trochita from the Pacific coast of
South America. It is commercially exploited in Chile and
Peru, however, the exact distribution range is somewhat
imprecise because of many names locally still in use
by local authors. Collin (2003a, b), Canete et al. (2007)
and, Paredes and Cardoso (2007) changed the name to
T. calyptraeformis (Born, 1778) without any explanation.
Born (1778) never proposed this name. Perhaps the con-
fusion started with the species Sigapatella calyptraeformis
(Lamarck, 1822) a valid species sometimes known under
the genus Calyptraea from Australia.
One of the main differences with the Atlantic species
is the size of the shell which is larger than both, T. pileus
and T. pileolus. Nevertheless, younger, small specimens
of T. trochiformis could he confused with T. pileus , how-
ever the number and appearance of ribs clearly differen-
tiate both species. T. trochiformis always has coarse ribs,
thick shells and more than 25 ribs per whorl. T. pileus
when present has thin, closely arranged ribs which con-
tinue through the suture of the next whorl. In addition
the septum of T. pileus is clearly sigmoid, with a char-
acteristic initial (central) portion of septum reflexed.
T. trochiformis usually has an almost straight septum
margin. Luthermore, the umbilicus is visible in T. pileus
and partially hidden in T. trochiformis.
Trochita pileolus (d’Orbigny, 1841)
(Ligures 4-12, 22-24, 34-39, 40)
Calyptraea (Trochatella) pileolus d’Orbigny, 1841: 463.
Infundibulum pileolus d’Orbigny, 1847: pi. 78, figs. 5, 6, 6’.
Calyptraea decipiens Philippi, 1845: 61.
Calyptraea Sigapatella pileolus d'Orb. — Tryon, 1886: 122,
pi. 35, figs. 91,100.
Calyptraea pileolus d’Orbigny, 1841. — Careelles, 1950: 57.
Calyptraea (Trochita) georgiana Powell, 1951: 127, Pig. E 3,
pi. 7, fig. 22; Zelaya, 2005: 119, fig. 24.
Description: Shell conical, white, short, of 3 convex
whorls, of less than 2 cm ol maximum diameter and less
than 1 cm of maximum height (H/W=0.53 ±0.07, n=36);
apex central; protoconch of about 1.5 whorls, slightly exca-
vated; first whorl of protoconch smooth, then 5-6 spiral
ribs that vanish towards the teleoconch; teleoconeh with
irregular concentric growth wrinkles; aperture circular;
umbilicus conspicuous, visible, always present; septum
straight, slightly sigmoid to the edge; periostracum usually
present, pale yellowish, transparent to dark brown.
Radula taenioglossate, rachidian tooth rectangular
with one central cusp and 3-4 small (obsolete) denticles
on each side; lateral tooth broad, with apical cusp toward
rachidian line and 7-8 irregular denticles on the external
side; two marginal teeth long, curved tall tip sharply
pointed without any trace of denticles.
Type Material: [Calyptraea (Trochatella) pileolus]
6syntypes, NHMUK1854.12.4.567; [Calyptraea (Trochita)
georgiana] holotype, NI1MUK 1961469; [Calyptraea
decipiens] not seen, probably in Museum fur Naturkunde,
Berlin (according to Ihering, 1907: 150).
Type Locality: [Calyptraea (Trochatella) pileolus] “ lies
Malouines et le continent americain, sur la cote cle la
Patagonie meridionale ”; [Calyptraea decipiens] “fretum
Magellanicum ”; [Calyptraea ( Trochita ) georgiana] near
Shag Rocks, west of Soutii Georgia, 53°43,40"S, 40°57' W
in 177 nr.
Other Material Examined: Argentina: 38°46/53" S,
55°50'30" W, in 97 m, St. 7 B/I Puerto Deseado, 1 sp.,
MACN-In39304; 38°50'56" S, 55°39'08" W, in 114 m, St.
8 B/I Puerto Deseado, 4 sp. MACN-In39303; 39°00' S,
57° 10' W, 1 sp. in 177-182 m, MACN-In 15225; Ria Santa
Cruz, 6 sp., MACN-In5171-1; 39°52/37.9" S, 59°38.6' W,
St. 22, B/I Puerto Deseado, in 96 m, 1 sp., MACN-
In39305; 54°28'58.5" S, 64°57/24.6" W, St. 1, B/I Puerto
Deseado, in 106 m, 4 shells, MACN-In39306; 54°54' S,
66° 19' W, 8 sp. in about 27 m, MACN-In24017; Punta
Sinaia, Tierra del Luego, 7 sp., MACN-In 12541; 51°46' S,
68° 45' W, 23 sp. in 22 m, MACN-In23858; 54°47' S,
63° 35' W, 2 sp. in 146 m, MACN-In22293; 54°50' S,
64°0T W, 3 sp. in 154 m, MACN-In22734-1; 55°4T S,
66° 34' W, 1 sp. in 1 15m, MACN-In24980-1; Bahia Buen
Suceso, Tierra del Luego, 3 sp., MACN-In25072-1;
Ushuaia, Tierra del Luego, 2 sp., MLP13485; Tierra del
Luego, 1 sp., MACN-In20496-1; Puerto Parry, Isla de los
Estados, 5 sp., MACN-In21918; Puerto San Juan, Isla de
los Estados, 1 sp., MACN-In22160-1; Punta Colnett, Isla
de los Estados, 1 sp., MACN-In22566-1; Puerto Olla, Isla
Observatorio, 1 sp. in 18-36 m, MACN-In22037-1; Bahia
Cumberland, South Georgia Is., I sp. in 22 m, MACN-
In 13537; Bahia Cumberland, 1 sp. in 36 m, MACN-
In 18969; Schlieper Bay, South Georgia Is., 1 sp., in 18 m,
MACN-Inl8972; Annenkov Is., Soudi Georgia, 3 sp. in
36 m, MACN-Inl8971; Larsen Harbour, Soudi Georgia
Is., 3 sp. in 27 m, MACN-In 18970; §3°32' S, 41°37' W,
Shag Rocks, 1 sp., MLP7252; 54° 30' S, 35° 50' W, 1 sp. in
94 m, MLP7284; Antarctic Bay, South Georgia Is., 2 sp. in
36 m, MACN-Inl8973; 53°58' S, 37°09' W, 1 sp. in 138 m,
MAC N-In36293 ; 53° 08' S, 35°25' W, 1 sp. in 115 m,
MACN-In36292. Chile: Isla Nueva, 2 sp., MACN-
In24957-1; Puerto Harris, Isla Dawson, 3 sp., MACN-
Inl2432-1; Punta Arenas, 2 sp. MACN-Inl2390; Punta
Arenas, 5 sp., MACN-Inl3118; Puerto Harris, Isla Dawson,
1 sp., MACN-Inl4000-1; Puerto Harris, Isla Dawson, 6 sp.,
MACN-Inl2482-1.
Distribution: Off Buenos Aires province, in about
100 m depth; subtidal from southern Santa Cruz province
(^50°S), Tierra del Luego, Isla de los Estados, Malvinas
(Lalkland) Is., South Georgia Is. in Argentina, to Punta
Arenas and Isla Dawson in Chile.
G. Pastorino and D. Urteaga, 2012
Page 71
Figures 4-12. Trochita pileolus (d’Orbigny, 1841). 4-6. Holotype of Calyptraea ( Trochita ) georgiana Powell, 1951
NHMUK1961469. 7-12. Three (7-9, 10, 11-12) syn types of Calyptraea (Trochatella) pileolus d’Orbigny, 1841
NHMUK1854. 12.4.567. Scale bars: 4-9 = 1 cm, 10-12 = 1 mm.
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THE NAUTILUS, Vol. 126, No. 2
Remarks: This is a typical example of a magellanic
species distributed along the whole malacological prov-
ince. It is subtidal in the southern region and becomes a
deep water (~100 m) species on the northern part of its
distribution. It was found usually mixed with specimens
ol Trochita pileus both on museum lots and on several
unsorted samples. However, it is easily distinguished
by the spirals on the protoconch, the open umbilicus
and the straight septum. In addition, it is smaller than
T. pileus and never presents ribs. Most ol the specimens,
including the holotype of T. georgiana , are covered,
sometimes by up to three different species of bryozoans
and/or sponges. Despite that both species, T. pileolus
and T. pileus , appear together in the samples the latter
is the only one that regardless the smooth surface partic-
ipates of this association.
Ihering (1907) mentioned T. decipiens as a synonym of
T. pileolus and according to him he has seen the type
material of Philippi's species. The size and short descrip-
tion of T. decipiens agree with this author idea.
Trochita georgiana Powell, 1951 was described without
no mention or comparison with T. pileolus (d’Orbigny,
1841). Also, several years later, Powell (1960) in his cata-
logue of Antarctic and Subantarctic Mollusca neglected
again d’Orbigny s species despite the type locality
(Malvinas Is. and Patagonian coast) is clearly part of
the subantarctic region. The holotype is housed at the
NHMUK. Its protoconch and the septum as well as the
distribution cited in the original description undoubtedly
refer to T. pileolus. It is interesting to see that Powell’s
draw of the holotype (1951: 127, pi. 7, fig. 22) is not
covered by bryozoans as it is the real shell and most of
T. pileolus specimens.
Trochita pileus (Lamarck, 1822)
(Figures 13-18, 19-21, 25-33, 41)
Trochus pileus Lamarck, 1822: 11.
Calijptraed costellata Philippi, 1845: 62; Strebel, 1906:
159, pi. 13, figs. 88-97; Carcelles, 1950: 57; Castellanos,
1970: 41, pi. 3, fig. 9.
?Trochita corrugate Reeve, 1859: pi. 2, sp. 9.
Trochita clypeolum Reeve, 1859: pi. 3, sp. 14; Powell,
1951: 127;
Trochita decipiens Rehder, 1943: 44, fig. 2 (non
Philippi, 1845).
Figures 13-18. Trochita pileus (Lamarck. 1822). 13-15. Syntype of Trochus pileus Lamarck, 1822, MPING INVE 51452/2
(ex 1095/54-2). 16-18. Probable holotype of Calijptraea costellata Philippi, 1845, MNHNS unnumbered. Scale bar = 1 cm.
G. Pastorino and D. Urteaga, 2012
Page 73
Figures 19-24. Trochita species. 19-21. Trochita pileus (Lamarck, 1822). Three views of the protoconch of MACN-In25072,
from Bahia Buen Suceso, Tierra del Fuego, Argentina. 22-24. Trochita pileolus (d’Orbigny, 1841). 22. Protoconch of MACN-
In25072-1, from Bahia Buen Suceso. 23-24. Two views of MACN-In5171 from Ria Santa Cruz. Scale bars = 500 pm.
Trochita trochiformis Powell, 1951: 126, fig. E 2 (non
Gmelin, 1791);
Calyptraea pileolus Carcelles, 1950: 57, pi. 2, fig. 25 (non
d’Orbigny, 1841). Castellanos, 1970: 40, pi. 3, fig. 5;
Trochita pileus (Lamarck, 1822). — Powell, 1960: 143;
Description: Shell conical, medium size, (HAV=0.56
±0.08, n=20) circular base of ~3 somewhat flat whorls
of about 3 cm of maximum diameter (x=1.74, n=20) and
about 1.7 cm of maximum height (x=1.05, n=20); apex
central, protoconch prominent of 1.5 whorls, first whorl
smooth, later multiple weak spiral threads that cross
growth lines and vanished on teleoconch; teleoconeh
with about 25 thin ribs on the two last whorls, first whorl
usually smooth, some specimens with irregular spiral
wrinkles only; ribs somewhat scaly when crossing growth
lines and continues thru suture to the next whorl. Aper-
ture circular, umbilicus present but sometimes covered
by septum; septum sigmoid reflexed on the umbilical
area and a sinus on the margin. Periostracum thick, yel-
low pale, thin, transparent to dark brown.
The radula showed no significant differences from
T. pileolus.
Type Material: [Trochus pileus ], three syntypes
housed at MHNG INVE 51452/1-3 (old registration
number MHNG 1095/54/1-3); [Calyptraea costellata],
one specimen probable syntype present in MNHNS;
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THE NAUTILUS, Vol. 126, No. 2
Figures 25-33. Trochita pileus (Lamarck, 1822). 25-27. MACN-In39309, Puerto San Julian, Santa Cruz, Argentina, on the clock
pilots. 28-30. MAC N -In 38191, 36°30'S, 54°44' W. 31-33. MACN-In22873, Puerto Parry, Isla de los Estados, Argentina. Scale
bar = 1 cm.
G. Pastorino and D. Urteaga, 2012
Page 75
Figures 34-39. Trochita pileolus (d’Orbigny, 1 <34 1 ) . 34-36. MACN-Inl2541 from Punta Sinaia, Tierra del Fuego. 37-39. MACN-
Inl8969, Bahia Cumberland, South Georgia Is. Argentina. Scale bar = 1 cm.
[Trochita corrugata], 3 syn types housed at NHMUK
1977148 (not seen) [Trochita clypeolum] holotype
NHMUK 1977149 (not seen).
Type Locality: [Trochus pileus] unknown, however,
there is a written label, not from Lamarcks, stating
Antilles? (Y. Finet, Com. Pers.); [Calyptraea costellata]
“fretum magellanicum” ; [Trochita corrugata] “Callao,
Peru”; [Trochita clypeolum] “Straits of Magalhaens".
Other Material Examined: Argentina: 36°30' S,
54° 44' W, 1 sp. in 26 m, MACN-In 38191; 37°40/ S,
56° 25' W, I sp., MACN-In 15805; Puerto Quequen,
6 sp., MACN-In 22921; 38°31' S, 55°42' W, 13 sp. in
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TIIE NAUTILUS, Vol. 126, No. 2
Figures 40—41. Radulae of Trochita species. 40. Trochita pileolus , MACN-In36293, 53°58' S, 37°09/ W. 41. Trochita pileus
MACN-In39309. scale bars = 100 pm.
109 m, MACN-In 23358; 38°18'38" S, 56°58'59.4" W, in
66 m, St. 5, B/I Puerto Deseado, 9 sp, MACN-In3931 1;
38°25' S, 56° 30' W, 1 sp., MACN-In30496; 38°46'53" S,
55°50'30" W, in 97 m, St. 7 B/I Puerto Deseado, 8 sp.,
MACN-In39310; 38°50'56" S, 55°39'08" W, in 114 m, St.
8 B/I Puerto Deseado, 16 sp. MACN-In39308; 39°0T40" S,
58° 10' 40" W, in 55 m, St. 11 B/I Puerto Deseado, 1 sp.
MACN-In39301; 39° 12' S, 56° W, in 110-128 m,
MACN-In20846; 39°00' S, 57° 10' W, 1 sp. in 177-412 m,
MACN-In 15225-T Sontli of Quequen, 2 sp., MACN-
In26290; 40°03' S, 57° W, 2 sp. in 91 m, MACN-
In 15689; 41°12' S, 62° 54' W, 2 sp. in 27 nr, MACN-
In20655; 42°22'02" S, 63°11'52.6" W, St. 11, B/I Puerto
Deseado, in 63 m, 1 sp., MACN-In39302; Punta Norte,
Peninsula Valdes, several sp., MACN-In 1 1480; North
of Puerto Madryn, Chubut, 1 sp., MACN-In29643;
Rawson, mouth ol Rio Chubut, 1 sp., MACN-Inl0718;
Bahia Cruz, Chubut, 1 sp., MACN-In9014-1; 44°16' S,
65° 12' W, Bahia Vera, 6 sp, in 46 m, MACN-In23745;
45°00'25.1" S, 65°16'44.2" W, St. 9, B/I Puerto Deseado,
in 85/89 m, 13 sp., MACN-In39312; 45° S, 65°32' W,
1 sp. in 86 nr, MACN-In24097; 45°08' S, 66°28' W, 3 sp.,
MACN-In23842; 45°09'S, 66°27' W, 77 sp. in 15 m,
MACN-In23809; 45°15' S, 65°26' W, 1 sp., in 97 m,
MACN-In9151; Dos Hermanns, Puerto Deseado, Santa
Cruz, 8 sp., MACN-In26188; Punta Cavendish, Puerto
Deseado, 1 sp., MACN-In26185; Isla Quiroga, Ria de
Puerto Deseado, 7 sp., intertidal, MACN-In35614; Rio
Deseado, 2 sp., in 16 m, MACN-Inl4970; Rio Deseado,
off Baliza Alianza, 2 sp., in 4 m, MACN-Inl4971; Ria
Deseado, 2 sp., 6 nr, MACN-In 14973; Bahia Uruguay,
Puerto Deseado, 21 sp., MLP266; Puerto Deseado,
Santa Cruz, 16 sp., MACN-Inl7746; 2 sp., MLP261; 9
sp., MLP4748; 1 sp., MLP4785-2 and 18 sp., MLP3892;
Punta Medanosa, Santa Cruz, 2 sp., MLP259; Cabo
Blanco, Santa Cruz, 2 sp., MACN-In23623; 47°47'39.9" S,
65°34'53.9" W, St. 5, B/I Puerto Deseado, in 46 m, 23
shells, 18 sp., MACN-In39313; Cabo Blanco, Santa
Cruz, 15 sp., MACN-Inl7745; San Julian, Santa Cruz, 5
sp., MACN-In 9200; San Julian, 2 sp, MACN-In9237;
Puerto San Julian, on the dock pilots, Santa Cruz, 15 sp.
iu 1-2 m, MACN-In39309; Punta Pena beach, San
Julian, Santa Cmz, 64 sp., MLP3024; Ria Santa Cmz,
7 sp., MACN-ln5171; Bahia Buen Suceso, Tierra
del Fuego, 9 sp., MACN-In25072; 54°28'58.5" S,
64°57'24.6" W, St. I , B/I Puerto Deseado, in 106 m, 62
shells, L sp. MACN-In39314; 54°25' S, 65°49' W, 1 sp.,
MLP4350; Puerto Parry, Isla de los Estados, 10 sp.
MACN-In22873; Puerto Parry, Isla de los Estados,
10 sp., MACN-In22874; Puerto San Juan, Isla de los
Estados, 2 sp., MACN-In22160; Puerto Roca, Isla de los
Estados, 2 sp. in 18 m, MACN-In21982; Puerto Cook,
Isla de los Estados, 14 sp., MACN-In22093; Puerto
Cook, 2 sp., MACN-In22094; Puerto Roca, Isla de los
Estados, 2 sp., MACN-In21982; Puerto Olla, Isla
Observatorio, 2 sp. in 18-36 m, MACN-In22037; Punta
Colnett, Isla de los Estados, 2 sp., MACN-In22566; Rio
del Fnego, Tierra del Fuego, 16 sp., MACN-Inl2539;
Rio Grande, Tierra del Fuego, 13 sp., MACN-Inl2540;
Punta Sinaia, Tierra del Fuego, 21 sp. MACN-Inl2357-
1; Cabo Santa Ines, Tierra del Fuego, 3 sp., MACN-
In 12538; Tierra del Fuego, 1 sp., MACN-In20496;
Tierra del Fuego, 3 sp., MACN-In21157-1; Canal Bea-
gle, Tierra del Fuego, 1 sp., MLP260; Islas Malvinas
(Falklands Is.) 3 sp., MACN-Inl0152; 51°46' S, 68°45' W,
68 sp. in 22 nr, MACN-In23858-1; 52°22' S, 68°29' W,
several sp., in 18 m, MACN-In24061; 53°6' S, 67°4'1.1 W,
in 37-46 m 1 sp. USNM896262; 53°6' S, 67°4'1.1W, in
86 m 2 sp. USNM896261; 54° 50' S, 64°01' W, 4 sp. in
154 m, MACN-In22734; 54°27' S, 63°35' W, 2 sp.,
MACN-In22292; 54°26'30" S, 64°53' W, 6 sp. in
112 m, MACN-In25025; 54°41'S, 64°01'17" W, in 55 m.
G. Pastorino and D. Urteaga, 2012
Page 77
MACN-In22613; Ushuaia, Tierra del Fuego, 1 sp.,
MACN-In 21157; Ushuaia, 5 sp., MLP4962; Patagonian
coasts, 1 sp., MACN-In9237-1; 54°4,58.7" S, 58052'L1" W,
Burdwood Bank, 1 sp. in 119 m, USNM881885; 55°41' S,
66° 34' W, 15 sp. in 115m, MACN-In24980. Chile: Bahia
Lomas, Isla Dawson, 7 sp., MACN-Inl2461; Bahia
Lomas, Isla Dawson, 2 sp., MACN-Inl2460; Puerto Harris,
Isla Dawson, 6 sp., MACN-Inl2431; Puerto Harris,
Isla Dawson, 5 sp., MACN-Inl2432; Puerto Harris, Isla
Dawson, several sp., MACN-Inl2482; Puerto Harris, Isla
Dawson, 5 sp., MACN-Inl4000; Isla Nueva, 11 sp.,
MACN-In24957.
Distribution: Off Buenos Aires province (~36° S),
Patagonian coast, Isla de los Estados, Islas Malvinas,
Straits of Magellan, Tierra del Fuego, Burdwood Bank
in the Atlantic, Isla Dawson, Puerto Harris in the Pacific.
Remarks: Trochita pileus is a common species living
subtidally on hard substrate. The costulate form was
confused with T. trochiformis but the radial ribs, nor-
mally absent, when present are thinner. The septum
margin is sigmoid and in the beginning is reflexed
while in T. trochiformis it is straighter and sharper.
In addition, the latter species has a thicker shell and
larger apertural diameter and height. The radula of
both species is also somewhat different, while T. pileus
has no denticles on the two marginal teeth, T. trochiformis
has several.
DISCUSSION
According to the results presented here only two species
of the genus Trochita are living in the Southwest-
ern Atlantic waters: T. pileohis (d’Orbigny, 1841) and
T. pileus (Lamarck, 1822). Both species have a similar
distribution, however while T. pileus is quite common
and subtidal in the northern part of the distribution,
T. pileohis is somewhat rare and usually lives deeper.
Representatives of the genus could be found from
Patagonian deposits of probably Miocene age. Ihering
(1907: 148) already mentioned the presence ol several
species of Trochita included what he identified as
T. pileus from Cenozoic deposits. A complete revision of
the Neogene species of Trochita could show that the
same species are living since that age in Patagonia.
Contrary to other species of Trochita (i.e., T. dhofarensis
and T. trochiformis sensu Taylor and Smythe, 1985), the
Atlantic species are apparently not related with upwell-
ing areas.
ACKNOWLEDGMENTS
Thanks to A. Tablado, A. Sodor (MACN), Kathie Way
(NHMUK), Ives Finet (MHNG) for their assistance
in the revision of the material. Ivan Cariete (Punta
Arenas) sent good information about the Pacific Trochita.
The manuscript benefitted from critical reading from
S. Nielsen (Kiel) and an anonymous reviewer. We are
grateful to CONICET ol Argentina, ol which G.P. is a
researcher and D.G.U. a fellow. Part of the material was
collected during a research trip onboard of the B/I Puerto
Deseado. This contribution was partially supported by
the project PICT 942 from the Agenda Nacional de
Pro morion Cientifica y Tecnologica (Argentina).
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THE NAUTILUS 126(2):79-85, 2012
Page 79
Two Neogene vesicomyid speeies (Bivalvia) from Japan
and their biogeographic implications
Kazutaka Amano
Department of Geoscience
Joetsu University of Education
Joetsu 943-8512, JAPAN
amano@juen. ac.jp
Steffen Kiel
Georg-August Universitiit Gottingen
Geowissenschaftliches Zentrum, Abteilung Geobiologie
Goldschmidtstr. 3
37077 Gottingen, GERMANY
[email protected]
ABSTRACT
A new species of the bivalve family Vesicomyidae, Calijptogena
veneriformis, is described from the Pliocene part of the
Kurokura Formation in Niigata Prefecture, Japan. This species
belongs to die Plio-Pleistocene Omma-Manganji fauna on the
coast of die Japan Sea. We document previously unknown char-
acters of the shell interior of “Vesicomya” kawadai (Aolti) from
lower to middle Miocene deposits in Honshu, Japan, showing
that the species belongs to the genus Pliocardia. The genus
Pliocardia might have a longer geologic history than previously
appreciated. When the currently known distribution of
Pliocardia is taken at face value, die genus might have colonized
the Atlantic Ocean only very recently, perhaps as late as the
Pliocene, despite its long geologic history.
Additional keywords: Pliocardia , Vesicomya, Calyptogena,
Plio-Pleistocene
INTRODUCTION
The Vesicomyidae is one of the six liivalve families living
in symbiosis with chemoautotrophic bacteria, with a geo-
logic history ranging back to the middle Eocene (Taylor
and Glover, 2010; Kiel, 2010). Due to the limited num-
ber of shell characters and the high morphologic plastic-
ity among the vesieomyids, Japanese malacologists often
subdivided the family into two genera, Calyptogena Dali,
1891 for large elongate shells, and Vesicomya Dali, 1886
for small subcircular or veneriform shells (e.g. Sasaki
et ak, 2005). Recently, we have examined fossil repre-
sentatives of large, elongate vesieomyids in the north-
ern Pacific region (Amano and Kiel, 2007, 2010, 2011;
Kiel and Amano, 2010) and distinguished four genera:
Calyptogena, Archivesica Dali, 1908, Adulomya Kuroda,
1931, and Hubertschenckia Takeda, 1953.
The genus Calyptogena is composed of ten or more
Recent species (Krylova and Sahling, 2006, 2010). Among
them, only one species, Calyptogena pacifica Dali, 1891,
lias fossil representatives, ranging from the upper Mio-
cene to the middle Pleistocene in the Japan Sea Border-
land (Kanno et ak, 1989; Amano, 2003; Amano and
Kanno, 2005; Amano and Jenkins, 2011). We have now
recovered one new species of Calyptogena from the Japan
Sea borderland, from the lower Pliocene part of the
Kurokura Formation in Niigata Prefecture.
Among the small vesieomyids ("Vesicomya” of Japanese
malacologists), Cosel and Salas (2001) recognized several
genera, namely Vesicomya , Isorropodon Sturany, 1896,
Waisiuconcha Beets, 1942, and Callogonia Dali, 1889. In
addition, Krylova and Janssen (2006) and Krylova and
Sahling (2010) redefined the small vesicomyid genus
Pliocardia Woodring, 1925, and included two Japanese
species, Vesicomya crenulomarginata Okutani, Kojima,
and Iwasala, 2002 and V. kuroshimana Okutani, Fujikura
and Kojima, 2000. According to these taxonomic revisions,
the genus Vesicomya is now confined to very small species
(3-13 mm long) having subcircular shells and thin cardi-
nal teeth that are arranged roughly in a linear fashion.
Three fossil species were reported as “ Vesicomya ” from
Cretaceous and Neogene deposits in Japan: Vesicomya
kawadai (Aoki, 1954), V. inflata Kanie and Nishida, 2000,
and V ellipsoidea Kanie and Kuramoehi, 2001. Among
them, Amano et al. (2008) showed that V. inflata from
Cretaceous deposits in Hokkaido is a lueinid, not a
vesicomyid, and established the new genus Ezolucina for
diis species. “ Vesicomya ” ellipsoidea has a large and elon-
gate shell (up to 188.6 mm long) and two cardinal teeth
in right valve and does therefore belong to Archivesica
or Adulomya rather than Vesicomya sensu stricto (Amano
and Kiel, 2011). The generic status of Vesicomya kawadai
has so far remained uncertain.
The scope of the present contribution is (1) to
describe and name the new species of Calyptogena , (2)
to clarify the generic status of “ Vesicomya ” kawadai
based on newly collected material, and (3) to discuss the
biogeographic significance of these species.
MATERIALS AND METHODS
Several small vesicomyid specimens were recovered
from calcareous concretions within the mudstone of the
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THE NAUTILUS, Vol. 126, No. 2
lower Pliocene part of the Kurokura Formation at
Sugawa in Joetsu City, Niigata Prefecture (Figure 1,
Loc. 1), and they occur in association with Calyptogena
pacifica Dali, 1891 and the thyasirid bivalve Conchocele
bisecta (Conrad, 1849). These specimens are described
as Calyptogena veneriformis herein.
The specimens of “Vesicomya” kawaclai examined
herein are from the type locality of this species, in the
lower Miocene Honya Formation at Donosaku in Iwaki
City, Fukushima Prefecture (Figure 1, Loc. 2). The spec-
imens were found in a large calcareous concretion where
they co-occur with Adulomya chitanii Kanehara, 1937
and Conchocele bisecta. Some specimens of “Vesicomya”
kawaclai from the Honya Formation in the Ishimoriyama
area of Iwaki City, Fukushima Prefecture (Figure 1,
Loc. 3) were examined. These specimens were collected
by the late Prof. Katsumi Hirayama of Rikkyo University,
and the exact locality and the associated fauna are
unknown. In addition, two specimens of “ Vesicomya ”
kawaclai were collected on the Rekifune River (Figure 1,
Loc. 4), about 400 m downstream from the fossil whale-
fall site described by Amano et al. (2007), in eastern
Figure 1. Localities of the fossil vesicomyids described
herein. 1: Sugawa in Joetsu City, Niigata Pref. (lower Pliocene).
2: Donosaku in Iwaki City, Fukushima Pref. (lower Miocene);
3: Ishimoriyama area of Iwaki City, Fukushima Pref. (lower
Miocene). 4: Rekifune River, Hokkaido (middle Miocene).
5: Shimo-sasahara, Toyama Prefecture (uppermost lower to
lowermost middle Miocene). Calyptogena veneriformis was
collected from Loc. 1. Pliocardia kawaclai was recovered from
Locs. 2-5.
I lokkaido. The specimens were collected from carbona-
ceous mudstones of the middle Miocene Nupinai For-
mation, which most likely represent an ancient eold-seep
site, inferred from the associated species Adulomya
chitanii , Conchocele bisecta, and Portlandia sp.
We also reexamined the specimens of “ Vesicomya ”
kawaclai described by Amano et al. (2001) from the upper-
most lower to lowermost middle Miocene Higashibessho
Formation at Shimo-sasahara in Toyama Prefecture (Fig-
ure 1, Loc. 5). All specimens are deposited in the Joetsu
University of Education (JUE).
SYSTEM ATICS
Family Vesicomyidae Dali and Simpson, 1901
Subfamily Plioeardiinae Woodring, 1925
Genus Pliocardia Woodring, 1925
Type Species: Anomalocardia bowdeniana Dali, 1903
from the upper Pliocene Bowden Formation in Jamaica;
by original designation.
Remarks: According to the redefinition by Krylova
and Janssen (2006), this genus is characterized by its
small- to medium-sized elliptical shells having a shallow
radial depression from beak to postero-ventral margin, a
deep lunular incision, a shallow pallial sinus and a stout
ventral tooth (1) overlying the subumbonal cardinal
teeth (3a, 3b) in the right valve.
Pliocardia kawadai (Aoki, 1954)
(Figures 2-4, 6-7, 9-12)
Lamelliconcha kawaclai Aoki, 1954: 36-37, pi. 2, figs. 1 ,
10, 12-15, 22.
Vesicomya kawadai (Aoki). — Kamada, 1962: 88-89, pi. 8,
figs. 2a-b; Amano et al., 2001: 192, figs. 3-5, 8-11;
Amano et al., 2007: figs. 3D, E, G, [.
Description: Shell thin, medium sized (up to 40.7 mm
long), ovate, with radial depression extending from beak
to postero-ventral corner, with distinct, shallow lunular
incision. Pallial sinus shallow and v-shaped, situated just
before radial inner ridge extending from beak to poste-
rior corner. On hinge of right valve, ventral tooth (1) thin
to moderately thick, overlain by arched subumbonal
teeth (3a, 3b); cardinal tooth 3b bifid in some specimens.
Left valve hinge with thin cardinal tooth 2a and stout
cardinal tooth 2b, connected with a thin tooth 4b. Ante-
rior adductor muscle scar ovate with rather straight and
sharp posterior margin; anterior pedal retractor scar with
semi-circular shape, deeply impressed with rough stria-
tions subparallel to shell margin, located between hinge
plate and anterior adductor muscle scar, from which it is
separated by a sharp step. Posterior adductor muscle
scar ovate or circular and deeply depressed.
Material Examined: Twenty-seven specimens: JUE
nos. 15895, 15896, 15897.
K. Amano and S. Kiel, 2012
Page 81
Figures 2-12. Pliocardia species. 2-4, 6-7, 9-12. Pliocardia kawadai (Aoki). 2, 3. Right valve surface and hinge. Length
40.7 mm, hinge length 22.8 mm, JUE no. 15895-2; Loe. 3. 4. Anterior pedal retractor sear. Illustrated hinge length 12.7 mm;
aa, anterior adductor scar; ap, anterior pedal retractor scar; |UE no. 15697; Loe. 5. 6, 11. Right valve hinge and weak lunule ineisa.
Length 24.7 mm, JUE no. 15897-1; Loe. 4. 7. Inner surface of right valve and left valve hinge. Length 32.6mm, JUE no. 15895-3,
Loc. 3. 9. Escutcheon and lunule ineisa. Length 35.4 mm, JUE no. 15895-1; Loc. 3. 10. Right valve. Length 40.0 mm, JUE
no. 15896-1; Loc. 2. 12. Inner surface of left valve. Length 35.4 mm, JUE no. 15896-2; Loc. 2. 5, 8. Pliocardia sp. from an
Oligocene seep carbonate in the Lincoln Creek Formation in Washington State, USA (LACMIP loc. 17447B, see Amano and Kiel,
2007 for details). 5. Dorsal view. 8. view on right valve showing the posterior radial depression; length 16.0 mm.
Distribution: Lower Miocene Honya Formation in
Fukushima Prefecture; uppermost lower to lowermost
middle Miocene Higashibessho Formation in Toyama
Prefecture; middle Miocene Nupinai Formation in
Hokkaido (from both seep and whale-fall sites).
Remarks: As shown by Amano et al. (2001), spec-
imens from the Higashibessho locality (Loc. 5) usually
have a stouter ventral tooth in the right valve than the
specimens from the type locality of "Vesicomya" kawadai
in the Honya Formation. Based on the hinge dentition of
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THE NAUTILUS, Vol. 126, No. 2
the right valve, the presence of a lunular incision and the
presence of a small pallial sinus, “Vesicomya” kawadai is
herein transferred to the genus Pliocardia.
Comparisons: Pliocardia kawadai resembles the
Recent species P. crenulomarginata (Okutani, Kojima
and Iwasaki, 2002) in having a similar hinge dentition in
both valves, a radial depressed area from the beak to the
postero-ventral corner and a weak ridge before the pos-
terior adductor muscle scar. However, P. kawadai differs
from P. crenulomarginata by having a bifid 3b tooth, no
sharply bounded escutcheon and no fine crenulation on
the posterior margin.
Similar in outline to P. kawadai is a species from a
seep carbonate in the upper Oligocene part of the
Lincoln Creek Lormation in western Washington State,
USA (Amano and Kiel, 2007). This species is herein only
tentatively assigned to Pliocardia. We examined addi-
tional specimens from this locality (Ligures 5, 8) that
show a depressed area running from the beak to the
posterior corner and a distinct lunular incision. These
features, together with the previously described hinge
characters (Amano and Kiel, 2007), confirm the assign-
ment of this species to Pliocardia , although the presence
or absence o( a pallial sinus is unknown. This species is
much smaller (16.0 mm long) than the up to 40.7 mm
long P. kawadai.
Genus Calyptogena Dali, 1891
Cah/ptogena veneriformis new species
(Ligures 13-19)
Diagnosis: Small-sized Cah/ptogena with veneriform
shell, lunular incision lacking, pallial sinus lacking,
subumbonal pit lacking. Middle hinge tooth (1) thin,
surrounded by U-shaped connection of anterior (3a)
and posterior (3b) teeth in right valve, with posterior
nymph al ridge.
Description: Shell up to 19.8 mm in length, rather
thin, weakly inflated, triangular veneriform (height/
length — 0.73-0.88), equivalve, and inequilateral.
Antero-dorsal margin concave, continuing to rounded
anterior margin; postero-dorsal margin nearly straight
into oblique posterior margin at obtuse angle; ventral
margin broadly arcuate. Beak prominent, prosogyrate
and located at anterior one-third to two-fifth of shell
length (i.e., at 29-45% of shell length from anterior
margin). Nymph narrow and short. Lunule and lunular
incision absent. Escutcheon very narrow, demarcated
and deeply depressed. Surface ornamented with growth
lines only. Right valve hinge wide for size, with three
cardinal teeth, distinct posterior nymphal ridge and
subumbonal pit absent. Posterior tooth of right valve
(3b) large and triangular; anterior tooth (3a) short and
thin, parallel with dorsal margin and connecting with
posterior tooth (3b), forming U-shaped connection; mid-
dle tooth (1) long and thin, and surrounded by anterior
and posterior teeth. Middle cardinal tooth of left valve
(2b) stout, connecting with anteriorly oblique anterior
tooth (2a); posterior tooth (4b) thin. Pallial line entire.
Anterior adductor scar ovate; posterior adductor scar
pear-shaped.
Figures 13-19. Cah/ptogena veneriformis new species. All specimens are from the type locality (Loc. 1). 13-15, 18-19.
Paratypes. 13. Left valve hinge; hinge length 10.0 mm, fUE no. 15899-5. 14. Right valve hinge; hinge length 6.6 mm, JUE
no. 15899-4. 15. Right valve hinge; hinge length 10.5 mm, JUE no. 15899-3. 18. Dorsal view of right valve; length 19.1 mm, JUE
no. 15899-1. 19. Inner structure of right valve; length 12.2 mm, JUE no. 15899-2. 16, 17. Holotype. 16. Right valve. 17. Dorsal
hew of figure 16; length 11.4 mm, JUE no. 15898. pnr, posterior nymphal ridge; aa, anterior adductor scar; pa, posterior
adductor scar.
K. Amano and S. Kiel, 2012
Page 83
Holotype: Length, 11.4 mm, height, 9.3 mm, JUE
no.15898, right valve.
Paratypes: Length, 19.1 mm, height, 15.1 mm, [UE no.
15899-1, light valve; length, 12.2 mm, height, 10.7 mm,
}UE no. 15899-2, right valve; length, 6.2 mm, height,
5.1 mm, JUE no. 15899-6, left valve"
Type Locality: Outcrop at Sugawa, Yasuzuka-ku, Joetsn
City, Niigata Prefecture (37°03'41" N, 138°29'22" E).
Material Examined: Twenty-two specimens from the
typo locality (Loe. 1 herein).
Remarks: At its type locality Calyptogena veneriformis
is associated with Cahjptogena pacifica. A veneriform
shell shape is unknown among the Recent specimens of
C. pacifica , despite the wide range of variation in shell
moiphology among Recent Calyptogena (Krylova and
Sahling, 2006; Krylova and Janssen, 2006; Cosel and
Olu, 2009). Furthermore, the morphological variation of
fossil C. pacifica from the Japan Sea borderland shows
narrower range than the Recent one (see Kanno et al.,
1989; Amano, 2003; Amano and Kanno, 2005; Amano
and Jenkins, 2011). We are therefore confident that
Cahjptogena veneriformis represents a new species inde-
pendent from C. pacifica.
Comparisons: Some specimens of Calyptogena pacif-
ica have a veneriform shell similar to that of C.
veneriformis new species (e.g. Krylova and Sahling,
2006, figure 4, H-M). However, C. veneriformis can Ire
distinguished from those specimens of C. pacifica by its
smaller and triangular shell, and its pear-shaped poste-
rior adductor muscle scar. Calyptogena veneriformis is
also similar to veneriform specimens of Calyptogena
valdiviae (Thiele and Jaeckel, 1931) from the Gull of
Guinea as illustrated by Cosel and Olu (2009). However,
C. veneriformis differs from C. valdiviae by having a
less inflated and triangular shape. Superficially C.
veneriformis resembles Wareniconcha guinensis (Thiele
and Jaeckel, 1931) in its veneriform shell. However,
Wareniconcha guinensis has a narrower hinge plate than
Cahjptogena and a subumbonal pit, unlike Calyptogena .
Cahjptogena veneriformis resembles some species
currently assigned to Waisiuconclia Beets, 1942 (e.g.,
Cosel and Salas, 2001; Krylova and Janssen 2006;
Kryl ova and Sahling, 2010) in having a similar shell form
and left valve hinge dentition. But C. veneriformis can lie
distinguished from Waisiuconcha by its broad posterior
cardinal tooth (3b) and the U-shaped connection
between the posterior and anterior cardinal teeth in the
right valve, and by the lack of a lnnnlar incision. How-
ever, we urge caution about the currently used concept
of Waisiuconcha because the type species (W. alherdinae
Beets, 1942) is known from a single left valve only but
the taxonomically informative hinge characters among
vesicomyids are usually those of the right valve.
Distribution: Type locality only; lower Pliocene part
of the Kurokura Formation in Niigata Prefecture, Japan.
Etymology: Named after its veneriform shell morphology.
DISCUSSION
Krylova and Sahling (2010) identified nine living species
of Pliocardia , which are found in many basins all over the
world ocean (Krylova and Sahling 2010). In addition,
there are several undescribed or misidentified extant
species in museum collections that are also likely to
belong to Pliocardia (S. Kiel, personal observation). The
fossil history of Pliocardia is complicated and requires
further research. The Miocene Pliocardia kawadai is so
far the only fossil representative of this genus in Japan.
Apart from P. kawadai and the late Pliocene type species,
the only other fossil species assigned to Pliocardia is
an as-yet unnamed species from the Oligoeene Lincoln
Creek Formation in western Washington State, USA
(Amano and Kiel, 2007; see also Figures 5, 8). However,
several Paleogene taxa from the North Pacific realm that
we have previously identified as Archivesica (Amano
and Kiel, 2007; Kiel and Amano, 2010), as well as
“ Vesicomya ” tschudi and “Vesicomya" ramondi from the
Oligoeene of Peru (Olsson, 1931), show some marked
differences to A. gigas, the type species of Archivesica .
They are all considerably smaller than A. gigas , many
have a lunular incision, unlike A. gigas , and some lack a
pallial sinus, unlike A. gigas. At least some of these
Paleogene taxa may belong to Pliocardia , or to new gen-
era. This applies also to the oldest known vesicomyid,
“ Archivesica ” cf. tschudi , from the middle Eocene
Humptulips Formation western Washington State, USA
(Amano and Kiel, 2007).
11 the currently known distribution of fossil Pliocardia
is taken at face value, it appears that the genus colonized
the Atlantic Ocean only very recently, despite its long
geologic history. Whereas the genus is known from the
northeastern Pacific at least since the Oligoeene and
from the northwestern Pacific since the early Miocene,
the earliest Atlantic record is the late Pliocene type spe-
cies P. bowdeniana. There are several older seep deposits
in the Caribbean region, but Pliocardia or Pliocardia- like
shells were reported from none of them (Gill et al., 2005;
Kiel and Peekmann, 2007). On the eastern side of
the Atlantic Ocean, several Miocene seep deposits are
known from Italy, but again, they seem to lack Pliocardia
(Taviani, 1994; S. Kiel, personal observation). One likely
pathway for the colonization of the Atlantic Ocean by
Pliocardia is the Isthmus of Panama which closed in
the early Pliocene about 4. 0-3. 5 Ma (e.g. Collins, 1996).
A passage through this isthmus was recently suggested by
Martin and Goffredi (2011) who found that the closest
relative (based on molecular evidence) of “ Pliocardia "
krylovata from the Pacific side of Costa Rica is the
Caribbean species “Cahjptogena" ponderosa Boss, 1968.
Whereas Calyptogena pacifica is widely distributed in
late Miocene to midtile Pleistocene deposits of the Japan
Sea borderland (see Amano and Jenkins, 2011), the new
species Cahjptogena veneriformis is only known from
Page 84
TPIE NAUTILUS, Vol. 126, No. 2
lower Pliocene deposit in the central part of the Japan
Sea borderland. Thus the geologic age of Cah/ptogena
veneriformis coincides with the development of the
Omma-Manganji fauna (cf., Otuka, 1939; Amano, 2001,
2007). This fauna is endemic for the semi-enclosed Japan
Sea and developed after the separation of the Japan
Sea from the Pacific Ocean by uplifting of the backbone
mountain range in northeastern Honshu (see Ijima and
Tada, 1990). Cah/ptogena veneriformis may thus be con-
sidered as part of the Omma-Manganji fauna and may
have evolved from C. pacifica , the only other species of
Cahjptogena present in this basin during the Pliocene.
ACKNOWLEDGMENTS
We thank Ryuichi Majima (Yokohama National University)
and Yukito Kurihara (Mie University) for their help in
examining some fossil specimens. We also thank Richard
Squires (California State University) and Elena M. Krylova
(P.P. Shirshov Institue of Oceanology) for their reviews and
useful comments. This study was partly supported by a
Grant-in-aid for Scientific Research from the Japan Society
for Promotion of Science (C, 23540456, 2011-2013) to KA,
and by the Deutsche Forsehungsgemeinschaft through
grant Ki802/6-l to SK.
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Late Pleistocene Coins stimpsoni (Moreh, 1868)
(Gastropoda: Bueeinidae) from a seabed core
(2520 m) in the western North Atlantic
We recovered a shell of Coins stimpsoni (Moreh, 1868)
from a seabed piston core (8.5 cm diameter) taken at a
depth of 2520 m, approximately 15 km south of the most
seaward extension of Block Canyon, off Long Island, New
York, USA (39°28.00'N, 71°03.78'W). The shell (Figure 1)
was positioned 4.6 m below the core top (seafloor), and
was radiocarbon dated to 51,700 ± 6,500 ybp (Lab ID:
OS-83306), during the last glacial advance of the Pleisto-
cene. The biota preserved in cores typically consists of
meiofaunal elements, principally foraminiferan tests, and
ostracod carapaces. It is rare to find nearly complete shells
of larger macrofaunal species like C. stimpsoni because
low temperatures and high pressures at great depths
cause high carbonate dissolution rates. There is very little
fossil evidence of any kind for deep-sea macrofaunal taxa.
Coins stimpsoni is a common species in present-day
benthic communities ol the continental shell and upper
bathyal zone of the western North Atlantic. It ranges from
34.5° to 56° N, from 50° to 75° W, and from 16 to 1103 m
in depth (Rosenberg, 2009; and M.G. Harasewych, per-
sonal communication). The core specimen is from 1400 m
deeper than the modem bathymetric range limit, and is
much larger in size than gastropods currently li\4ng at
2500 nr. The deep sea is an energy-poor environment that
selects for small body size in mollnsks (McClain et ah,
2009). The size of the shell (Figure 1), which is broken, is
95.0 mm (height + width). Based on the shape of intact
living shells, we estimate the complete specimen to have
measured 136.4 mm (height + width). A statistical analy-
sis ol shell size, using quantile regression, performed on
an extensive collection of modern deep-sea gastropods
(3426 individuals of 83 species) from the western North
Atlantic, predicted a maximum size (height + widtir) of
only 28 mm at 2500 m (McClain et ah, 2009). A complete
specimen of C. stimpsoni would be five times this size
in terms of linear dimensions, and even larger in terms
of biovolume.
It is conceivable that ocean cooling and increased pro-
ductivity during glaciation (Yasuhara and Cronin, 2008)
could have permitted downward range extensions and
larger body size in deep-sea macrofaunal species. Bathy-
metric range shifts during glaciation are known for
meiofaunal species (Knrihara and Kennett, 1988; Cronin
and Raymo, 1997). Deep-sea ostracods increased in size
with global cooling during the Cenozoic (Hunt and Roy,
2006). In modern deep-sea gastropods in the western
North Atlantic, body size increases with depth across the
bathyal zone to around 3000-4000 m, and then decreases
at abyssal depdis (McClain et al., 2005). Since productiv-
ity in the form ol particulate organic carbon flux to the
Figure 1. Shell of Coins stimpsoni (Moreh, 1868) from a deep
seabed core taken at 2520 m south of Long Island, New York,
USA (core number OC463-19PC, 39°28.00' N, 71°03.78' W, IW
Oceanus Cruise 463, June 19, 2010). The specimen was located
4.6 m below the core top, and was radiocarbon dated to 51,700 ±
6,500 ybp. Shell height 62.8 mm, width 32.2 mm. Catalogue
number 374361, Department of Mollusks, Museum of Compar-
ative Zoology, Harvard University. Photograph by Leo Kenney.
seafloor decreases exponentially with depth (Rex et ah,
2006), body size and productivity are inversely correlated
in the bathyal zone. This suggests that increased produc-
tivity would not favor larger size at this depth, although it
is clearly associated with larger size in coastal species. It
is difficult to reconstruct paleoenvironments and their
effects on the deep-sea macrofauna in a precise way,
but a downward range shift of 1400 m and such a huge
increase in body size seem remarkable.
The Pleistocene was a period of active canyon forma-
tion and large-scale downslope transport of sediment
(Emery and Uelnipi, 1984). The area where the core was
M.A. Rex etal., 2012
Page 87
taken shows evidence of extensive debris flow (O’Leary,
1996; Chaytor et al., 2011). Several mechanisms that
could be responsible for the presence of Coins stimpsoni
in a seabed core at 2520 m are being investigated, includ-
ing downslope transport as part of a sediment-laden grav-
ity flow and deposition via ice-rafting. While the shell s
occurrence in the core is probably a spurious locality
record for the range of living populations, it does provide
an important temporal reference point. Coins stimpsoni
was a constituent of Pleistocene communities extending
back to at least 45,000 ybp. It survived the last glaciation,
which either must have obliterated much of its geographic
range through lower sea level and ice cover or shifted it
southward, as well as subsequent episodes of abrupt cli-
mate change that appear to have caused widespread
extinction of the upper bathyal deep-sea fauna (Yasuhara
et al., 2008).
ACKNOWLEDGMENTS
We thank M. G. Harasewych for confirming the iden-
tification of the shell, and Naval Health Clinic New
England for providing access to x-radiograph equipment
used to find the shell within the core. M.G. Harasewych
and B.A. Marshall read the manuscript. L. Kenney
photographed the specimen in Figure 1. This research
is supported by NSF grant OCE-1129612 to M.A. Rex,
S. Brault and C.T. Stuart. Funding for core collection and
radiocarbon dating by the USGS was supported by U.S.
Nuclear Regulatory Commission Project N6480: Physical
Study of Tsunami Source.
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Michael A. Rex
Department of Biolog)'
University of Massachusetts
100 Morrissey Blvd.
Boston, MA 02125
[email protected]
Jason D. Chaytor
U. S. Geological Survey
Woods Hole Coastal and Marine Science Center
384 Woods Hole Rd.
Woods Hole, MA 02543
Carol T. Stuart
Department of Biology
University' of Massachusetts
100 Morrissey Blvd.
Boston, MA 02125
Sponsored in part by the State of
Florida, Department of State,
Division of Cultural Affairs and the
Florida Council on Arts and Culture
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Volume 126 , Number 3
October 25, 2012
ISSN 0028-1344
CONTENTS
Corrigendum
117
THE NAUTILUS 126(3):89-97, 2012
Page 89
Detrital feeding in Xeniostoma inexpectans , new genus,
new species, and new subfamily Xeniostomatinae
of Calliostomatidae (Gastropoda: Vetigastropoda), liosted by
hexactinellid sponges of the Aleutian Islands, Alaska
James Hamilton McLean
Natural Histoiy Museum of Los Angeles County
900 Exposition Blvd.
Los Angeles C A 90007 U S A
[email protected]
ABSTRACT
The new genus Xeniostoma, type species X. inexpectans new spe-
cies, occurs from 200-400 m, off the western Aleutian Islands,
Alaska, in association with vase-shaped hexactinellid sponges of
the family Rossellidae. The feathery radula of Xeniostoma indi-
cates an allocation to the Calliostomatidae. It is assigned to the
new subfamily Xeniostomatinae because it lacks the massive
inner marginal teeth that function in the carnivorous feeding of
other genera in the family. There is no evidence of spongivory;
its feathery teeth are suitable for feeding on detritus that accu-
mulates on the host sponge.
Xeniostoma is also unusual in the family Calliostomatidae for
its shell with rounded whorls, lack of spiral sculpture, large
protoconch, and lack of teleoconch I, its lack of jaws, and lack
of the pseudoproboscis; it is also unusual in having a large right
suboptic tentacle. The very large protoconch suggests that only
a few large eggs are produced and that larvae can gain protec-
tion among the large projecting spicules on the outer surface of
the host sponge. Within the Calliostomatidae, Xeniostoma most
resembles the nearly smooth-shelled Magellanic genera Photinula
Adams and Adams, 1854 and Pliotinastoma Powell, 1951, which
have morphology of the lateral teeth in common and a similar
shell profile, but have the other characters expected in carnivo-
rous calliostomatids of the subfamily Calliostomatinae.
The typical subfamily Calliostomatinae exhibits an extreme
level of specialization that sets it apart from other trochiform
vetigastropods. If other living or fossil genera become
known, there would be further support for the hypothesis that
Xeniostomatinae could represent a surviving member of a
nearly extinct basal group within Calliostomatidae.
One other trochiform vetigastropod family, the smaller-shelled
Trochaelididae, is also hosted by deep-water hexactinellid sponges
of the family Rosellidae. A reconsideration of the feathery rad-
ula of Trochaelididae leads to the conclusion that it too is
suitable for detrital feeding.
Additional keywords: Calliostomatidae, Xeniostomatinae,
Trochaelididae, Rosellidae, spongivory, Hexactinellid sponges
INTRODUCTION
Benthic sampling for resource-monitoring purposes
has been conducted in recent years by the National
Marine Fisheries Service in the vicinity of the Aleutian
Islands, Alaska. New species of mollusks have been
recognized by the late Rae Baxter and by Roger
N. Clark; descriptions of further new species will
be forthcoming.
Among the new findings is a remarkable new mono-
typic genus Xeniostoma and new species inexpectans,
for which the smooth shell resembles the trochid
genus Margantes in the subfamily Margaritinae, and the
Magellanic genus Margarella in the subfamily Gibbulinae,
but radular characters of the new genus are indicative of
the family Calliostomatidae. Within the Calliostomatidae,
the closest comparison in shell profile is with the
Magellanic genera Photinula II. and A. Adams, 1854 and
Pliotinastoma Powell, 1951, which have nearly smooth
shells unlike those of most other calliostomatids. Compar-
isons are made in the descriptions of the new genus and
species to the type species Photinula coerulescens (King
and Broderip, 1831), from southern Argentina and Chile
(Figures 9-11) and Pliotinastoma taeniata (Wood, 1850),
from southern Argentina (Figures 12-13).
A number of questions about the affinity and its pos-
sible phylogenetic significance are raised by this dis-
covery. The taxonomic question requires resolution by
the placement in a new subfamily of Calliostomatidae.
Another question of major interest concerns the asso-
ciation of the new species with hexactinellid sponges.
This requires comparison with two other groups of
trochiform gastropods that have previously been reported
to occur on vase-shaped sponges of the family Rosellidae
in the sponge order Hexactinellida. No information about
this group of sponges has been provided in the accounts
of the gastropods; this omission is addressed here with
references to the literature on hexactinellid biology.
The best known gastropod family associated with
hexactinellid sponges is the small-shelled trochiform
vetigastropod family Trochaelididae Thiele, 1928, all
members of which are known to be associated with
hexactinellid sponges (Marshall, 1995b, and references
therein). In previous reports on the Trochaelididae, there
has been no information pertaining to the sponges.
Page 90
TPIE NAUTILUS, Vol. 126, No. 3
Another species of comparable size and morphology
identified by Gutt and Schicken (1998) as “Margarella
sp." [likely M. antarctica Lamy, 1905], is also associated
with hexactinellid sponges of the family Rosellidae in the
Antarctic fauna. The genus Margarella Thiele, 1893, has
most recently been assigned to the subfamily Gibbulinae
of the Trochidae by Zelaya (2004).
The new ealliostomatid genus Xeniostoma represents
the diird known vetigastropod group known to be associ-
ated with vase-shaped sponges of the order Hexactinellida.
Questions about the association of these gastropod
families with the hexactinellid sponges are raised: Are
these gastropods feeding directly on the sponges or are
they feeding on detritus that accumulates on the sponges?
What other benefits are derived from the association of
these gastropods with the sponges?
MATERIALS AND METHODS
The new species has come to light due to collecting
efforts of the late Rae Baxter and subsequently by Roger
N. Clark, both of whom have obtained specimens in the
process of salvaging invertebrate specimens from the
trawl catches, while aboard survey vessels during sum-
mer cruises at the Aleutian Islands.
Specimens were preserved directly in ethyl alcohol of
unknown strength, without relaxation or fixation. The
preservation of all material for the new species is not
adequate for thorough analysis. Preserved bodies of the
specimens have been twisted out of the shell, but bod-
ies of all specimens are torn, with the visceral mass not
remaining attached to the head-foot. The head-foot
material is suitable for study of the external anatomy
and the radula; the macerated remains of the viscera are
suitable for examination of gut contents, but not for
determining the condition of the gonads.
Although none of tire records of associated vetigastropods
are accompanied by identified sponges, some informa-
tion on the biology of the hexactinellid sponges is pro-
vided here. All three records (. Xeniostoma , one species of
Margarella, and all Trochaclididae) mention vase-shaped
sponges, which are here noted to be members of the
family Rossellidae. A substantial number of genera are
known in the Rosellidae, based on the morphology of the
siliceous spicules and the size and general moiphology of
the sponge (see Leys, et al. [1998], and Leys, et al. [2007]
for general review of hexactinellid biology).
Abbreviations: F/V, Commercial fishing vessel
contracted for survey cruises, by National Marine Fish-
eries Service; LACM, Natural History Museum of Los
Angeles County; USNM, United States National
Museum of Natural History, Washington; SEM, Scan-
ning electron microscopy.
SYSTEMATICS
Superorder Vetigastropoda Salvini-Plawen, 1980
Superfaniily Trocboidea Rafinesque, 1815
Family Calliostomatidae Thiele, 1924
Remarks: Hickman and McLean (1990) and Hickman
(1996) previously treated the group Calliostomatinae as a
subfamily of Trochidae, but other recent authors includ-
ing Marshall (1995a), Waren and Bouchet in Bouchet
and Rocroi (2005), Williams et al. (2010), have treated this
as a full family, as done so here. The most current and
comprehensive taxonomic treatment is that of Marshall
(1995a), who provided a complete list of generic level taxa
of Calliostomatidae. Marshall distinguished two sub-
families in which the sculpture consists of beaded spiral
cords, typical Calliostomatinae and Thysanodontinae
M arshall, 1985, members of which have a completely
different radula of slender teeth with backward directed
barbs. Thysanodontinae are not further mentioned here.
A third subfamily Xeniostomatinae is proposed here.
Subfamily Calliostomatinae Tbiele, 1924
Diagnosis: Spiral sculpture of raised cords, usually
strongly beaded; protoconch diameter usually relatively
small (diameter 320-420 pm); protoconch with raised hex-
agonal pattern and apertural rim; short teleoeoneh I of first
quarter whorl; pseudoproboscis well developed; jaws well-
developed; inner marginal teedi of radula enlarged.
Remarks: The typical subfamily was further divided into
two tribes by Marshall (1995a), the typical Calliostomatini
and Fautricini Marshall, 1995, the latter with shell char-
acters not unlike Calliostomatini, but considered by
Marshall to exhibit a basal, plesiomoiphic condition of
the radula for the family.
Subfamily Xeniostomatinae new subfamily
Diagnosis: Profile low, spiral sculpture lacking at all
growth stages; protoconch smooth, very large (900 pm);
teleoeoneh I lacking; pseudoproboscis lacking; jaws lack-
ing; inner marginals not enlarged.
Included Genera: Monotypic for Xeniostoma , new
genus. Other species or genera may await discovery.
Remarks: The lack of enlarged inner marginal teedi
distinguish this subfamily from typical members of the
subfamily Calliostomatinae. The lack of any spiral sculp-
ture, the lack of jaws, and the lack of the pseudoproboscis
are further differences from Calliostomatinae.
Xeniostoma new genus
Type Species: Xeniostoma inexpectans new species,
by original designation .
Description: Generic level description as in species
description below.
Remarks: The genus is monotypic; once additional
species become known, the limits of the genus will be
better understood; differences in size at maturity, shell
proportions, and shell sculpture can be expected.
J.H. McLean, 2012
Page 91
Xeniostoma inexpectans new species
Description: Shell (Figures 1-3): Color silvery white,
periostracum fully intact, shiny, thin, tightly adhering;
nacre of shell interior strongly iridescent. Shell diameter
about 10 nun, whorls 2.7. Whorls rounded but slightly
subangulate on shoulder, suture moderately impressed.
Surface smooth, axial and spiral sculpture lacking, peri-
stome nearly complete. Aperture large, height ol aper-
ture more than half height of shell. Umbilicus narrow,
umbilical wall rounded. Height 8.7 mm, diameter 9.4 mm
(holotype). Protoconch (Figure 3) smooth, bulbous, about
900 jam in maximum diameter, lip not thickened, transi-
tion from protoconch to teleoconch weakly indicated.
Operculum corneous, multispiral, about 7 evenly expanding
volutions, growing edge fitting the parietal area ol
the aperture.
External Anatomy (Figure 4): Head/foot with evenly
tapered suboptic tentacle below light cephalic tentacle
and anterior to short eyestalk, contracted length equiva-
lent to that of cephalic tentacle, diameter twice that ol
cephalic tentacles, not with apparent fold or groove. Ten-
tacle tip tapered. Cephalic lappets lacking, neck lobes of
simple narrow flaps. Eyes large, on short stalks, darkly
pigmented. Oral disk lacking pseudoproboscis. Anterior-
most pair of epipodial tentacles the largest, more poste-
rior pairs small (contracted in preserved specimens).
Jaws not detected. Ctenidium bipeetinate, free tip short.
Osphradium with simple lamellae.
Radula (Figures 5-6): Central field broad, with about
20 pairs of laterals, with inverted-V alignment of tooth
rows. Rachidian tooth with tapered tip and broad base.
Lateral teeth with broad bases to shafts, tips long and
narrow, extending toward rachidian tooth nearly across
central field, with deeply serrated edges away from distal
end. Extended tips emerging from inner side of tooth
base. Basal part of tooth at least four times broader
than emerging tip. Lateral edges of tooth bases thin and
Figures 1-4. Xeniostoma inexpectans new genus, new species. 1-2. Holotype, off Kiska Island, Rat Islands, Aleutian Islands,
Alaska, 272 m depth, maximum diameter of shell 10.0 mm. 1. Basal view. 2. Apertural view 3. Apical view ol immature paratype,
showing protoconch and first teleoconch whorl. Scale bar -- 1 mm. 4. Dorsal view of presented head-foot of holotype, mantle skirt
with gill attached (not showing) folded over to left. Abbreviations: LT, left cephalic tentacle; E, eye stalk; F, foot; M, mantle skirt; SO,
right suboptic tentacle; RT, right cephalic tentacle; S, snout. Scale bar = 1 mm.
Page 92
THE NAUTILUS, Vol. 126, No. 3
Figures 5-8. Radulae. 5-6. Xeniostoma inexpectans new genus, new species. 5. Full width of radular ribbon, showing rachidian
tooth at center, approximately 20 pairs of curved lateral teeth with broad bases, and numerous, closely adjacent, relatively straight
marginal teeth. Scale bar = 200 pm. 6. Enlarged view of rachidian and adjacent lateral teeth, showing that the rachidian tooth is only
slightly broader than adjacent lateral teeth. Scale bar = 100 pm. 7. Photinula coerulescens (King and Broderip, 1831), partial view of
ribbon with 4 complete rows of teeth, at center left showing elongate rachidian teeth with long tapering cusps, flanked by four pairs
of lateral teeth on left and right; on right showing large inner marginal teeth, and smaller outer marginal teeth (outmost marginals
had been stripped away). Scale bar = 200 pm. 8. Photinastoma taeniata (Wood, 1850), central field of radula, showing rachidian and
6 pairs of lateral teeth; on left and right showing enlarged inner marginal teeth and on right showing adjacent lateral teeth. Scale bar
] 1 ■" ) pm. [SEMs, 5-6, D. Geiger, 7. C. Hickman; 8, A. Waren],
flap-like, lacking interlock, concealing inner edge of adja-
cent lateral teeth. Marginal teeth numerous, with long
shafts and much shorter denticle bearing tips. Inner mar-
ginal teeth with short denticles, size of inner marginal
teeth not larger than other marginal teeth.
Holotype: LACM 2971 (ex LACM 97-162)
Type Locality: 272 nr, W of Kiska Island, Rat Islands,
Aleutian Islands, Alaska (51°54.3T N, 176° 35. 89' E), F/V
Dominator (sta. 23-971-202), R. N. Clark, 1 August 1997.
One live-collected specimen (Figure 1).
Paratypes: LACM 2972 (ex LACM 86-330), 219 m, NE
of Semisopochnoi Island, Rat Islands, Aleutian Islands
(52°24.09' N, 179° 42. 84' E), Rae Baxter, 10 September
1986; one live-collected specimen and two smaller,
dead specimens; maximum diameter 4.8 mm (used lor
initial radular preparation); one small shell coated for
SEM of protoconch (Figure 3); LACM 2973 (ex LACM
97-168), 325 m, SW of Buldir Island, Aleutian
Islands (52°18.50' N, 175°49.0' E), F/V Dominator
(sta. 23-971-243), R. N. Clark, 9 August 1997; 3 spec-
imens, 2 live-collected.
LACM 2974 (LACM 97-156), 384 m, SWof Amchitka
Island, Rat Islands, Aleutian Islands, Alaska (51° 27.70' N,
178° 35.0' E), F/V Dominator (sta. 23-971-181), R. N.
Clark, 27 July 1997. 2 live-collected specimens; LACM
3233, 221 m, NW of Kiska Island, Rat Islands, Aleutian
Islands, Alaska (52°9,58' N, 175° 11.67' E), F/V Gladiator
(sta. 147-04-195), R. N. Clark, 25 July 2004. 7 large spec-
imens (lof 3 LACM specimen used for radular SEM,
figures .5-6); one para type, USNM 1184071, and 3 paratypes
R. N. Clark Collection.
Distribution and Depth: Rat Islands, western Aleutian
Islands, Alaska, from Buldir Island toward die west, to
Semisopochnoi Island toward the east, from moderately
deep water, depths ranging from 219-384 m. An associ-
ation of the new species with vase-shaped hexactinellid
sponges of the family Rosellidae has been confirmed,
based on the eight additional specimens collected with
the sponge from Kiska Island, Aleutian Islands, during
the summer of 2004 by Roger Clark.
Etymology: The prefix xeno is Greek, xenos , a stranger,
suggested by the strange combination of characters in the
new genus. The root -iostoma is based on Calliostoma ,
J.H. McLean, 2012
Page 93
the nominate genus of the typical subfamily. The specific
name means unexpected.
Comparative Remarks: Shell. The smooth shell
resembles that of the trochid subfamily Margaritinae, par-
ticularly die Aleutian Island species Marga rites hickmanae
McLean, 1984, in its large size and silvery white colora-
tion, but differs in its complete lack of spiral sculpture and
its larger protoconch size. See McLean (1984) for illustra-
tion of M. hickmanae.
The smooth shell is unusual for Calliostomatidae; most
species of Calliostoma and related genera have a high
shell profile, often with flat sides. The sculptures of typ-
ical species consist of complexly beaded spiral cords.
Among the Calliostomatidae, the rounded whorls and
the smooth shell of Xeniostorna are comparable to the
genera Photimda (Figures 9-1 f), which has faint spiral
sculpture, and to Photinastoma (Figures 12-15), which
has more strongly indicated spiral cords in the early
teleoeonch (Figure 14, 15).
Protoconch. The protoconch at a diameter of 900 pm
lacks the typical hexagonal micro-sculpture of most
calliostomatids (see Hickman and McLean, 1990,
fig. TOD), and is larger than that known for most
troehoideans, in which the usual protoconch diameter
is about 200-300 pm (Hickman and McLean, 1990).
Loss of the hexagonal sculpture would be expected
with size increase of the protoconch. The protoconch
of Photinula coerulescens is shown here for compari-
son (Figure 9). It has a diameter of about 500 pm, lacks
the hexagonal sculpture and lacks the inflated lip edge.
The protoconch of Photinastoma is also large, but has
broad pits that diminish in size near the edge.
Teleoconch I. Marshall (1995: 385, figs. 2-3) indi-
cated that typical calliostomatines have a short segment
known as teleoconch !, followed by a growth scar on
the first quarter teleoconch whorl, which marks a change
to the mature sculpture. This is lacking altogether in
Xeniostorna (Figure 3), but very faintly indicated in
Photinula coerulescens (Figure 9), and in Photinastoma
taeniata (Figure 15).
Radula. The radula of Xeniostorna (Figures 5, 6) has
the rachidian and lateral teeth with a long, narrowly
constricted upper region of the shaft, with deep serra-
tion on the edges, having a strong resemblance to the
outer lateral teeth of the calliostomatid genus Photinula
(Figure 7) and Photinastoma (Figure 8), but having a
much narrower rachidian tooth and not the enlarged
inner marginals. The inner marginal teeth of Xeniostorna
have the appearance of calliostomatid marginal teeth at
the outer zone, but these inner marginal teeth are not
enlarged, as in Photinula and Photinastoma , and all other
calliostomatid radi ilae.
Jaw Plates. Jaw plates in calliostomatids are large,
dark brown, and readily observed in preserved spec-
imens through a dorsal cut made on the snout between
Figures 9-15. Shells and early whorls of Photinula and Photinastoma. 9-11. Photimda coerulescens (King and Broderip, 1831).
9-10. Mature shell (LACM 54671), intertidal, Posesion, Chile, height 20.0, diameter 27.9 mm. 9. Apertural view. 10. Basal Mew.
11. SEM view of apex of small specimen, showing smooth protoconch with pinched tip and lacking apertural rim (LACM 71-30),
75 m, SE of Tierra del Fuego, Chile. Scale bar = 200 pm. 12-15. Photinastoma taeniata (Wood, 1850), Mature shell (LACM 29116),
Punta Foca, Puerto Deseado, Argentina, height 9.3, diameter 11.6 mm. 12. Apertural view. 13. Basal Mew. 14-15. SEM views of
apical whorl and protoconch. 14. Apical whorl. Scale bar 1 mm. 15. Protoconch. Scale bar = 100 pm. [14-15, SEM by A. Waren],
Page 94
THE NAUTILUS, Vol. 126, No. 3
the cephalic tentacles (see Hickman and McLean, 1990,
fig. 68G). No jaws were found for Xeniostoma , but the
jaws of Photinula are large, as in other calliostomatids.
Pseudoproboscis. Xeniostoma lacks the calliostomatid
pseudoproboscis; a pseudoproboscis is present in Photinula.
The pseudoproboscis ol calliostomatids is a shallow
groove on the oral disk (see Hickman and McLean,
1990: fig. 70B).
Suboptic tentacle. Xeniostoma is the first known
calliostomatid genus to have a large right suboptic tentacle
in mature specimens, as shown in Ligure 4. All specimens
examined have this structure. This might be construed as
a cephalic penis (see Kano, 2008), but it is found on all
specimens, which would additionally require the assump-
tion that the species is hermaphroditic. The sex is not
known for any of the specimens. The material is not
sufficiently well preserved for me to establish a function
for the structure, and to establish that the gonad is her-
maphroditic. The most likely interpretation (A. Waren,
pers. comm.) is that it represents a hyper-development of
the anterior right “lateral appendage,” of the calliostomatid
larval stage, as illustrated by Ramon (1990: fig, 3).
DISCUSSION
Association of Troehiform Vetigastropods with
Hexaclinellid Sponges: Specimens of Xeniostoma ,
the trochid Margarella antarctica (as reported by Gutt
and Schicken, 1998), and all species in the family
Trochaelididae are associated with glass sponges of the
Class Hexactinellida, family Rosellidae (the deep-water
vase sponges). The association of Margarella and the
Trochaelididae are further discussed below.
The most recent general summary of hexactiuellid
biology is that of Levs et al. (2007). One species of rosellid
sponge, Rhabdocah/ptus dawsoni (Lambe 1892), has been
well studied because it accessible to divers off southern
Vancouver Island, British Columbia. Studies on the biol-
ogy of this species are those of Boyd (1981), and Leys and
Lauzon (1998), with further details provided by Leys et al.
(2007). This species lives at depths below 20 m on the
walls of fjords and sloping bottoms below walls of the
fjords. Long spicules project beyond the outer wall of
Rhabdocah/ptus , forming a dense veil of spicules that
harbors a microfauna of invertebrates (Boyd, 1981),
including small crustaceans, polychaetes, small bivalves,
and cerithiopsid gastropods, all of which live within the
spicule veil for protection from predators and benefit
from the flow of particles produced by the pumping
action of the sponge. The veil may become clogged with
detritus, but the sponge is capable of sloughing off the
outer layers of the mat, which is usually accomplished
in the winter months.
Another northeastern Pacific genus is Stauroeah/ptus,
which produces long spicules that project straight out
from the body of the sponge (see S. solidus Shulze, 1889,
as shown on the cover illustration of Green and Bakus,
1994); the long spicules of this genus also produce a micro-
habitat that provides protections for small invertebrates.
According to Reiswig (pers. comm.), the large atrial cavities
of rosellid sponges are kept free of sediment by the force
of the out-current flow generated by the sponge; these
cavities are utilized by larger invertebrates and fishes,
and are thus not suitable as a refuge for small gastropods.
Small-shelled vetigastropods of the families
Calliostomatidae ( Xeniostoma ), Troehidae ( Margarella ),
and Trochaelididae ( Trochaclis ) would be able to find
protection from predators within the outer veil of
rosellid sponges and would be provided with a contin-
uous flow of detritus. Species in each of these genera are
equipped with smooth shells devoid of projecting sculp-
ture, which would enable navigation through a veil or
thicket of long, external spicules.
Modifications of the Calliostomatid Radula and
Feeding in Xeniostoma: Marshall (1995: 386) noted:
“As showi by Waren (1990), the ealliostomatine radula is
distinctive among trochoideans in that the innermost pair
of marginals become greatly enlarged at an extremely
early stage of ontogenesis, after which the central and
lateral teeth arise by intercalation in the central field."
The large inner marginal teeth of calliostomatids are also
illustrated by Hickman and McLean (1990, figs. 71A, E).
It is these enlarged marginal teeth that are the working
teeth of calliostomatids, whether feeding on enidarians
or demosponges. Sponge feeding in the non-silicate
Demospongia has been infrequently reported in
Calliostomatidae (see Marshall, 1988: 228), but there is
no previous mention of any member of Calliostomatidae
feeding on hexactiuellid sponges.
The fact that the innermost marginal teeth of Xeniostoma
are not enlarged at any stage is a clear indication that this
is a radula otherwise unknown in the Calliostomatidae.
This indicates that Xeniostoma is not carnivorous, as
expected in all other calliostomatids. It seems most rea-
sonable that, because Xeniostoma does not have any
teeth in the radula that are robust enough to function in
sponge feeding, the only other option left for such a
feathery radula is to assist in feeding on detrital particles,
available within the veil of long spicules that project
away from the body of the rosellid sponge. There is a
source of detritus that settles on hard bottoms in sublit-
toral zones, in addition to pumping activity of the sponge
that would provide a source of detritus flowing toward
the outer body wall of the sponge.
Marshall (1995a) considered detritivory to be the mode
of feeding in the post-veliger larval stages of calliostomatids.
With regard to the termination of the teleoeonch I stage
at the end of the first quarter of teleoeonch growth,
Marshall (1995: 385) stated: “It seems clear that the
post-larval scar represents a growth pause of a crisis
period and I suggest that it may denote the transition
to exclusive detritivory (later transitional to carnivory).
It may actually mark the transition to carnivory in
some species, though the radula may be insufficient to
deal with Cnidaria or sponges at such an early stage of
development. This interpretation differs from that of
J.H. McLean, 2012
Page 95
Hickman (1992, fig. 5G), who identified the terminal
protoconch varix and the varix following it as denoting
the times of hatching and settlement respectively.”
Retention of this mode of feeding at maturity is there-
fore a simple hypothesis lor the feeding of Xeniostoma.
This is confirmed by my finding no sponge spicules in
the macerated visceral residue of any examined spec-
imens of Xeniostoma.
Egg Size and Protoconch Size in Xeniostoma: Males
of most species of Calliostomatidae broadcast their gam-
etes, which stimulates females to produce eggs, which are
laid in gelatinous strings, with fertilization taking place
externally (e.g., Ramon, 1990: 322). Larvae of many spe-
cies hatch and exit the egg strings as post-torsional veli-
gers (Hickman, 1992). The unusually large protoconch of
Xeniostoma is indicative of direct development from
large, yolky eggs. Relatively few eggs capable of forming
a protoconch at such a large diameter of 900 microns
could be produced. The veil of spicules on the outer wall
of the sponge would provide an ideal site for the early
stages of growth in Xeniostoma , in affording protection
as well as a steady source of detritus.
Loss of the usual micro-sculpture of calliostomatid
protoconchs is a function of increased size. Large
protoconchs are indicative of large egg size, direct devel-
opment, and reduced dispersal capabilities. Other large
protoconchs lacking the typical hexagonal microsculpture
are known in Calliostomatidae; in the deep-sea species
Falsimargarita nauduri Waren and Bouchet 2001 the
1 .2 mm diameter protoconch is smooth (Waren and
Bouchet, 2001: 133, fig. 8e).
Feeding in Margarella Antarctica: Gutt and Schiekan
(1998: 403) stated that the Antarctic species “Margarella
sp. L occurs on the glass sponges of the Rosella/
Scolymastra group, and that “ Margarella sp. 1 cleans but
also feeds on the sponges, thus having both a positive and
negative effect on the host.” I have not found a detailed
account of this relationship in papers cited therein, but
the radula of all species of Margarella has the rachidian
and the lateral teeth with strong cusps (see Zelaya, 2004),
so it is conceivable that the teeth of the central field
would be capable of feeding directly on the tissue of
the hexactinellid sponge, and the marginal teeth would
also clean the detritus that accumulates on the sponge.
Feeding in Troehaelididae: The radula of Acremodonta ,
a genus now assigned to the Troehaelididae, was illustrated
by Marshall (1983); that of Trocliaclis was first illus-
trated with SEM by Waren (1989). These genera have
finely divided tips to the numerous teeth, in which the
rachidian, laterals and marginals are of similar mor-
phology. The shell of Trocliaclis species (Figures 17,
10), is much smaller and of higher profile than that of
Xeniostoma. Here I illustrate the radula (Figure IS) of
a yet undescribed species of Trocliaclis, which shows
fully extended lateral teeth with feathery tips. This was
prepared by critical point drying of the whole radu-
lar ribbon, which provides an improvement over that
of most illustrations of the radula of Trocliaclis , in
which the most terminal divisions of the cusps are
often clumped together.
All records of trochaelidids indicate an association
with hexactinellid, vase-shaped sponges (see Marshall,
1995a), although none of the sponges have been identi-
fied in previous accounts. In a review of the species of
the southwestern Pacific in the vicinity of New Zealand,
M arshall (1995a) stated for the family that "gut contents
indicate spongivory.” In an effort to confirm this assump-
tion, I examined the macerated viscera of an unnamed
species of Trocliaclis collected from a vase sponge from
deep water in the Gulf of Alaska by the late Rae Baxter.
Figures 16-18. Family Troehaelididae, shell and radula of undescribed species of Trocliaclis from Gulf of Alaska. 16-17. Shell,
height 3.9 mm. 18. C ritical-point dried radula. Seale bar = 20 pm.
Page 96
THE NAUTILUS, Vol. 126, No. 3
I found only a few of the smaller spicules that comprise
the body in a rosellid sponge, rather than finding the
gut to be packed with sponge spicules. There should be
a few spicules that would be shed by the sponge if
Trochaclis is feeding on detritus, for which its radula is
suited. I therefore conclude that Trochaclis is also a
detrital feeder.
Implications for the Evolution and Higher
Classification of Calliostomatidae: Two genera from
southern South America, Photinula and Photinastonia , lack
the strongly beaded sculpture of typical ealliostomatids,
but have other features of typical ealliostomatids, includ-
ing jaws and the pseudoproboscis. Their protoconchs are
relatively large; that of Photinula (Figure 1 1) is smooth
and that of Photinastoma (Figure 15) has pits that sug-
gest a condition comparable to the hexagonal structure
of typical ealliostomatids with smaller protoconchs.
There is early spiral sculpture in the teleoconeh of
Photinastoma (Figure 14). These two genera are there-
fore retained in the subfamily Calliostomatinae.
Characters of the shell, anatomy, and radula of
Calliostoma and related genera are sufficiently unlike
those of all other trochoi deans to justify the family
level status currently accepted for Calliostomatidae.
On the shell alone, the species of Calliostoma in its
familiar sense are unlike all other trochoideans in their
intricately beaded sculpture and in having the short
phase of teleoconeh I, representing a suite of charac-
ters by which the shell, whether living or fossil, may
readily be distinguished from all other trochoideans.
These features are so elaborate that they are unlikely
to be basal in the family anti would have been derived
from something much simpler in shell sculpture (Mar-
shall, pers. comm.).
The nearly smooth shell of Xeniostoma may well rep-
resent a remnant of the basal or stem condition for the
family, with shells so unremarkable that there may be a
hidden fossil record with unrecognized roots that are
deep in geologic time. I am therefore proposing a new
subfamily Xeniostomatinae, which could prove to be the
basal subfamily for Calliostomatidae, if other genera for
living or extinct species are recognized in the future. The
other possibility is that all the derived character states of
Calliostomatidae have been lost in Xeniostoma .
It may be possible to get useful information from
molecular sequence information from Xeniostoma. If it
fits near other species, it is probably a loss of derived
characters. If it stands apart, there would be evidence
for a deeper division in the family.
ACKNOWLEDGMENTS
I am grateful to the late Rae Baxter, who first collected
the new species, and to Roger Clark for subsequent
material of Xeniostoma inexpectans. Daniel L. Geiger
(Santa Barbara Museum of Natural Histoiy) provided
SEM images of the radula of a mature specimen. Carole
Hickman (University of California, Berkeley), generously
provided the SEM image of a previously prepared
radula of Photinula coerulescens . Anders Waren (Swedish
Museum of Natural History) furnished images of the
early whorls of Photinastoma taeniata. Michelle Sehwengel
(Figures 1-4) and Brian Koehler (Figures 5-10) assisted
in the imaging tasks in the preparation of the illustra-
tions. For enlightenment about hexaetinellid biology
and references to the current literature 1 thank Henry
Reiswig (University of Victoria, British Columbia). I thank
Bruce Marshall (National Museum of New Zealand), for
helpful suggestions and for sharing his views about the
unknown stem group for Calliostomatidae. I thank Daniel
L. Geiger, Angel Valdes (formerly of LACM), and Anders
Waren (Swedish Museum of Natural History, Stockholm)
for helpful suggestions, and Carole Hickman and Bruce
Marshall, for thorough reviews of current and previous
versions of the manuscript.
LITERATURE CITED
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of gastropod families. Part 1. Nomenclator of Family-Group
Names (Bouehet and Rocroi). Part 2. Working Classifica-
tion of the Gastropoda (modem “archaeogastropods" by
Waren and Bouehet). Malacologia 47: 1-397.
Boyd, I. 1981. The spicule jungle of Rhabdocalyptus dawsoni : a
unique microhabitat. Unpublished senior honors thesis,
University of Victoria, Vancouver, 34 pp.
Green, K.D. and G.J. Bakus. 1994. Taxonomic Atlas of the
Benthic Fauna of the Santa Maria Basin and Western
Santa Barbara Channel, edited by J.A. Blake, A. L. Lissner,
and P.H. Scott. Vol. 2, The Porifera, 82 pp.
Gutt, f. and T. Schickan. 1998. Epibiotic relationships in the
Antarctic benthos. Antarctic Science 10: 398-405.
Hickman, C.S. 1992. Reproduction and development of
trochacean gastropods. The Veliger 35: 245-272.
Hickman, C.S. 1996. Phylogeny and patterns of evolutionary
radiation in trochoidean gastropods. In: J.D. Taylor (ed.)
Origin and Evolutionary Radiation of the Mollusca. Oxford
University Press, pp. 177-198.
Hickman, C.S. and J.H. McLean. 1990. Systematic revision
and supragenerie classification of trochacean gastropods.
Natural Histoiy Museum of Los Angeles County, Science
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Kano, Y. 2008. Vetigastropod phylogeny and a new concept
of Seguenzioidea: independent evolution of copulatory
organs in the deep-sea habitats. Zoologica Scripta 27:
1-21.
Leys, S.P. and N.R.J. Lauzon. 1998. Hexaetinellid sponge ecol-
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Trochidae (Mollusca: Gastropoda). Records of the National
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Marshall, B.A. 1988. Thysanodontinae: A new subfamily of
the Trochidae (Gastropoda). Journal of Molluscan Studies
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Marshall, B.A. 1995b. Recent and Tertiary Trochaelididae from
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THE NAUTILUS 126(3):98-104, 2012
Page 98
A new species of Felimare (formerly Mexichromis)
( Gastropoda: Opisthobranchia: Chromodorididae)
from the Yucatan Peninsula, Mexico
Deneb Ortigosa
UMDI-Sisal, Facultad de Ciencias
Universidad Nacional Autonoma de Mexico
Apartado Postal 70-153, C.P. 04510
Mexico, D.F., MEXICO
[email protected]
Angel Valdes
Department of Biological Sciences
California State Polytechnic University
3801 West Temple Avenue
Pomona, CA 91768 USA
[email protected]
ABSTRACT
A new species of chromodorid nudibranch from the western
Atlantic is described based on three specimens collected in the
Campeche Bank, Gulf of Mexico. This new species is assigned
to Felimare, an allocation based on a new classification of the
Chromodorididae established with molecular data. The anat-
omy and molecular data for this new species are compared
to those of other Atlantic species previously assigned to
Mexichromis (now part of Felimare). The new species is char-
acterized by having a primarily white dorsum with two longitu-
dinal blue lines, as well as a yellow band surrounding the
mantle margin. Mitochondrial 16S gene data confirms that the
new species is genetically different from other Atlantic species
of Felimare for which molecular data is available.
Additional ket/ words: Nudibranehia, Gulf of Mexico, 16S rntDNA,
anatomy
INTRODUCTION
Rudman (1984) defined Mexichromis Bertseh, 1977
(type species Chromodoris antonii Bertseh, 1976), as
well as other genera of Chromodorididae, based on
several anatomical characteristics, primarily the mor-
phology of the mantle glands, oral tube, jaw rodlets,
radular teeth, and reproductive system. According to
Rudman (1984), Mexichromis includes species with
mantle glands open ventrally and restricted to a few
large glands posteriorly and a few smaller ones along
each side; oral tube at least four times the length of the
buccal bulb and at least twice the diameter; jaw rodlets
ranging from bicuspid to multicuspid; radular teeth
bicuspid and denticulate, the denticles being as large or
nearly as large as the cusps; reproductive system with a
large ramifying vestibular gland covering the ventral
surface of the female gland mass and a large exoge-
nous sperm sac (bursa copulatrix) lying down the
wide muscular vagina. Although Mexichromis initially
included only eastern Pacific species (Bertseh, 1977),
Rudman (1983, 1984) transferred the tropical Indo-
Pacific species M. mariei (Crosse, 1872), M. festive
(Angas, 1864), M. macropa Rudman, 1983, and
M. miiltitubercidata (Baba, 1953) to diis genus. Since then,
four additional species have been added to this group
(Ortea et ah, 1996), including the western Atlantic spe-
cies M. kempfi (Ev. Marcus, 1971) and M. molloi Ortea
and Valdes, 1996, and the eastern Atlantic species
M. francoisae (Bouehet, 1980) andM. garciagomezi Ortea
and Valdes, 1996.
Recently, Johnson and Gosliner (2012), based on
molecular data, reorganized the classification of the
Chromodorididae and found that the traditional group
Mexichromis is paraphyletic. According to this new
scheme, the eastern Pacific and Caribbean species
M. porterae (Cockerell, 1901) and M. kempfi are trans-
ferred to Felimare Marcus and Marcus, 1967 along with
other eastern Pacific and Atlantic species previously
assigned to Hypselodoris Stimpson, 1955, whereas
Mexichromis is maintained for M. antonii, the type spe-
cies, as well as species previously assigned to Durvilledoris
Rudman, 1984 and Pectenodoris Rudman, 1984, and all
the tropical Indo-Pacific species of Mexichromis.
Although not explicitly tested in their phylogenetic
analysis, Johnson and Gosliner (2012) hypothesized
that most Atlantic species traditionally assigned to
Mexichromis likely belong to Felimare, including
M. francoisae, M. molloi, whereas eastern Pacific species
belong to Mexichromis, including M. tica Gosliner,
Ortea and Valdes, 2004 and M. turn (Marcus and
Marcus, 1967).
In this paper, a new species of Atlantic chromodorid
nudibranch is described based on specimens collected
in the Yucatan Peninsula, Mexico. The external mor-
phology and anatomy of this new species is consis-
tent with those of the group traditionally defined as
Mexichromis, but now considered Felimare (Johnson
and Gosliner 2012).
D. Ortigosa and A. Valdes, 2012
O
Page 99
MATERIALS AND METHODS
Collection and Preservation: The Campeche Bank
is composed of small reefs and cays located in the
northwestern sector of the Yucatan Peninsula. The bank
reaches 60 m depth (Spalding, 2004). The specimens
here studied were collected by hand in the Madagascar
Reef, within the Campeche Bank (Ortigosa-Gutierrez,
2009) as part of a multidisciplinary project aiming to
describe the diversity of the main groups of invertebrates
(corals, echinoderms, crabs, and shrimps) that inhabit
some reefs of the Campeche Bank. The Madagascar
Reef is located 40 km offshore and ranges in depth from
4 to 14 m (Zarco-Perello, 2008). All the specimens were
photographed alive, then relaxed with clove oil, and fixed
and preserved in absolute ethanol. The type material is
deposited in the collections of the Coleccion Nacional de
Malacologia, Instituto de Biologfa, Universidad Nacional
Autonoma de Mexico (CNMO) and the Natural History
Museum of Los Angeles County (LACM).
Morphological Examination: The specimens were
dissected and the reproductive system examined and
drawn using a dissecting microscope with camera lucida.
The buccal mass of one individual was removed and
dissolved in 10% sodium hydroxide, and the radula and
jaw examined using a scanning electron microscope
(SEM) Hitachi S-3000N at the LACM.
DNA Extraction: DNA extraction was performed
using either a hot Clielex’ protocol or the DNeasy®
Blood and Tissue Kit (Qiagen). Approximately 1-3 mg
of the foot was cut into fine pieces for extraction for both
protocols. For the Clielex™ extraction, the foot tissue was
rinsed and rehydrated using TO mLTE buffer (10 mM
Tris, 1 mM EDTA, pH 8.0) for 20 minutes. A 10% (w/v)
Chelex® 100 (100-200 mesh, sodium form, Bio-Rad)
solution was prepared using TE buffer. After rehydra-
tion, the tissue mixture was then centrifuged, 975.00 pL
of the supernatant was removed, and 175.00 pL ol the
Clielex®1 solution was added. Samples were then heated
in a 56°C water bath for 20 minutes, heated in a 100°C
heating block for 8 minutes, and the supernatant was
used for PCR. The DNeasy protocol supplied by the
manufacturer was followed, with some modifications.
The elution step was modified such that the first elution
was collected using 100.00 pL of Buffer AE and was
allowed to incubate at room temperature for 5 minutes
before centrifugation. In a new test tube, a second elu-
tion step was conducted using 200.00 pL of Buffer AE
and was also allowed to incubate at room temperature
for 5 minutes before centrifugation. The first elution was
used for PCR.
PCR Amplification and Sequencing: Pahnnbi s uni-
versal 16S primers (Pahnnbi, 1996), as well as internal
primers for 16S designed for another group of opistho-
branchs (Ornelas-Gatdula et ah, 2011) were used to
amplify the regions of interest. Multiple attempts to
obtain CO I or complete 16S sequences using different
primers were unsuccessful.
The master mix was prepared using 34.75 pL IDO,
5.00 pL Buffer B (ExACTGene, Fisher Scientific),
5.00 pL 25 mM MgCL, 1.00 pL 40mM dNTPs, 1.00 pL
lOmM primer 1, 1.00 pL primer 2, 0.25 pL 5 mg/mL
Taq, and 2.00 pE extracted DNA. Reaction conditions
were as follows: an initial denaturation for 2 min at
94°C, 35 cycles of (1) denaturation for 30 sec at 94°C,
(2) annealing for 30 sec at 50°C, and (3) elongation for
I min at 72°C, and a final elongation for 7 min at 72°C.
PCR product yielding a band of appropriate size, each
approximately 250 bp in length, for I6S (16Sar-L +
1 6Sbr-FAP) was purified using the Montage PCR Cleanup
Kit (Millipore). Cleaned PCR samples were quantified
using a NanoDrop 1000 Spectrophotometer (Thermo
Scientific). Each primer was diluted to 2.0 pmol/pL for
sequencing with the PCR products. PCR products were
diluted to 6.0 ng/pL. Samples were sequenced at the
City of Hope DNA Sequencing Laboratoiy (Duarte,
CA) using chemistry types BigDye VI. I .
Phylogenetic Analyses: Sequences were assembled
and edited using Geneious Pro 4.7.4 (Biomatters Ltd.).
Geneious was also used to extract the consensus sequence
between the primer regions and to construct the align-
ment for each gene using the default parameters. A total
ol 250 bp were amplified from the new species and used
for the phylogenetic analyses (GenBank accession number
JX101321). For comparison purposes, several GenBank
sequences belonging to Mexichromis (as defined by
Johnson and Gosliner, 2012) were downloaded from
GenBank: Mexichromis antonii (EU982800), Mexichromis
macropa (EF534050), Mexichromis mariei (EF534049),
Mexichromis festive (EF534051), Mexichromis aurora
(EU982805), Mexichromis trilineata (EU982806),
Mexichromis lemniscata (EU982790), and Mexichromis
similaris (EF534055), and so were sequences belong-
ing to Felimare (as defined by Johnson and Gosliner,
2012): Felimare orsinii (AJ225189), Felimare villafranca
(AF249237), Felimare bilineata (EF534052), Felimare
califomiensis (EU982796), Felimare picta (AF249238),
Felimare picta verdensis (HM 162594), Felimare ruthae
(EU982799), Felimare kempfi (EF534047), Felimare
porterae (EF534067). Hi/pselodoris infucata (FJ917426)
was selected as the outgroup.
Tl re Akaike Information Criterion (Akaike 1974) was
executed in MrModeltest v2.3 (Nylander 2004), to deter-
mine the best-fit model of evolution. MrModeltest
selected GTR+I-f-G as the best-fit evolutionary model
for the data set and estimated the following parameters:
Base frequencies (A = 0.3623, C = 0.1202, G = 0.1679, T =
0,3496); Rate matrix ([A-C] = 0.9228, [A-G] = 9.0170,
[A-T] = 3.1 131, [C-G] = 0.1367, [C-T] = 9.2412); Pro-
portion of invariable sites = 0.3260; Gamma distribution
shape parameter = 0.5376. The resulting MrBayes block
model line was: lset nst=6 rates = invgamma.
A Bayesian analysis was executed in MrBayes v3.1.2
(Huelsenbeck and Ronquist 2001). The Markov chain
Page 100
THE NAUTILUS, Vol. 126, No. 3
Hypselodoris infucata
— Hypselodoris picta
0.7
■ Hypselodoirs villafrartca
0.68
0.71
0.95
Hypseledoris orsinii
— Hypselodoris bilineata
Hypselodoris californiensis
Hypselodoris ruthae
0.87
Hypselodoris picta verdensis
Mexichromis porterae
— Mexichromis kempfi
0.72
Felimare sisalensis
Pectenodoris trilineata
Pectenodoris aurora
Felimare
0.8
Mexichromis festiva
Mexichromis macropa
• Mexichromis mariei
- Durvilledoris lemniscata
Durvilledoris similaris
Mexichromis antonii
Mexichromis
0.05
Figure 1. Bayesian tree of 16S sequences. Posterior probabilities are shown only for nodes with values over 0.5. Binominal
combinations are the same as used in GenBank, reflecting the established nomenclature before Johnson and Gosliner (2012).
Monte Carlo analysis was run with two runs of six chains
for ten million generations, with sampling every 100 gen-
erations. All other settings remained in the default. The
default 25% burn-in was applied before constructing
majority-rule consensus tree. The remaining 150,000 trees
were used to construct majority rule consensus trees and
calculate posterior probabilities. All clades and support
values are shown in the resulting phytogenies. All posterior
probabilities are mapped on all trees.
RESULTS
Molecular Data (Figure 1): The Bayesian consensus
phylogram (Figure 1) is consistent with the results
obtained by Johnson and Gosliner (2012) despite the fact
that only one gene was analyzed, although it lacks sup-
port in several areas. This phylogeny confirms that
Felimare is monophyletic (posterior probability = 1.0)
and includes Atlantic species previously included in
Hypselodoris and Mexichromis as well as F. porterae.
Within Felimare, species with bicuspid radular teeth
(previously classified as Hypselodoris) are also mono-
phyletic (posterior probability = 1.0), whereas eastern
Pacific and Atlantic species previously classified as
Mexichromis are clustered in a poorly supported clade.
The new species described herein is in this group, con-
firming the morphological hypothesis of classification
proposed in this study.
The rest of the species included in the analysis (for-
merly Pectenodoris , Durvilledoris and Indo-Pacific and
one eastern Pacific Mexichromis ) are not supported as
monophyletic, but both sample size and gene coverage
are too limited to reach any conclusion about the classi-
fication of larger groups of the Chromodorididae.
This analysis confirms the position of the new spe-
cies here described in Felimare. It also shows that
this species appears to be sister to F. kempfi (posterior
probability = 1.0), but these two species are genetically
distinct in the 16S gene. In the short fragment obtained
(250 bp) there were 96.6% of identical sites Jretween
the new species and F. kempfi which is equivalent to
level of sequence similarity between other sister spe-
cies of Chromodorididae; in the same fragment there
were 97.0% of identical sites between M. aurora and
M. trilineata.
D. Ortigosa and A. Valdes, 2012
Page 101
Figure 2. Photographs ol live animals. A. Dorsal view of a
paratype of Felimare sisalensis sp. nov., 11 mm long (LACM
3223), B. Dorsal view of the holotype of Felimare sisalensis,
12 mm long (CNMO 3037). C. Dorsal view of a juvenile
specimen of Felimare kempfi from Campeche Bank, 14 mm
long. D. Lateral view of the mantle margin of a juvenile
specimen of F. kempfi (7 mm long) showing the characteristic
black spots.
Morphological Data (Figures 2A-B, 3, 4): The
examination of the morphological data conformed that
the material here examined represents a new species.
The following sections contain the formal description.
SYSTEMATICS
Chroniodorididae Bergh, 1981
Felimare
Felimare sisalensis new species (Figures 2 A B, 3, 4)
External Morphology (Figures 2A-B): The body is
opaque white with two blue lines that run from each
rhinophore to the gill, surrounding it. The mantle edge
is surrounded by a yellow band, followed by a thinner
blue band. The branchial leaves and rhinophores are blue
and retract into pockets. There are nine unnipinnate
branchial leaves. The loot sole is opaque white and is
surrounded by a blue line. The mantle margin contains
one band of small, rounded mantle glands, except for the
anterior end of the body.
Reproductive system (Figure 3): The deferent duct
is wide and short, followed by a long and narrow pros-
tate. The vagina is short and narrow. The uterine duct
connects into the vagina. The bursa copulatrix is large
and oval in shape. The seminal receptacle is somewhat
elongated and connects into the vagina, at the base of
dd
Figure 3. Reproductive system of a paratype of F. sisalensis
(CNMO 2981). Abbreviations: am: ampulla; be: bursa
copulatrix; dd: deferent duct; fg: female gland complex; pi-:
prostate; sr: seminal receptacle; ud: uterine duct; va: vagina;
vg: vestibular gland.
the bursa copulatrix. The ampulla is relatively short, and
straight, with no visible folds. The female gland complex
is large, almost the same size that the rest of the repro-
ductive systems. A vestibular gland was not observed,
but due to the small size ol the reproductive systems it
possible that it was overlooked.
Radula and Jaw (Figure 4): The radular formula is
38 x 20.0.20 in a paratype (CNM029S1). There are no
raehidian teeth. All lateral teeth are similar in shape and
size, with no clear distinction between inner, mid and
outer lateral teeth. All teeth are hook-shaped, elongate,
with 4-6 denticles. The jaw consists of numerous short,
tricuspid rodlets.
Type Material: HOLOTYPE: 12 mm long (CNMO 3037).
PARATYPE S: I specimen 11 mm long (LACM 3223) and
1 specimen 12 mm long (CNMO 2981).
Type Locality: Madagascar Reef, Campeche Bank,
Yucatan, Mexico (21°26'28.3" N, 90°17'22.8" W), 4 Sep-
tember 2007, 7 m depth. All specimens were collected
on green algae.
Geographic Distribution: This species lias only
been collected on Madagascar Reef, Campeche Bank,
Yucatan, Mexico.
Etymology: The species is named in honor of the
village of Sisal, Yucatan, Mexico, were the Umdi-Sisal,
UNAM station is located.
DISCUSSION
Among the Atlantic species of Felimare, only four (those
previously assigned to Mexichromis ) have radular teeth
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THE NAUTILUS, Vol. 126, No. 3
Figure 4. Scanning electron micrographs of a paratype of F. sisalensis (CNMO 2981). A. Outer teeth. B. |aw rodlets.
with all denticles similar in size (non bicuspid) and short
jaw rodlets with more than two cusps. These species
are Felimare kempfi , F. molloi , F. garciagomezi , and
F. francoisae. Additionally, molecular data confirms a
closer relationship between these species versus species
of Felimare with bicuspid teeth. Thus, the new species
F. sisalensis , which has a similar radular and jaw mor-
phology, is compared only to Atlantic species of Felimare
with non-bicuspid teeth.
All Felimare species previously assigned to Mexichromis
are very uncommon, with no more than eight specimens
found per survey even in the case of F. kempfi (Meyer,
1977), which is the most common species. Most of these
Felimare species have restricted geographic ranges.
Felimare francoisae has been reported only for Cape
Verde and Senegal (Bouchet and Ortea, 1980; Ortea
et ah, 1996). The material from Ghana assigned to
Hypselodoris tricolor by Edmunds ( 1981 ) was described as
the new species M. garciagomezi by Ortea et al. (1996),
because of its distinct external coloration, radula and
jaw morphology. This species has not been collected
ever since Edmunds’s (1981) record. Felimare molloi
Table 1. Character comparison of Atlantic species of Felimare traditionally assigned to Mexichromis.
D. Ortigosa and A. Valdes, 2012
Page 103
F. francoisae
F. garciagomezi
F. molloi
F. sisalensis
F. kempfi
Figure 5. Schematic representation of the color pattern of the Atlantic species of Felimare previously assigned to Mexichromis.
was originally described based on a single specimen
collected in Isla Picuda, Venezuela (Ortea et ah, 1996)
and never cited again. Felimare kempfi is the only spe-
cies with a relatively broad geographic range including
Florida, Mexico, Panama, Costa Rica, Puerto Rico, and
Brazil and also in two reefs of the Campeche bank
(Table 1). All these species seem to live in shallow
waters; the deepest record is for F. kempfi in Brazil,
at 37 m depth (Marcus, 1971).
Felimare sisalensis can be distinguished easily from
those other four species by its external coloration
(Figure 5). Felimare kempfi is a primarily blue species
with a longitudinal white band that runs from between
the rhinophores to the gill and a series of elongate black
spots on each side of this band, whereas in F. sisalensis is
mainly white with two blue lines that run from each
rhinophore to the gill. Ortigosa-Gutierrez (2009) reported
specimens of F. kempfi collected from another reef
of the Campeche Bank and similar in size to the type
material of F. sisalensis (7 and 12 mm long), and the
black spots are visible (Figure 2C-D). Valdes et al.
(2006) suggested the possibility that F. molloi could be
a synonym of F. kempfi , which would lack black spots
as juveniles, but Ortigosa-Gutierrez’s (2009) record sug-
gests that F. molloi is indeed a distinct species. Felimare
molloi has also a white band that runs from between the
rhinophores to the gill and instead of the black spots of
F. kempfi , it has a blue band surrounded by irregular
areas of white color. Felimare garciagomezi is almost
completely blue, with a central white line that runs
from the midi lie of the rhinophores to the gill and two
shorter pale blue lines on each side ol the central line
that do not surround the rhinophores.
The radular and jaw morphology of F kempfi and
F. sisalensis are veiy similar, both species lack rachidian
teeth, and the outer radular teeth have 6 cusps; the jaw
rodlets have 3-4 cusps in both species (Table 1). The
radula of the eastern Atlantic species F. francoisae is
different from that of F sisalensis as it has more cusps
on the outer radular teeth (10-11) and the jaw rodlets
have 3-6 cusps instead of 3-4 in F sisalensis. Finally
F. garciagomezi has finely denticulate outer radular teeth
(Edmunds, 1981) and jaw rodlets with 3-4 cusps (Table 1).
The only two species for which the reproductive sys-
tem has been described and illustrated are F francoisae
(in Bouchet and Ortea, 1980; Ortea et al., 1996) and
F. molloi (in Ortea et al., 1996). The reproductive
system of F. sisalensis is clearly distinguishable from
that of F. francoisae and F molloi because the semi-
nal receptacle is not connected directly into the bursa
copulatrix as in these two species. Additionally, in
F. sisalensis the deferent duct is wider than that of
F francoisae. A vestibular gland was not observed
in F. sisalensis , but it was reported in F. francoisae
and F. molloi by Ortea et al. (1996).
ACKNOWLEDGMENTS
CONACyT (Mexico) provided a Master Scholarships
through the PCMyL, UNAM (202617) to the senior
author. The Ecology research group of UMDI-Sisal,
UNAM supported the fieldwork. The SEM work was
conducted at the Natural Histoiy Museum of Los Angeles
County. Jermaine Malmib, Jennifer Alexander and Dieta
Hanson helped with DNA extraction, amplification and
sequencing. This work was funded partially by the UNAM
(PAPIMe" PE207210) and CONACyt-SEMARNAT
(Project # 108285) and the Programa de Biodiversidad
Marina de Yucatan (BDMY). Vinicius Padula and
Rebecca Johnson provided valuable comments that
greatly improved the quality of this paper.
Page 104
THE NAUTILUS, Vol. 126, No. 3
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THE NAUTILUS 126(3): 105-112, 2012
Page 105
A new genus and new species of freshwater mussel from the
mid Late Triassic rift lakes of eastern North Carolina
(Bivalvia: Unionida: cf. Unionidae)
Arthur E. Bogan
Patricia G. Weaver
North Carolina State Museum of Natural Sciences
11 West Jones St.
Raleigh, NC 27601-1029 USA
[email protected]
ABSTRACT
In eastern North America, surface exposures of Triassic
basins extend from Nova Scotia southwestward to South
Carolina. This interrupted series of half-grabens resulted
from early Mesozoic rifting of the supercontinent Pangaea.
In south-central North Carolina, the Deep River basin is
comprised of the Durham subbasin, the Colon cross-structure,
the Sandford subbasin, the Pekin cross structure, and the
Wadesboro subbasin. Deposits within the Durham subbasin
are recognized as the Chatham Group, part of the Newark
Supergroup and form part of a series informally designated
as Lithofaeies Association II. These strata are considered
to be alternating fluvial and lacustrine sediments. Though
research on the vertebrate fauna from this lithofaeies has
been ongoing for over a century, considerably less research,
particularly in North Carolina, has been done on the lacus-
trine invertebrate fauna.
Ongoing field work at a brick-clay quarry in tire village of
Genlee, Durham County, North Carolina, has yielded three
distinct forms of Triassic freshwater bivalves, one is considered
here to be of the order Unionoida and the others tentatively
a mytiloid in shell outline and a sphaeriid in shell outline.
The unionoid specimens are assigned to Triaslacus new genus
and Triaslacus carolinesis new species and are tentatively
assigned to the Unionidae. These new specimens are compared
with specimens described as belonging to the unionoid fami-
lies Unionidae, Hyriidae, and Mulleriidae [ + Myeetopodidae]
from Massachusetts and Pennsylvania. However, none of the
North Carolina specimens exhibit the umbonal sculpture
exhibited by the northeastern specimens. The North Carolina
unionoid specimens lack any evidence of hinge development
or umbonal sculpture.
Non-unionoid bivalves are quite rare in these deposits.
The bivalve fauna is found in association with ostracods of
the genus Darwinula; clam shrimp, Euestheria, represented
by carbonized impressions of the shells; and fish and plant
remains. This freshwater environment is preserved in a mud-
stone or clayey siltstone sediments.
Additional keywords: Unionida, Uniomorphi, Newark Supergroup
INTRODUCTION
Freshwater bivalves in the Order Unionida have been
placed in the Cohort Uniomorphi (=Paleoheterodonta),
based on shell characters used to identify the sister rela-
tionship of Trigonida and Unionida (Weir, 1969; Bieler
et al., 2010; Carter et al., 2011). Trigonida and Unionida
are considered sister taxa in the Uniomorphi based ini-
tially on shell morphology and the presence of nacre.
This relationship is supported by DNA analyses (Hoeh
et ah, 1998; Hoeh et ah, 2001; Giribet and Wheeler,
2002; Taylor et ah, 2007).
Subdivision of the Unionida lias varied over the past
hundred years from a single family to six modern fami-
lies and various included fossil groups (e.g., Haas, 1969;
Starobogatov, 1970; Carter et ah, 2011). Taxonomic place-
ment of fossil freshwater bivalves in modern extant fami-
lies and genera is not supported. This practice is fraught
with problems of convergence of shell morphology
(Ortmann, 1912; Prashad, 1931; Hartman and Bogan,
2009). Allocations of bivalve taxa discussed here are
based on shell shape and morphology and are considr
ered tentative until there is a phylogenetic analysis of
the modern and fossil Unionida.
History of North American bivalve fossils assigned
to the Order Unionida was summarized by Henderson
(1935) and more recently reviewed by Watters (2001).
Triassic freshwater bivalves described from the eastern
United States are primarily from Pennsylvania and
Massachusetts (Table 1). North American Triassic fresh-
water bivalves described from the southwestern United
States including Texas, New Mexico, Arizona, California,
and Utah, are assigned to Unionidae and Hyriidae
(Good, 1998).
Pilsbry (1921, 1926) placed the Triassic freshwater
bivalve taxa from the northeastern United States in
the modern South American families Mycetopodidae
based on similarity of shell shape to the modern genus
Page 106
THE NAUTILUS, Vol. 126, No. 3
Table 1. List of freshwater bivalves from the Triassic of the
eastern United States.
Mulleriidae (+Mycetopodidae)
Mycepoda? diliculi Pilsbry in Wanner, 1921, Pennsylvania
Ilyriidae
Diplodon borealis Pilsbry in Wanner, 1921; Pilsbry,
1926, Pennsylvania
Diplodon carolus-simpsoni Pilsbry in Wanner,
1921, Pennsylvania
Diplodon pennsylvanicus Pilsbry in Wanner,
1921 . Pennsylvania
Diplodon wanneri Pilsbry in Wanner, 1921, Pennsylvania
Diplodon yorkensis Pilsbry, in Wanner, 1921 Richards
1944, Pennsylvania
Unionidae
Unio emersoni Troxell, 1914, Massachusetts
Anoplopliora wilbrahamensis Emerson, 1900, Massachusetts
[moved to Unio by Lull, 1915]
Unionidae sp. Bain and Harvey, 1977; Good, 1995a; Good,
1995b, North Carolina
Trace fossils (Lull, 1915, as Mollusks?; Henderson, 1935,
as freshwater mussel trails).
Bisulcus undulatus Hitchcock, 1865, Massachusetts
Trisulcus laqueatus Hitchcock, 1S65, Massachusetts
Mycetopoda, and Ilyriidae based on radial umbo sculpture,
similar to the modern genus Diplodon. Remaining taxa
were described or moved to the genus Unio and assumed
to belong in the family Unionidae.
Two trace fossils described by Hitchcock (1865)
Bisulcus undulatus Hitchcock, 1865 and Trisulcus
laqueatus Hitchcock, 1865, both from Massachusetts,
were described as annelids. Trisulcus was listed as a
worm track by Lesley (1890), who failed to list Bisulcus.
Lull (1915, 1917) placed them in Mollusca? noting they
differ from worms in being multiple-, double-, or triple-
ridged trails. Both of these taxa were listed as possible
tracks of freshwater mussels but with no further elabo-
ration by Henderson (1935). There is some doubt that
these represent tracks of freshwater mussels and not
some other invertebrate.
GEOLOGICAL SETTING
The Deep River Basin, located in east-central North
Carolina, resulted from early Mesozoic rifting of the
supercontinent Pangaea (Clark et ah, 2001). This north
to northeast-trending half graben is about 226 km long
and averages about 16 km in width (Bain and Harvey,
1977). Deposits of the Deep River Basin consist of varying
amounts of conglomerate, sandstone, siltstone, claystone,
shale, coal, and small amounts of limestone and chert
(Clark et ah, 2001). These deposits are recognized as the
Chatham Group, part of the Newark Supergroup as
defined by Olsen (1978) and Luttrell (1989). The Deep
River Basin is bordered on the east by the Jonesboro fault,
a west-dipping, high-angle normal fault (Campbell and
Kimball, 1923). The Jonesboro fault separates Triassic
sedimentary rocks of the basin from the Raleigh meta-
morphic belt and Carolina zone metavolcanics and
metasediments (Clark et ah, 2001). Minor faults form
the basin's western border, where Triassic sedimen-
tary rocks unconformably overlie Late Proterozoic and
Cambrian metavolcanic and metasedimentary rocks
(NCGS, 1985). Several minor faults have also been
recognized throughout the basin. Lrom north to south,
the Deep River Basin is traditionally divided into four sub-
structures: Durham subbasin. Colon cross-structure,
Sanford subbasin, and the Wadesboro subbasin (Bain
and Harvey, 1977). Some workers also recognize The
Pekin cross structure between the Sanford and
Wadesboro Basins (Clark et ah, 2001).
Bivalves described herein were recovered by North
Carolina Museum of Natural Sciences research staff
and volunteers in the late 1990s and early 2000s from
a brick/clay quarry in Durham County, North Carolina
(Figure 1). The quarry is located in the northeast
corner of the Green Level 7.5-minute Quadrangle at
35°52'14. 12" N latitude and 78° 53'51.77" W longi-
tude (coordinates are from Acme Mapper 2.0 http://
mapper.acme.com/ on June 10, 2012 using WGS 84) in
the Ullage of Genlee, Durham County, North Carolina
(Olsen and Huber, 1997). Strata in this quarry occur
in the south-central part of the Durham subbasin of
the Deep River Basin (Chatham Group of the Newark
Supergroup) (Sues et ah, 2003a), and form part of a
series informally designated as Litho facies Association II
(Hoffman and Gallagher, 1989). Kozur and Weems
(2007) placed this lithofacies in the Sanford Forma-
tion. The main quarry exposes about 60 m of red,
purple and gray fissile to bioturbated massive mud-
stones interbedded with gray, brown and red arkosic
sandstones (Olsen and Huber, 1997) and are considered
by Hoffman and Gallagher (1989) to be a series of
alternating fluvial and lacustrine facies. The recovery
of the vertebrate Aetosaurus from the quarry provides
a Norian age for the local section (Lucas et ah, 1998).
However, using conchostracan biostragraphy, Kozur and
Weems (2007) determined a latest Tuvalian (Carnian)
age for the section. As the eonehostracans and unionoid
bivalves come from layers well below the layers con-
taining the vertebrates, it is possible that quarry sedi-
ments straddle the Carnian/Norian boundary.
A modified version of the exposed section at the
quarry is given in Olsen and Huber (1997, fig. 3) and
earlier versions of the section are given in Olsen (1977)
and Olsen et ah (1989, 1991). From the fluvial facies
of this quarry, Parker (1966) and Lucas et ah (1998)
described a partial articulated skeleton of Aetosaurus
(— Stegomus ). Sues et ah (2003b) also reported an
articulated rauisuchian, Postosuchus sp., with gut con-
tents consisting of a partial skeleton of Aetosaurus
(—Stegomus), limb bones of a juvenile traversodont, a
partial dicynodont digit and a dermal bone of an inde-
terminate temnospondyl from one of the fluvial facies.
Also, from that same lithofacies, beneath the torso of
A. E. Bogan and P. G. Weaver, 2012
Page 107
Figure 1-6. Freshwater bivalves from the mid Late Triassic rift lakes of eastern North Carolina Triangle. 1. Brick Quarry,
Carpenter Plant brick pit. 2. Triaslacus carolinesis holotype, NCSM 9169A, total shell length is 10.54 mm. 3. Triaslacus carolinesis,
partaypes, NCSM 10781, internal mold. Total shell length = 9.74 mm. 4. Triaslacus carolinesis, specimen NCSM 11156, Cedar
Run, Virginia. Total shell length = 8.94 mm. 5. Possible Snhaeriidae? NCSM 10525. 6. Possible Mvtilidae? NCSM 10520.
crayfish; fragmentary and articulated fish; occasional
phytosaur teeth; and coprolites (Olsen and Huber,
1997). Also found in association with the bivalves, are
conchostraeans described by Kozur and Weems (2007)
as Euestheria buravasi, Euestheria sp. cf. E. lmusmanni ,
and Anijuanestheria ? new species.
the rauisuchian. Sues et al. (2003a) described an articu-
lated skeleton of sphenosuchian crocodylomorph,
Dromicosuchus grallator. The bivalves described herein
come from one of the lower lacustrine facies. Faunal
elements formyl in association with the bivalves include
darwinulid ostracodes; the trace fossil Scouenia ; cambarid
Page 108
THE NAUTILUS, Vol. 126, No. 3
MATERIALS AND METHODS
All specimens used in this description are housed in
the North Carolina State Museum of Natural Sciences,
Invertebrate Paleontology section, herein abbreviated
NCSM.
Shell measurements were made to the nearest milli-
meter using digital calipers and included total length and
height. Total length was defined as distance between
anterior and posterior margins, measured parallel to the
hinge line. Shell height is distance between dorsal and
ventral margins, measured near the midpoint of the
hinge line, perpendicular to shell length.
SYSTEMATIC PALEONTOLOGY
Class Bivalvia
Cohort Uniomorphi (=Palaeoheterodonta of authors)
Order Unionoida
Family cf. Unionidae
Genus Triaslacus new genus
Type Species: Triaslacus carolinensis new species
Diagnosis: A small edentulous freshwater bivalve, oval
in outline, lacking any umbonal sculpture.
Description: Triaslacus is distinguished from other
fossil unionoid species by a combination of the follow-
ing characteristics: Apparently thin, somewhat inflated,
shell, elliptical outline; hinge teeth absent; umbo only
slightly elevated above hinge line; umbo sculpture is
absent. Triaslacus can be distinguished from other Triassic
bivalves from northeastern United States assigned to the
Unionida by the following combination of characters.
It differs in shell shape from the elongate shell shape
assigned to Mulleriidae or the more rectangular shape
of species assigned to the Hvriidae. Triaslacus appear to
have a smooth umbo at or only slightly above the hinge
line, while other Triassic bivalve taxa have radial umbonal
sculpture and are assigned to the Hvriidae. Triaslacus
appears to be edentulous.
Distribution: Known from the Late Triassic rift lake
deposits in North Carolina and Virginia.
Etymology: The genus name Triaslacus stands for the
Triassic lake deposits vdrere it was collected.
Triaslacus carolinensis new species
(Figures 2-4)
Unionidae sp. — Bain and Han ey, 1977;
Good 1995a; 1995b.
Diagnosis: Triaslacus carolinensis is distinguished
from all other knowm Late Triassic unionoid bivalves
occurring along the east coast of North America by the
oval shell outline in contrast to the elongate shell in
species of Mycetopoda . It lacks the radial umbonal sculp-
ture of Diploclon species and does not have the square
shell outline of Unio emersoni.
Description: Length to 12.8 mm, height to 7.5 mm;
shell thin, moderately inflated, outline oval. Posterior
margin narrowly rounded to bluntly pointed. Anterior
margin broadly rounded. Dorsal margin straight. Ven-
tral margin convex. Posterior ridge rounded. Posterior
slope moderately steep. Umbo broad, moderately inflated,
barely elevated above hinge line. Umbo sculpture absent.
Pseudocardinal and lateral hinge teeth absent. Umbo
cavity wide, shallow.
Comparison with Similar Species: Triaslacus
carolinesis shells do not resemble the shell referred
to as Mycepoda ? dilicidi Pilsbry in Wanner, 1921and
placed in the Mycetopodidae, which is nowadays con-
sidered to be restricted to South America. Triaslacus
carolinesis does not have the radial umbonal sculpture of
those taxa referred to the genus Diplodon (Table 1) and,
by extension, to the family Hyriidae, today restricted
to South America, Australia, New Zealand, and New
Guinea. Diplodon carolussimpsoni from Pennsylvania
is more rectangular in shell shape and has 'a well-
marked posterior ridge. Unio emersoni is more elongate
is shell outline, larger in maximum shell length and
possess large hinge teeth, while Triaslacus carolinesis
is smaller in size, more oval in shell outline and is
edentulous. Anoplophora wilbrahamensis is more elon-
gate oval in outline, larger in maximum shell length
than Triaslacus carolinesis and has a long lateral tooth,
while T. carolinesis is edentulous.
Type Material: Holotype, NCSM 9169, internal mold,
both valves. Paratypes, NCSM 10747, negative relief;
NCSM 10757, NCSM 10760, cluster; NCSM 10767,
negative relief; NCSM 10769, NCSM 10778, positive
relief; NCSM 10781, positive relief; NCSM 10790,
positive relief. All from the type locality.
Type Locality: United States, North Carolina, Durham-
Wake County, Triangle Brick, Carpenter Plant in the
village of Genlee. Northeast corner of the Green
Level 7.5' quadrangle at 35° 52' 14.12" N latitude and
78°53/ 51.77" W longitude (coordinates are from Acme
Mapper 2.0 http://mapper.acme.com/ on June 10, 2012)
(Figure 1).
Other Material Examined: United States, North
Carolina, Durham-Wake counties, Triangle Brick,
Carpenter Plant: NCSM 5035.1-5035.9 all positive/
negative; NCSM 5036, 38 specimens; NCSM 5037.1-
5037.9, some are both positive/negative; NCSM 6135,
total of 7 specimens; NCSM 10745-10746; NCSM
10748-10756; NCSM 10758-10759; NCSM 10761-
10766; NCSM 10768; NCSM 10770-10780; NCSM
10782-10788; NCSM 10791; NCSM 10794-10818;
United States, Virginia, Fauquier County, Carriage
A. E. Bogan and P.G. Weaver, 2012
Page 109
Ford, Nokesville. Collector: Robert Weems coll. NCSM
11132-11139; United States, Virginia, Hanover County,
Stagg Creek transect, Taylorsville Basin, Hannover
Academy Quadrangle, Robert Weems coll.; NCSM
11140-11150; United States, Virginia, Fauquier County,
Cedar Run. Catlett Quadrangle, Robert Weems coll.,
NCSM 11151-11158
Etymology: The species is named for the state of
North Carolina, where the species was first collected.
Distribution: Triangle Brick Quarry Carpenter Plant,
Gen Fee, Durham-Wake County, North Carolina.
Carriage Ford, Nokesville, Fauquier County, Virginia.
Stagg Creek transect, Taylorsville Basin, Hannover Acad-
emy Quadrangle, Hanover County, Virginia and Cedar
Run, Catlett Quadrangle, Fauquier County, Virginia.
Stratigraphy: Mid Fate Triassie (Carnian-Norian
Stage interval) Newark Supergroup, Lithofacies II.
Habitat: Triaslacus carolinesis is found in similar
sedimentary deposits bearing similar associated faunal
elements in Durham County North Carolina, and Fauquier
County, Virginia. Good (1995a, b) suggested the small or
“dwarfed” size of the unionoid is due to “environmental
conditions not optimum for this group, probably indi-
cating a deep water lacustrine habitat.” Small shell size
is common in modern unionoids and some species are
found at depths over 15 m and they are not dwarfed
(Bailey and Green, 1989). The lake environment was likely
rather shallow and soft bottomed, containing aquatic
vegetation, associated fish, ostracods and clamshrimp.
The latter species is found in lacustrine facies sug-
gesting soft-bottomed deposits in quiet water similar
to habits of the common modern thin-shelled eastern
North American anodontine, Utterbackia imbecillis
(Say, 1829).
Order Cardiida?
Family Sphaeriidae?
(Figure 5)
Description: Shell almost round in outline with some
valve inflation, reminiscent of modern Sphaeriidae,
None of the specimens have exposed hinge plates
that would allow for family level confirmation (see
Mackie, 2007).
Habitat: Sphaeriids occur today throughout the world
in freshwater habitats similar to those described here
for Lithofacies II.
Geologic History: Keen (1969) lists Late Jurassic? as
the oldest record for the family Sphaeriidae. II cor-
rectly identified, this record would extend the geologic
range of the family back to the Late Triassie.
Material Examined: NCSM 10521-10524, 10819,
I 1 157, 11158, from Lithofacies II, Triangle Brick Quarry
Carpenter Plant, Gen Lee, Durham-Wake County,
North Carolina and Cedar Run, Catlett Quadrangle,
Fauquier County, Virginia NCSM 1 1157—11 160. Speci-
mens 1 1 157 and 1 1 158 also have Triaslacus on the
same block.
Cohort Mytilomorphi
Order Mvtilida?
Family Mvtilidae?
(Figure 6)
Description: Specimens with a mvtilid shell shape,
with umbo area very anterior and a rather elongate oval
shape are tentatively referred to Mytilidae.
Geologic History: This family is reported as occur-
ring from the Devonian to the Recent (Soot-Ryen, 1969).
Material Examined: NCSM10520, 10820-10822,
Lithofacies II, Triangle Brick Quarry Carpenter Plant,
Gen Lee, Durham-Wake County, North Carolina.
Remarks: Pilsbry (1926) described Naiaclites tnassicus
Pilsbry in Wanner, 1926 and Naiaclites wanneri Pilsbry in
Wanner, 1926, recognizing that this genus was known
from the Coal Measures [Upper Carboniferous] of Nova
Scotia and far earlier than the Triassie deposits these
specimens were collected. Weir ( 1969) placed Naiaclites
in the family Myalinidae.
Good (1995a, b) claimed that the identification of
the small associated bivalves from these deposits are
non-marine Myalinidae. As Good pointed out, they are
not Cyrenidae (+ Corbiculidae), but some appear to
resemble Mytilidae (Figure 6) and others Sphaeriidae
(Figure 5). This does not, as yet [?], call for a Lazarus dis-
tribution between the Pennsylvanian and the Late Triassie.
DISCUSSION
Freshwater bivalves from the Durham subbasin and
the Taylorsville Basin of Virginia are described here as a
new genus and species of freshwater bivalve tentatively
assigned to the family Unionidae. Triaslacus carolinesis
differs from the Triassie freshwater bivalves described
from Connecticut, Massachusetts, and Pennsylvania in
shell outline, inflation and umbonal sculpture. Triaslacus
carolinesis is only tentatively assigned to the Unionidae
based on the shell outline, lack of radial umbonal sculp-
ture and lack of hinge teeth. The ambiguity of family
assignment is due to rampant shell shape convergence
or shell homeomorphy in freshwater bivalves (Ortmann,
1912; Prashad, 1931; Watters, 1994)).
White (1907), writing on the origin of the North
American fossil freshwater bivalve fauna before the
work of Pilsbry (1921), felt that the modern unionoid
fauna was probably derived from the Triassie fauna of
the southwestern United States.
Pilsbry (1921) described Triassie freshwater bivalves
for the northeast United States and suggested that this
fauna is composed of taxa assigned to the modern families
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THE NAUTILUS, Vol. 126, No. 3
Hyriidae and Mycetopodidae and suggested that proba-
bly all of the North American fossil Triassic freshwater
bivalves were related to the South American fauna. He
felt the hyriid mussels would have disappeared with the
migration of "Old World forms in the Upper Trias
or Jurassic’ .
Parodiz (1969) followed this line of reasoning and
claimed the Hyriidae of the Triassic of North America
migrated into South America in the Paleocene.
The evolution of freshwater bivalves in North America
has most recently been summarized by Watters (2001),
who agreed that Hyriidae occurred in North America
in the Triassic. However, Watters does raise the problem
of convergence in shell sculpture and notes the lack
of shell characters to separate these fossils from the
modern Hyriidae.
One of the five major extinction events occurred at
the end of Triassic. Timing and extent of this extinction
event is controversial, catastrophic vs. gradual (e.g. Olsen
et ah, 1987; Palfy et ah, 2000; Hallam, 2002; Tanner
et ah, 2004; Deenen et ah, 2010). Taking into consider-
ation terrestrial extinctions at the end of the Triassic and
the expanding rift zone along eastern North America,
animals living in these rift lakes appear to have been
slated for extinction. Triassic freshwater bivalves col-
lected from the rift lakes of the eastern United States
most likely represent an extinct radiation and as such,
may not have any relation to the currently recognized
six families in the Unionida and may not represent the
origin of any of the modern Unionida families.
ACKNOWLEDGMENTS
We want to thank Richard Chandler for his assistance
with bivalve and conchostracan photography; Vince
Schneider for assistance with field photography; Rob
Weems for assistance with conchostracan photography;
Vince Schneider, Verne Lee, Heniy Rust, Valeria Rice,
John Adams, Rob Weems and Heinz Kozur are all
acknowledge for field assistance; Earle E. Spamer,
formerly with the Academy of Natural Sciences,
Philadelphia, PA, assisted with the location of other
Triassic unionoid specimens in Academy’s collections.
Mary Dejong, formerly with the American Museum of
Natural History for assistance with literature. Cindy
Bogan reviewed and commented on a draft of the
manuscript. Dr. Joseph Hartman is acknowledged for
his very thorough review and thoughtful comments.
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America, and its implications for the Worldwide fauna.
In: G. Bauer and K. Wachtler (eds.) Ecology and Evolu-
tion of the Freshwater Mussels Unionoida. Ecological
Studies, v. 145. Springer- Verlag, Berlin, pp. 281-307.
Weir, J. 1969. Nonmarine and brackish-water Myalinidae.
In: R.C. Moore (editor). Treatise on Invertebrate Paleon-
tology. Geological Society of America and the University
of Kansas, Part N, Volume 1 [of 3], Mollusca 6. Bivalvia,
pp. N29 1-295.
White, C. A. 1907. The ancestral origin of the North American
Unionidae, or fresh-water mussels. Smithsonian Miscella-
neous Collections 48: 75-88, plates 26-31.
THE NAUTILUS 126(3): 113-116, 2012 Page 113
Research Note
An albino shell of Nautilus pompilius
(Cephalopoda: Nautilidae)
from the Philippines
ABSTRACT
An albino shell of Nautilus is described. The shell is 174.5 mm
in diameter with a whorl width at the aperture of 81.9 mm.
It bears a black band at the aperture indicating that it is a
mature animal. The morphology of the shell conforms to the
general species description of Nautilus pompilius Linnaeus,
1758, in all respects, except that it lacks any reddish-brown
color stripes. The lack of color stripes on the shell is interpreted
as a case of albinism, although it is unknown if the soft parts
also lacked color. This is the first reported case of an albino
Nautilus although such shells have periodically cropped up
over the years for sale on the internet.
Additional keywords: Albinism, Nautiloidea, Nautilida
INTRODUCTION
All shells of Nautilus and Allonautilus bear reddish-brown
color stripes (Saunders, 1981) although their exact pattern,
hue, and intensity vary. The function of the color pattern is
interpreted as camouflage (Cowen et al., 1973). The stripes
appear in early ontogeny and persist onto the juvenile shell.
They disappear on the adoral one-third of the body cham-
ber coincident with the attainment of maturity. We report a
specimen of Nautilus that lacks any color bands.
REPOSITORY COLLECTION
The specimen is reposited at the American Museum
of Natural Histoiy (AMNH 275269) and was donated
by Mrs. Peter R. Fleischner between 1992 and 1995.
According to tire accompanying label, the specimen was
"collected in traps off Zamboanga, Mindanao, Philippines”.
However, as described below, the inside surface of the
body chamber bears several bryozoan colonies that must
have grown after the death of the Nautilus , indicating that
the shell must have been collected post-mortem.
DESCRIPTION
AMNH 275269 is a mature specimen based on the pres-
ence of a deep ocular and hyponomic sinus and a thick-
ened apertural margin with a black band 0.5 to 1.5 mm
wide (Figures 1-4). The shell conforms to the general
description of Nautilus pompilius Linnaeus, 1758. It is
1 74.5 mm in diameter, close to the average size at maturity
ol this species from the Philippines (165 mm reported in
Saunders, 2010: p. 39), with a whorl width at the aperture
of 81.9 mm. The umbilicus is small and covered with a
well-developed umbilical callus, a hallmark of this species.
The most unusual feature of this specimen is the por-
cellaneous white color of the shell and the absence of
any reddish-brown color stripes. This contrasts with the
well -developed black layer on the dorsum. In small
areas where the black layer on the dorsal surface of
the shell has chipped away, the shell is also white. The
specimen does not show any scratches or polishing or
any indication of bleaching that would have altered its
appearance. Growth lines are prominent with approxi-
mately 8 to 16 growth lines/5 mm as measured on the
flanks approximately 70 mm adapieal of the aperture.
Faint longitudinal lirae are also present on the lateral sides
and venter of the conch. There are two minor repaired
shell breaks, one on the venter on the adoral one-third
of the whorl, and one on the right apertural margin.
Each is associated with deposits of black material.
The shell bears evidence of epizoans. A bryozoan
colony is covered over by the black layer on the dorsal
side. Several traces of bryozoan colonies also appear on
and around the umbilical callus (Figure 5). There are
also traces of two possible serpulid worm tubes on the
left lateral side. Most importantly, the inside surface of
the shell near the aperture is covered with several cir-
cular to semicircular bryozoan colonies with diameters
of up to 15 mm (Figure 6). These bryozoans indicate that
the specimen was dead when it was recovered. It was
probably floating near the surface and was accidentally
caught in a fish trap or net, which is not an uncommon
occurrence in the Philippines (Dunstan et al., 2010).
DISCUSSION
Normally, shells of Nautilus develop irregular brown
to reddish-brown bands or stripes that extend from
the umbilicus and branch across the flanks, sometimes
coalescing on the venter. In Nautilus pompilius , the ini-
tial part of the embryonic shell is ivory colored, but
at approximately I whorl adoral of the apex, the shell
develops a pale orange color that subsequently becomes
darker. The white and brown color pattern appears
gradually, first on the flanks and later on the venter
(Arnold et al., 2010). The pattern becomes visible on
the flanks about one quarter to one half whorl adapieal
of the nepionic constriction (the point of hatching). Ini-
tially, the boundaries between the color bands are indefi-
nite, but with later growth, they become better defined.
After hatching, the color pattern becomes darker reddish
brown with sharply defined white bands. In juvenile ani-
mals, the color stripes persist to the aperture. The color
Page 114
THE NAUTILUS, Vol. 126, No. 3
Figures 1-4. Mature albino shell of Nautilus pompilius (AMNII 275269) from the Philippines. 1. Right lateral. 2. Apertural.
3. Ventral. 4. Left lateral. Specimen diameter = 174.5 mm.
R. H. Mapes and N. H. Landman, 2012
Page 115
Figures 5-6. 5. Bryozoan colonies on the left umbilical callus. 6. Bryozoan colonies inside the apertural margin, which is marked
by a black band (top of photo).
bands disappear on the venter approximately one third
whorl adapical of the mature aperture, corresponding to
the approximate position of the last septum. This pro-
duces a white venter, so that the colored area is restricted
to the dorsal part of the shell (Collins and Ward, 2010).
Only one specimen of Nautilus pompilius was ever
reported in which a color band reappears on the white
ventral part of a mature shell (Trego, 1993).
The color bands are secreted by pigment secreting
cells at the edge of the mantle. The pigment is produced
by reddish-brown melanin and is restricted to the outer
spherulitic-prismatic part of the shell (Stenzel, 1964).
It can be removed by polishing oil this layer revealing
the iridescent nacreous layer below. Because the pattern
involves alternating color and no color and crosses the
growth lines, the secretory activities of the cells must
sequentially start and stop.
The lack of any color striping on our shell suggests that
this animal was an albino. As seen through chips in the
black layer, the shell must have been white throughout
ontogeny. It is unknown if the hood or other tissues
reflected this unusual lack of color. However, it is note-
worthy that the black area on the dorsum and the black
band at the aperture are present. Both of them consist of
black melanin. The black area on the dorsum is secreted
by the dorsal part of the mantle covering the earlier
secreted part of the shell (Signor, 1985). The black band
at the aperture is secreted by the mantle at maturity in
association with a termination in shell growth (Collins
and Ward, 2010), implying that the cessation of arago-
nitic secretion is accompanied by the deposit of melanin.
Color striping on Nautilus has been interpreted as
camouflage to enhance protection from visual predators
such as sharks and other fish (see the discussion by
Mapes and Davis, 1996). Viewed from above, the normal
adult shell exhibits transverse bands of reddish-brown,
which serve for counter-shading so that incident light
does not illuminate the surface (Cowen et al, 1973).
Page 116
O
THE NAUTILUS, Vol. 126, No. 3
Additionally, Stenzel (1964) argued that the color bands
disrupt the contour of the animal, breaking up its shape
and outline. Viewed from below, the adult shell is white,
blending in with the surface above.
This albino Nautilus would presumably have been
more easily detected by visual predators from above or
from the side because of its lack of color striping com-
pared to normal animals, thereby reducing its probability
of surviving to maturity. Alternatively, the white color
may have fooled potential predators because they did
not recognize the animal as their usual food item. How-
ever, the shell bears two repaired injuries implying that
the animal experienced sublethal predation. It is pos-
sible that it survived to maturity because it spent most
of its time below the photic zone out of sight of visual
predators, the preferred habitat of Nautilus (Saunders
and Ward, 2010).
Recently, we learned that two other albino speci-
mens of Nautilus pompilius were recovered from the
Philippines (W. B. Saunders, pers. comm., 2012). They
were advertised on the internet by a shell dealer, who
sold them to a collector. The dealer mentioned that
another albino shell had been collected in the area
ten years ago. Both of the shells advertised on the
internet exhibited heavily modified apertures. One
shell, with a diameter of 119.4 mm, was collected by
trawling at 400 m off Mindoro, Philippines, and the
other shell, with a diameter of 125.8 mm, was recovered
by trawling at 350-400 m off Laminusa Island, approxi-
mately 600 km south of Mindoro. The occurrence of
these shells, plus our own, suggests that albino specimens
are rare but present in the population of N. pompilius in
the Philippines.
ACKNOWLEDGMENTS
We thank Steve Thurston (AMNH) who prepared the
photographs of the specimen, Bushra Hussaini (AMNH)
who assisted R.H.M. in the collections, and Peter Ward
(University of Washington) who reviewed an earlier draft
of this manuscript and made many helpful suggestions.
LITERATURE CITED
Arnold, J.A., N.I1 Landman, and H. Mutvei. 2010. Develop-
ment of the embryonic shell of Nautilus. In: W. B.
Saunders and N.H. Landman (eds.), Nautilus : The Biology
and Paleobiology of a Living Fossil Springer, New York,
pp. 373-400.
Collins, D. and P. D. Ward. 2010. Adolescent growth and matu-
rity in Nautilus. In: W. B. Saunders and N.H. Landman
(eds.) Nautilus : The Biology and Paleobiology of a Living
Fossil. Springer, New York, pp. 421-432.
Cowen, R., R. Gertman, and G. Wiggett. 1973. Camouflage
patterns in Nautilus , and their implications for cephalopod
paleobiology. Lethaia 6: 201-214.
Dunstan, A., A. Alanis, and f. Marshall. 2010. Nautilus
pompilius fishing and population decline in the Philippines:
a comparison with an unexploited Australian Nautilus
population. Fisheries Research 106: 239-247.
Mapes, R . II.. and R.A. Davis. 1996. Color patterns in anuno-
noids. In: N.II. Landman, K. Tanabe, and R.A. Davis
(eds.) Ammonoid Paleobiology, Topics in Geobiology 13,
Plenum Press, New York and London, pp. 103-127.
Saunders, W. B. 1981. The species of Nautilus and their dis-
tribution. The Veliger 24: 8-17.
Saunders, W.B. 2010. The species of Nautilus. In: W.B.
Saunders and N.II. Landman (eds.) Nautilus : The Biology
and Paleobiology ol a Living Fossil. Springer, New York,
pp. 35—52.
Saunders, W. B., and P. D. Ward. 2010. Ecology, distribu-
tion, and population characteristics in Nautilus. In: W.B.
Saunders and N.H. Landman (eds.) Nautilus : The Biology
and Paleobiology of a Living Fossil. Springer, New York,
pp. 201-212.
Signor, P. 1985. Surficial shell resorption in Nautilus
macromplialus Sowerby, 1849. The Veliger 28: 195-199.
Stenzel, II. B. 1964. Living Nautilus. In: Treatise on Invertebrate
Paleontology (Part K) Geological Society America and Uni-
versity of Kansas Press, Lawrence, Kansas, pp. K59-K93.
Trego, K.D. 1993. Band color pattern on the venter of a
mature shell of Nautilus pompilius Linnaeus, 1758. The
Veliger 36: 430-431.
Royal H. Mapes
Department of Geological Sciences
Ohio University
Athens, OII 45701 USA
[email protected]
Neil H. Landman1
Division of Paleontology (Invertebrates)
American Museum of Natural History
79th St. and Central Park West
New York, NY 10024
[email protected]
O
1 Corresponding author.
In the ACKNOWLEDGMENTS section of the article by Lee and Huber (2012), please replace “Ms. Way facilitated
the delivery of the images taken by S. Peter Dance of C. medium in the Linnean Society Collection" for: "The images of
the Linnaean collections available via the Society’s website were taken by staff of the NHMUK Science Photography unit
in 2008. The A4 sheet which accompanies each set of images is a transcription of the information on the label written by
Peter Dance in the mid-1960s for each lot in the collection strongroom."
LITERATURE CITED
Lee II. G. and M. Huber. 2012. Americardia lightboumi new species and A. cohnnbella new species compared to A. media
(Linnaeus, 1758), A. speciosa (A. Adams and Reeve, 1850), and the extinct A. Columba (Heilprin, 1886) (Bivalvia: Cardiidae).
The Nautilus 126: 15-24.
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THE NAUTILUS 126(4): 1 19-126, 2012
Page 119
The fossil record and phylogeography of the family Cerionidae
(Gastropoda: Pulmonata), with the description of a new species
from the Pleistocene of Florida
M.G. Harasewych
Department of Invertebrate Zoology, MRC-163
National Museum of Natural History
Smithsonian Institution
PO Box 37012
Washington, DC 20013-7012 USA
[email protected]
ABSTRACT
Cerion petuchi , new species, the first record of the genus from
the Pleistocene ol Florida, is described from Loxahatchee,
Florida, from deposits of the Loxahatchee Member of the
Bermont Formation (Aftonian Pleistocene). This new species
is more similar to Cerion agassizii from the Pleistocene of the
Great Bahamas Bank, and to the Recent Cerion incanum
saccharimeta, from the Florida Keys, than to either of the spe-
cies from the Late Oligocene — Early Miocene Ballast Point
Silex Beds of Tampa, Florida. Data on the geographic distribu-
tion and geological age of all known cerionids is compiled and,
together with models of die geologic and tectonic history of the
Caribbean region, used to construct a hypodiesis of the origins,
ages, and relationships among the various lineages within the
family since the Upper Cretaceous. The early distribution
of the family was governed primarily by overland dispersal
and vieariance. It is suggested that the significant proliferation
of diversity that began during the Pleistocene is due to the
increased prevalence of stochastic dispersal of small propagules
(either by rafting or hurricane-born) among the islands of the
Cuban and Bahamian archipelagos. Sea level changes caused by
Pleistocene glaciations amplified diversity by repeatedly and
sequentially recombining / hybridizing and isolating neighboring
populations. Amplification of diversity would have been greatest
on the Great Bahama Bank, as it had the highest number of
islands that were isolated during interglacial periods and con-
joined during glaciations.
Additional keywords: Cerion , diversity, zoogeography, vieariance,
dispersal
INTRODUCTION
The family Cerionidae has long been known for its excep-
tional morphological diversity and the profusion of species
level taxa in the Recent fauna (e.g., Maynard, 1889-96;
Pilsbry, 1901-02; Clench, 1957; Woodruff, 1978). In con-
trast, the fossil record of the Cerionidae, which extends
from the Upper Cretaceous (Roth and Hartman, 1998),
is poorly known and represented by comparatively few
taxa, most based on rare specimens. A notable exception
is in the fauna of the Bahamas, where Cerion are locally
common in Pleistocene and younger deposits (Hearty
and Kaufman, 2009). The geographic and ecological
ranges of the family have recently been expanded con-
siderably by inclusion of the genera Brasilennea from the
Paleocene of the Itaboraf Basin, Brazil (Salvador et al.,
2011; Salvador and Simone, 2012), and Mexistrophia ,
from cool, coniferous forests in the highlands of central
Mexico (Thompson, 2012).
Within Florida, Cerionidae is represented by two spe-
cies (one with two subspecies) from the Late Oligocene-
Early Miocene Ballast Point Silex Beds of Tampa, Florida,
and by a single native Recent species (subdivided into
four subspecies) that is confined to the Florida Keys.
Thirteen non-native species from the Bahamas, Cuba,
Puerto Rico, and Curasao were intentionally introduced
into the Florida Keys and Dry Tortugas between 1912
and 1924 by Paul Bartsch as part of a long-running and
well-documented series of experiments (see Harasewych
and Strauss, 2006: Table 1). An additional taxon, Cerion
tridentata costellata Pilsbry, 1946, despite a type locality
of Garden Key, Florida, is a hybrid of two of the intro-
duced Cuban species (Harasewych et ah, 201 1 ).
One complete and one partial specimen of Cerion
have recently been discovered in Loxahatchee, Florida,
from fossil deposits of Pleistocene age (Bermont Forma-
tion). These specimens represent a previously undis-
covered species that is described as new and compared
to fossil and Recent congeners.
The diversity of the family Cerionidae is reviewed
from geographic and temporal perspectives. These
results are combined with paleogeographic reconstruc-
tions of the Caribbean region (Iturralde-Vinent, 2006) to
produce a zoogeographic hypothesis for the dispersal of
Cerionidae during the Late-Cretaceous and Cenozoic.
A recently published molecular phytogeny of Cerionidae
Page 120
THE NAUTILUS, Vol. 126, No. 4
Table 1. Diversity of species level taxa within the family Cerionidae throughout its geographic and geological range. Data derived
from the Cerion website [http://invertebrates.si.edu/cerion/] (Harasewych, 2009).
(1) These figures do not include the species introduced into Florida by Bartsch during the first half of the 20th Century (see
Harasewych and Strauss, 2006: Table 1), nor does it include Cerion tridentata costellata , which, despite a type locality of Garden Key,
Florida, is a hybrid of introduced Cuban species (see Harasewych et ah, 201 1 ).
(2) Some species were reported both as Recent and Pleistocene.
(3) Clench (1961) synonymized six taxa from Turks Island under Cerion regina Pilsbry and Vanatta, 1895.
(4) Gould and Pauli (1977) synonymized all living Cerion taxa from Hispanola, Mona Island, and Puerto Rico and the Virgin
Islands under Cerion striatellum Guerin-Meneville, 1829, but retained as separate the fossil C rude (Pfeiffer, 1855) from St. Croix.
(5) The Cerion of Aruba, Bonaire, and Curasao had been subdivided into four subspecies some with large and dwarf named forms
(Baker, 1924). Gould (1984) showed that the four subspecies correspond to geographic subdivisions, while the forms are non-
adaptive ecophenotypic variations.
(Harasewych et ah, 2011) is used to evaluate portions of
this hypothesis.
SYSTEMATICS
Cerion petuchi new speeies
(Figures 1-1 1 )
Diagnosis: Shell large (to 36 mm), with evenly taper-
ing, conical spire, thick, finely ribbed, with broadly ovate
aperture, widely flaring, simply recurved outer lip. Inner
lip with parietal region broader than columella, each
with a single tooth at mid-length. Umbilicus imperforate.
Description: Shell (Figures 1-9) large (holotype
36.05 mm long, 13.87 mm in diameter), thick, roundly
conical in profile, with each whorl slightly broader than
the preceding whorl. Protoconch (Figure 10) initially
smooth, increasing in diameter from 504 pm to 3.26 mm
in 2.6 whorls, with sculpture of 36 weak, evenly spaced
axial ribs as broad as the intervening spaces appearing
on the final half whorl. Transition to teleoeonch (Fig-
urelO, p/t) marked by an abrupt increase in shell diame-
ter and onset of coarser surface sculpture. Teleoeonch of
9>/s evenly rounded whorls. Suture tightly adpressed. Axial
sculpture of multiple (~96 on first teleoeonch whorl, ~74
on final whorl) low rounded prosocline ribs (~18° to coil-
ing axis) that generally align with interspaces of prior
whorl. Spiral sculpture absent. Aperture expands, deflect-
ing suture adapically ~‘/s whorl prior to formation of
thickened and flared terminal lip. Aperture broadly ovate,
with the parietal wall forming an angle of 112° with the
shorter columella. Single parietal tooth (Figure 1 1, par)
emerges from mid-point of parietal callus. A weaker,
broader tooth (Figure 11, col) recessed along mid-length
of columella. Umbilicus imperforate, obscured by expanded
terminal lip.
Type Locality: Palm Beach Aggregates, Inc. (old
GKK) pit number 7, Loxahatchee, Palm Beach County,
Florida [26°42.20' N, 80°20.97 W], at 15 m depth.
Stratigraphy: From the Holey Land Member of the
Bermont Formation. Aftonian Pleistocene (~1.6 million
years before present) (Petueh and Roberts, 2007: 147-162).
Type Material: Holotype, USNM 1191690; Paratype 1,
USNM 1191691, both from the type locality.
Distribution: Known only from the type locality. This
species was living in tire Tomeu Paleoislands, along
the northern end of Palm Beach Paleoarchipelago,
Loxahatchee Subsea, Okeechobean Sea during the
Aftonian Pleistocene (Petueh, 2003: fig. 31).
Etymology: This species honors Dr. Edward ].
Petueh, who discovered it, in recognition of his many
contributions to our understanding of the Recent and
M.G. Harasewyeh, 2012
Page 121
Figures 1-9. Cerion petuchi new species. 1. Apertural, 2. Lateral, 3. Dorsal, 4. Apical and 5. Anterior views of the holotype
(USNM 1191690). 6. Apertural, 7. Lateral, 8. dorsal, and 9. Anterior views of paratype fragment (USNM (XI 191691). Palm Beach
Aggregates, Inc. pit number 7, Loxahatchee, Palm Beach County, Florida. [26°42.20' N, 80°20.97' W] At 15 m depth. Holey Land
Member of the Bermont Formation. Aftonian Pleistocene (~1.6 million years before present).
fossil molluscan faunas of Florida and the tropical west-
ern Atlantic.
Remarks: The large size, conical shell shape, presence
of multiple, fine, axial ribs and a simple, flared lip serve
to distinguish Cerion petuchi new species from all fossil
cerionids reported from the Late Oligoeene / early Mio-
cene silex beds of Ballast Point, Florida. Cerion
( Microcerion ) floridanum Dali, 1915 had a much smaller
shell with smoothly rounded whorls and a distinctive,
extremely broad, eoncavely rounded outer lip. Both
Cerion ( Eostrophia ) anodonta Dali, 1890 and Cerion
( Eostrophia ) anodonta var. floridanum Dali, 1915 had
shells that are cylindrical in shape, with a smooth surface
and smaller, rounder apertures. Pilsbry (1946: 161)
reported a single fossil specimen that he identified as
Cerion incanum from Clewiston, Florida, from the youn-
ger Fort Thompson Formation (Pleistocene). This spec-
imen, originally in the McGinty collection, was not
illustrated and could not be located.
Of the Recent Cerion inhabiting the Florida Keys,
Cerion petuchi new species most closely resembles
Cerion incanum saccharimeta Pilsbry and Vanatta, 1899,
which is also characterized by its large size and conical
shell shape, but differs in having coarser and more irreg-
ularly spaced axial sculpture, a rounder aperture and a
simpler, less flared outer lip. Cerion petuchi new species
most closely resembles Cerion agassizii Dali, 1894, from
the Pleistocene (ca. 120,000 years BP) dunes of New
Providence Island (Bahamas), a species that is readily
distinguished on the basis of having fewer, more irregu-
larly spaced axial ribs, more pronounced parietal and
Page 122
THE NAUTILUS, Vol. 126, No. 4
Figures 10-11. Cerion petuchi new species. 10. Apical view
of protoconch of holotype. 11. Details of apertural dentition
of holotype.
columellar teeth, and a “duplex” outer lip, which forms a
concavely rounded, bow-like structure most pronounced
along the anterior margin of the aperture. Cerion
picturata (Maynard and Clapp in Maynard, 1921) from
Pleistocene/Holocene deposits on Cabbage Key in the
Berry Islands (Bahamas) is also similar. It lacks the
“duplex” outer lip, but differing in having coarser, more
axially aligned sculpture. A similar phenotype survives in
the Recent fauna of Guana Key, Berry Islands as Cerion
arbusta (Maynard and Clapp in Maynard, 1919).
GEOGRAPHIC AND TEMPORAL DIVERSITY
OF THE CERIONIDAE
Data on geographic distribution and geologic age for the
600 species level taxa within the family Cerionidae were
compiled based primarily on information from the Cerion
website (Harasewych, 2009). Taxa were assigned to islands
or island groups based on their type localities. Objective
synonyms and nomina nuda were excluded, as were taxa
with unknown [e.g., Cerion cumingiana (Pfeiffer, 1852)]
or clearly erroneous type localities [e.g., Cerion antonii
(Kiister, 1847) from British Guiana], but subjective syno-
nyms were generally counted as separate taxa. Results
are shown in Table I
Several authors have noted the multitude of taxonomic
names applied to the abundant phenotypes within the
Cerionidae and speculated that the actual number of
valid species may be on the order of 1-20% of the num-
ber of names now extant (e.g., Clench, 1957; Woodruff,
1978). Although the numbers of taxa appearing in Table 1
will inevitably be revised downward based on detailed
systematic studies, certain patterns are striking and will
likely persevere. Most conspicuous is the observation
that by far the greatest diversity of Cerionidae occurs on
the Great Bahama Bank (47.2%, of all named Recent
species level taxa), while the fauna of Cuba comprises
32.8% of all Recent species level diversity. Both these
areas have alternated between being large, contiguous
land masses and archipelagos during the glacial and
interglacial sea-level fluctuations of the Pleistocene and
Holocene. Iturralde-Vinent (2006:figs. 7,8,10,11) has shown
that Cuba consisted of a series of isolated islands from
the Oligocene to the Late Pleistocene, with the present
shape of the island attained 8,000-6,000 years ago.
By contrast, the combined fauna of the southeastern
Bahamian Islands and the Turks and Caicos Islands, which
would have remained isolated during the lowest sea levels,
comprises only 9.3% of total Recent cerionid diversity.
None of the remaining regions populated by cerion ids
contribute more than 3.5% of the Recent diversity.
This pattern is even more pronounced during the
Pleistocene / Holocene, when the Great Bahama Bank
accounted for 90.0% of the 50 described species level
taxa. The isolated southeastern Bahamian Islands collec-
tively accounted for 4.0% of the named fauna, with only
single species (2.0% of the fauna) reported from Cuba,
Florida and tire western Virgin Islands. Prior to the Pleis-
tocene, the fossil record is exceedingly sparse, with only
three taxa reported from the Oligocene, three from the
Paleoeene, and one from tire Cretaceous.
A ZOOGEOGRAPHIC HYPOTHESIS
FOR CERIONIDAE
The early fossil record of the Cerionidae consists of
few taxa represented by rare specimens that neverthe-
less provide insights into the zoogeographic history of
the family. The earliest record for the family ( Cerion
acherontis Roth and Hartmann, 1998) is based on a
single, damaged specimen from the Late Cretaceous of
Montana. The genus Brasilennea, represented by three
species from the Paleoeene of Itaborai Basin, Brazil,
(Salvador et ah, 2011; Salvador and Simone, 2012) has
M.G. Harasewyeh, 2012
Page 123
recently been transferred from the family Streptaxidae
to the Cerionidae. Western Laurasia (North America)
and western Gondwana (South America) were sepa-
rated by a marine waterway during the Jurassic
(Callovian), with land bridges developing during the
Late Cretaceous (Campanian / Maastrichtian) (75-65 Ma)
via a proto-Antillean island arc, and again during the
Plio-Pleistocene (2.5 Ma) via the Panama land bridge
(Iturralde-Vinent, 2006). Parodiz (1969: 189, Map 7)
noted that many of the families of South American non-
marine mollusks are of Nearctic origin, having migrated
to South America at the close of the Cretaceous. Given
the presence of cerionids in South America during the
Paleocene, it is probable that members of this family
were part of this Late Cretaceous migration.
The recent description of the genus Mexistrophia
from cool, coniferous forests in the highlands of central
Mexico (Thompson, 2012), suggests that this genus was
an early offshoot that diverged during the Late Creta-
ceous, prior to the adaptations to near shore, halophilie
habitats of most living cerionids.
Uit de Weerd (2008: 323, fig. 8) explored a vieariance
model in which the proto-Antillean island arc may have
carried North American lineages, including Cerionidae
and Uroeoptidae, northeastward during the early Ter-
tiary until it collided with the Bahamas platform during
the Middle Eocene (Pindell, 1994). H owever, Iturralde-
Vinent (2006) noted that it was not until the Middle
Eocene that permanent lands, required for the develop-
ment of a terrestrial fauna, were present within the
Caribbean realm. An alternative hypothesis, that a mid-
Cenozoic GAARlandia (GAAR = Greater Antilles +
Aves Ridge) land bridge provided the means for coloni-
zation of the Greater Antilles from northwestern South
America (see Iturralde-Vinent, 2006: fig. 6) during a 1-2
Myr interval at the Eocene-Oligoeene boundary has
been proposed (Iturralde-Vinent and MaePhee, 1999,
see Ali, 2012 for a review). Iturralde-Vinent (2006: fig. 13)
provided a detailed account of the patterns and chro-
nology of interconnections among the various land
masses within the Caribbean. This model is applied to
produce a zoogeographic hypothesis for the distribution
of Cerionidae since the Cretaceous (Figure 12), and is
concordant with the absence of cerionids in famaica
and the Lesser Antilles.
As there was never a land connection between Cuba
and the Bahamas, the Great Bahamas Bank must have
been populated initially by propagules from Cuba or
the Cuban archipelago dispersed by hurricanes, with the
Little Bahama Bank colonized by propagules from the
Great Bahama Bank. The role of hurricanes in dispersing
propagules of Cerion among neighboring islands lias
been accepted as stochastic events with a major influ-
ence on biogeographic patterns of cerionids (e.g., Pilsbry,
1907; Mayr and Rosen, 1956; Clench, 1957). Like Cuba,
the islands of the Little and Great Bahama Banks were
sequentially conjoined and separated by changes in sea
level resulting in multiple instances of secondary contact
between populations isolated during interglacial high
Figure 12. 1 lypothesized zoogeographic history ol the fam-
ily Cerionidae based on the age and distribution of fossil and
Recent Tax a (Table 1) and paleogeographie reconstructions
of the Caribbean region from the Cretaceous to Recent
(Iturralde-Vinent, 2006). ABC, Aruba, Bonaire and Curagao;
B, Brasilennea ; C, Cuba; Cl, Cayman Islands; F, Florida;
GBB, Great Bahama Bank; H, Hispaniola; LBB, Little
Bahama Bank; M, Mexistrophia ; PB, Puerto Rico; VI, western
Virgin Islands
stands. Tims, populations that were isolated on small,
individual islands during interglacial periods were able to
expand their ranges and interbreed during glacial periods.
The islands of the Turks and Caicos and many of the
southeastern Bahamian islands never had direct connec-
tion with tire Bahama Banks or Cuba, and remained
isolated by deep channels even during glacial sea level
minima. The cerionid faunas of these islands are thus the
summation of lmrricane-born propagules reaching their
shores from Cuba, Hispaniola, and the Bahamian Banks
and islands.
The Cayman Islands were joined to southeastern
Cuba by shallow banks during the Pliocene and are likely
to have been colonized by cerionids from this region.
Early authors (e.g., Binney, 1851; Pilsbry, 1902, 1907,
1946; Dali, 1905) considered the living Cerion of the
Florida Keys to be derived from populations of north-
ern Cuba. However, a molecular phylogeny based on
partial sequences of tire cytochrome e oxidase I gene
(Harasewyeh et al. 2011:fig. 16) shows them to be most
closely related to samples from Andros Island on the
Great Bahama Bank. The occurrence of Cerion petuchi
new species in the Pleistocene of southeastern Florida
is consistent with a Bahamian origin for the cerionid
fauna of the Florida Keys. However, the origin and
THE NAUTILUS, Vol. 126, No. 4
Page 124
relationships of the upper Oligoeene-lower Miocene
cerionids of the central west coast of Florida are enig-
matic. There was never a direct connection between
GAARlandia and Florida, although this large peninsula
altered current patterns that might have facilitated
rafting of a propagule from the northern portions of
GAARlandia or one of its subsequent archipelagos to
tire west coast of Florida (Iturralde-Vinent, 2006: fig. 12).
Alternatively, a separate offshoot of the Late Cretaceous
Cerion acherontis may have extended eastward along
the northern shores of the Gulf of Mexico, reaching
the coast of central western Florida by the late Oligo-
eene. These are, at best, speculative scenarios for the
origin of a cerionid fauna that does not appear to be
ancestral to the Pleistocene or Recent Cerion of south-
eastern Florida.
The zoogeographic hypothesis depicted in Figure 12
was converted to a phylogenetic tree (Figure 13), with
dates attributed to several nodes based on Iturralde-
Vinent (2006: figs. 10, 13). Dashed lines represent faunas
for which molecular data is lacking. The solid lines rep-
resent branches that are supported by a molecular phy-
logeny of Cerionidae based on amino acid sequences of
a portion of the mitochondrial COI gene (Harasewych
et ah, 2011: fig. 17).
The early history of the Cerionidae was determined
by overland dispersal and vicariance caused by tectonic
events. Mexistrophia was isolated from the South American
cerionids since the Upper Cretaceous by the formation
of a seaway brought about by the displacement of the
proto-Antillean island arc by the eastward movement of
the Caribbean Tectonic Plate from the eastern Pacific.
Cerion sensu stricto , which inhabits the islands of Aruba,
Curasao and Bonaire, was isolated from the remain-
ing Caribbean lineages of Cerion by the break-up of
GAARlandia and continued displacement of the Antil-
lean blocks and terranes northeastward by the advance-
ment of the Caribbean Tectonic Plate. The opening of
the Mona Passage during the lower Miocene separated
the terrestrial faunas of Cuba and Hispaniola from those
of Puerto Rico and the Virgin Islands. Hispaniola was
separated from eastern Cuba during the mid to late Mio-
cene, while the Virgin Islands and Puerto Rico were last
joined during the Pliocene (Iturralde-Vinent 2006: fig. 13).
With the exception of Cuba, the Cerion faunas of these
islands are not diverse.
The absence of a land connection between the islands
of the Bahamas (or the Turks and Caicos Islands) and
the Greater Antilles precluded overland colonization.
Rather, these islands must have initially been populated
by small propagules dispersed from the Greater Antilles
either by rafting or carried by hurricanes. Over time,
Cerion likely reached the more remote islands of the
Bahamas and Florida by a stepping stone pattern. It is
interesting to note that Bartsch's introductions of Cerion
into the Florida Keys generally consisted of large
numbers of individuals (n=500) and the resulting colo-
nies remained unchanged for multiple generations (see
Bartsch [1920] for a summary). However, experimental
<o
c
CD
C
-5
5
CD
S'
CD
5
o
tn
o
8.
CD
CD -5 —
o « <r
_§.£■§
-q cn cd
o o
o
Figure 13. Phylogenetic relationships among living and fos-
sil cerionid faunas derived Irom phylogeographie hypothesis in
Figure 12. Solid lines represent portions of tree that are con-
cordant with a molecular phylogeny of Cerionidae based on
amino acid sequences of a portion of the mitochondria] COI
gene (Harasewych et ill., 2011:fig. 17). Dashed lines represent
faunas for which molecular data is lacking. Dates attributed to
nodes are from Iturralde-Vinent (2006: figs. 10, 13).
introductions of Anolis lizards within the Bahamas using
much smaller propagules (5 or 10 individuals) resulted
in rapid differentiation over a 10-14 year period (Losos
et al. 1997).
Sea-level fluctuations caused by Pleistocene and Holo-
cene glacial / interglacial cycles altered the geography
and faunal diversity of individual islands to varying
degrees. During interglacial periods, terrestrial faunas
would be confined to isolated refugia on hill tops and
mountains that remained above the elevated sea-levels.
Drops in sea level during glacial periods would join the
M.G. Harasewych, 2012
Page 125
islands of such archipelagos into larger platforms per-
mitting previously isolated populations to expand their
ranges and come into contact. Thus, such sea-level
changes amplified diversity by repeatedly isolating and
recombining / hybridizing neighboring populations of
islands and banks defined during glacial periods.
The degree to which tire diversity of a particular bank
or island was amplified during a glacial cycle varied sig-
nificantly, being a function of the number of islands that
were united during glacial periods. The diversity of
distinctive cerionid phenotypes (hybrids) on the Great
Bahama Bank would have been greatly amplified by
multiple cycles of glaciation due to the high number of
islands that were isolated and conjoined. By contrast, the
diversity of isolated islands such as San Salvador in the
Bahamas, which remained isolated throughout the gla-
cial cycles, would have changed little.
This ability of Cerion to hybridize, producing dis-
tinctive phenotypes and alozymes not present in either
parent population, have contributed to the proliferation
of taxonomic names, especially during the late lb11' and
early 20th centuries. Subsequent genetic and morpho-
metric studies have shown that such novel phenotypes
are often ephemeral, surviving on the order centuries
(Woodruff and Gould, 1987) to millennia (Goodfriend
and Gould, 1996). Further, all documented instances
of hybridization have been between geographically and
probably also phylogenetically proximal taxa [e.g..
Hybrid on Bahia Honda Key, Florida = Cerion incanum
(Florida) + Cerion cassablancae (Andros Island, Bahamas);
Hybrid on Newfound Harbor Key, Florida^ Cerion
incanum (Florida) + Cerion viaregis (Andros Island,
Bahamas); Cerion tridentata costellata Fort Jefferson,
Garden Key, Dry Tortugas, Florida = hybrid of Cerion
tridentata (Rincon de Guanabon, Cuba) + Cerion
sculptum (Mariel, Cuba); Fossil hybrid on Great Inagua
= Cerion excelsior (Great Inagua) + Cerion rubicundum
(unspecified, but presumably a nearby island)]. None of
Bartschs experimental attempts to hybridize Cerion from
Curasao, Puerto Rico or Cuba with each odier, or with
Cerion from Florida or the Bahamas were successful.
ACKNOWLEDGMENTS
I am grateful to Dr. Edward J. Petuch for making
available the type series of this new species and for many
helpful discussions of the paleogeography of Florida, the
Bahamas and the Caribbean. Thanks also to Dr. Fred G.
Thompson and John Slapcinsky for their helpful and
constructive reviews. This is Smithsonian Marine Station
at Fort Pierce Contribution Number 895.
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cene genus Brasilennea Maury, 1935. Journal of Mollus-
can Studies 77: 445-447.
Salvador, R.B. and L. R.L. de Simone. 2012. New fossil pulmo-
nate snails from the Paleocene of Itaborai Basin, Brazil
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Thompson, F.G. 2011. Mexistrophia, a new genus of Cerionidae
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Uit de Weerd, D.R. 2008. Delimitation and phylogenetics of
the diverse land-snail family Uroeoptidae (Gastropoda:
Pulmonata) based on 28S rRNA sequence data: a reunion
with Cerion. Journal of Molluscan Studies 74: 317-329.
Woodruff, D.S. 1978. Evolution and adaptive radiation of
Cerion: a remarkably diverse group of West Indian land
snails. Malacologia 17: 223-239.
Woodruff, D.S. and S.J. Gould. 1987. Fifty years of interspe-
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Keys. Evolution 41: 1022-1045.
THE NAUTILUS 126(4): 127-135, 2012
Page 127
A new species of Fusivoluta Martens, 1902
(Gastropoda: Volntidae) from Mozambique
Patrice Bail
2 Square La Fontaine
75016 Paris, FRANCE
[email protected]
Nicolas Puillandre
UMR 7138, Museum National d’Histoire Naturelle
Departement Systematique et Evolution,
43, Rue Cuvier, 75231 Paris, FRANCE
[email protected]
ABSTRACT
During a recent expedition to Mozambique, several specimens
attributed to the genus Fusivoluta von Martens, 1902 were
collected between 1100 and 1820 m deep. Among them, a
new species has been found and is here described and com-
pared with the other East African Fusivoluta . Several live-
collected specimens, belonging to the newly described species
and to Fusivoluta clarkei Rehder, 1969 were sequenced for a
nuclear gene (28S), revealing fixed differences between the
two species.
Additional keywords: Molecular taxonomy, 28S gene, Fusivoluta
profundorum nov. sp.. Gastropoda, Mozambique, Volutidae
INTRODUCTION
The MAIN BAZA expedition took place in April 2009 on
board R/V Vizconde de Eza, a research vessel from the
Secretaria del Mar of the Spanish Ministerio de Medio
Amhiente, Medio Rural y Marino, as part of a joint pro-
ject between Museum National d’Histoire Naturelle
(MNHN) and Instituto Espanol de Oceanografia (IOE).
Forty-six hauls were conducted at depths between
100 and 1820 meters on four transects off Maputo,
Inhambane, Bazaruto, and the mouth of the Zambezi
River. The expedition discovered many new species of
benthic invertebrates, several of which have already been
named (Alf et ah, 2010; Cabezas et ah, 2010; Komai and
Chan, 2010; Richer de Forges, 2010). Among them, a new
bathyal Volutidae has been identified, and is described as
new in this article. Morphological differences clearly sep-
arate it from other Fusivoluta species. Additionally, sev-
eral live-collected specimens of this new species and of
F. clarkei were sequenced for the 28S gene to molecularly
confirm the validity of the two species.
MATERIALS AND METHODS
Study Material: The studied material identified as
Fusivoluta spp. was collected in six different stations
(details are provided below) during the Mainbaza expe-
dition. Three of them (CP3139, CP3141, and CP3154)
provided live material (five specimens in total) that was
used for molecular analyses (Table 1). Two of them were
morphologically identified as Fusivoluta clarkei- the
three others were assigned to the new species. All
vouchers are deposited in the MNHN; specimen data
are also recorded in BOLD and sequences are published
in GenBank (accession numbers are given in Table 1).
Another species, Athleta mozambicana (Rehder, 1972),
also belonging to the family Volutidae, was collected
during the same expedition; it was added to the analysis
as closely related outgroup. Two other Volutidae species
were available in GenBank ( Amoria hunteri - accession
number DQ916584.1 and Alcithoe aillaudonim - acces-
sion number GU440153.1). A distant outgroup belong-
ing to the family Murieidae was used to artificially root
the phylogenetic tree ( Chicoreus subpalmatus, accession
number GU440174.1).
DNA Analysis: Live-collected specimens were pre-
served in alcohol and their DNA was extracted following
the same protocol as in Puillandre et al. (2011). A frag-
ment of the 28S gene was amplified using the primer CT
(ACC CGC TGA ATT TAA GCA T) and D2 (TCC GTG
TTT CAA GAC GGG) (Jovelin and Justine 2001). PCR
reactions were performed in 25 pL final volume,
containing approximately 3 ng template DNA, 1.5 mM
MgCl2, 0.26 nrM of each nucleotide, 0.3 pM of each
primer, 5% DM SO and 0.75 U of Taq Polymerase
(Qbiogene). Amplification products were generated by
an initial denaturation step of 5 min at 94 °C followed
by 35 cycles at 94 °C for 40 s, annealing at 52°C and by
an extension at 72°C for 1 min. The final extension was at
72°C for 5 min. PCR products were purified and
sequenced by a sequencing facility (Eurofins). All genes
were sequenced in both directions for increased accu-
racy. Sequences were manually aligned, and phyloge-
netic analyses were performed using the bayesian
approach implemented in MrBayes (Huelsenbeck et al.
2001). Two parallel analyses were run, consisting each of
eight Markov chains of 5,000,000 generations. All other
parameters were set to default.
Abbreviations: DNMH: Delaware Museum of Natu-
ral History, Greenville, Delaware; MNHN: Museum
National d’PIistoire Naturelle, Paris; RSA: Republic of
South Africa; Stn: station.
RESULTS
Phylogenetic Analysis: The phylogenetic tree
(Figure 1) clearly shows that till the specimens identified
as Fusivoluta belong to a single well-supported clade (Pos-
terior Probability PP = 1). Within this clade, two recipro-
cally monophyletic groups are defined, one corresponding
GU440174.1 Chicoreus subpalmatus
DQ916584.1 Amoria hunter!
0.49
1.00
0.01 subs/site
Figure 1. Phylogenetic tree.
GU440153.1 Alcithoe aillaudorum
IM 2009 9487 Athteta mozambicana
r IM 2009 7340 Fusivoluta clarkei
0 59
L IM 2009 9495 Fusivoluta clarkei
1 00
r IM 2009 9488 Fusivoluta profundorum
1 00
r- IM 2009 9489 Fusivoluta profundorum
0 35
- IM 2009 7392 Fusivoluta profundorum
P. Bail and N. Puillandre, 2012
Page 129
IM_2 00 9- 734 0
IM_2 009-9495
IM_2 009-9489
IM_2009-7392
IM 2009-9488
461
GGGCAGGAGA
GGGCAGGAGA
GGGCAGGAGA
GGGCAGGAGA
GGGCAGGAGA
511
CCGTGTGCAC
CCGTGTGCAC
CCGTGTGCAC
CCGTGTGCAC
CCGTGTGCAC
561
AGAAGC T 1GC
AGAAGC T 1GC
AGAAGC C 1GC
AGAAGC C 1GC
AGAAGC C 1GC
611
CAGCCTC- GC
CAGCCTC-GC
CAGCCTC- GC
CAGCCTC-GC
CAGCCTC-GC
661
T-G-TAGGCC
T-G-TAGGCC
T-G-TAGGCC
T-G-TAGGCC
T-G-TAGGCC
711
CAACCGTGTC
CAACCGTGTC
CAACCGTGTC
CAACCGTGTC
CAACCGTGTC
761
GGGCC C
GGGCC.C
GGGCC T
GGGCC T
GGGCC T
TCTTCAACGA
TCTTCAACGA
TCTTCAACGA
TCTTCAACGA
TCTTCAACGA
TTTCCGCGGG
TTTCCGCGGG
TTTCCGCGGG
TTTCCGCGGG
TTTCCGCGGG
- GAGGAT GGT
- GAGGAT GGT
-GAGGAT GGT
-GAGGAT GGT
-GAGGAT GGT
CTGTCCCGAT
CTGTCCCGAT
CTGTCCCGAT
CTGTCCCGAT
CTGTCCCGAT
GA
GA
GA
TCTCCCG
TCTCCCG
:tctcccg
:tctcccg
TCTCCCG
GCCCC
GCCCC
GCCCC
GCCCC
GCCCC
-GC
-GC
-GC
-GC
-GC
AGGCGCG
AGGCGCG
AGGCGCG
AGGTGCG
AGGCGCG
C-
C-
TT
TT
TT
CCGCCC ■
CCGCCC ■
CCGCCC-
CCGCCC -
CCGCCC -
-GCCC
-GCCC
CGCCC
CGCCC
CGCCC
TGCTGACCGC
TGCTGACCGC
TGCTGACCGC
TGCTGACCGC
TGCTGACCGC
CGGCACAGGG
CGGCACAGGG
CGGCACAGGG
CGGCACAGGG
CGGCACAGGG
GCGC
GCGC
GCGC
GCGC
GCGC
•GGGA
•GGGA
1JGGGGA
T GGGGA
T GGGGA
CAGAGCGCCA
CAGAGCGCCA
CAGAGCGCCA
CAGAGCGCCA
CAGAGCGCCA
CGACCGGTTC
CGACCGGTTC
CGACCGGTTC
CGACCGGTTC
CGACCGGTTC
TCTCCGC
TCTCCGC
TCTCCGC
TCTCCGC
TCTCCGC
G
G
a:
a:
a:
g
CG
G
G
G
TCCTGGGATG
TCCTGGGATG
TCCTGGGATG
TCCTGGGATG
TCCTGGGATG
TTCCTTGGAC
TTCCTTGGAC
TTCCTTGGAC
TTCCTTGGAC
TTCCTTGGAC
TCGGTGGCGA
TCGGTGGCGA
TCGGTGGCGA
TCGGTGGCGA
TCGGTGGCGA
CGCGTGCCGG
CGCGTGCCGG
CGCGTGCCGG
CGCGTGCCGG
CGCGTGCCGG
TTGGGCGGTC
TTGGGCGGTC
TTGGGCGGTC
TTGGGCGGTC
TTGGGCGGTC
TGCGCTGGTA
TGCGCTGGTA
TGCGCTGGTA
TGCGCTGGTA
TGCGCTGGTA
GGG GACCGAGGAG
■GGG GACCGAGGAG
GGG GACCGAGGAG
GGG GACCGAGGAG
GGG GACCGAGGAG
CCGCCGC
CCGCCGC
CCGCCGC T
CCGCCGC
CCGCCGC
1G
1G
1G
jG
jG
TTCGACTGGC
TTCGACTGGC
TTCGACTGGC
TTCGACTGGC
TTCGACTGGC
GGCAAAAACC
GGCAAAAACC
GGCAAAAACC
GGCAAAAACC
GGCAAAAACC
ATCGGTCGGC
ATCGGTCGGC
ATCGGTCGGC
ATCGGTCGGC
ATCGGTCGGC
AGAGAC TGGG
AGAGAC TGGG
AGAGAC TGGG
AGAGAC TGGG
AGAGAC TGGG
AAGGGC
AAGGGC
AAGGGC
AAGGGC
AAGGGC
— :
- :
GTG :
GTG :
GTG :
CCTCCACCTG
CCTCCACCTG
CCTCCACCTG
CCTCCACCTG
CCTCCACCTG
Figure 2. 28S alignment of the two specimens of Fusivoluta clarkei (above) and three specimens of F profundorum (below)
showing the diagnostic characters that discriminate the two species.
Page 130
THE NAUTILUS, Vol. 126, No. 4
to Fusivoluta clarkei, with low statistical support (Posterior
Probability PP = 0.59), the second to Fusivoluta
profundorum with a high statistical support (PP = 1).
Furthermore, the analysis of the 28S alignment reveals
the existence of several fixed mutations and insertions
(10 in total, shown on figure 2) that distinguish the two
species. These results thus confirm the morphological
identifications and support the validity of the two species.
In this partial tree, the relative closeness of Fusivoluta
to Athleta is noticeable. Nevertheless, no conclusion
must be drawn yet, pending the construction of a com-
plete phylogenetic tree of the Volutidae.
SYSTEM ATI CS
Family Volutidae Rafinesque, 1815
Subfamily Callioteetinae Pilsbry and Olsson, 1954.
Genus Fusivoluta Von Martens, 1904.
Type Species: Fusivoluta anomala von Martens, 1902
Fusivoluta profutuiorum new species (Figures 3-21)
Diagnosis: Shell of moderate size, fusiform, with high
spire. Protoconch bulbous, with 2-3 whorls. Teleoconch
sculptured with ax ud ribs. Columella without plaits. Aper-
ture with flared outer lip when adult. Fasciole indistinct.
Siphonal notch absent. Thin periostracum and horny
operculum present.
Description: Adult shell of moderate size for genus
(length 45-55 mm), light, fusiform. Surface dull covered
by thin, adherent, gray periostracum. Protoconch small
(average width: 2 mm), with 2 whorls, first whorl flat-
tened, subsequent very convex. Transition protoeoneh-
teleoconeh sharp. Spire moderately high, forming average
of 48% of total length of shell. Protoconch and abapical
pint of spire usually eroded. Teleoconch of 5 whorls, last
slightly inflated. Suture indented. Sculpture of axial ribs
(14-15 on antepenultimate whorl), blunt or knobbed on
some specimens, prominent onto spire, then becoming
obsolete on half abapical part of penultimate whorl and on
whole last whorl. Spiral sculpture of numerous, regularly
spaced, minute cords covering whole teleoconch, becom-
ing weaker on last whorl, crossed by faint axial lirae giving
cancellate appearance (Figure 40, left). Aperture large,
semi-circular, with outer lip thin and anteriorly flared. Col-
umella slightly sigmoid without plaits. Anterior canal very
short without siphonal notch. Surface of teleoconch and
aperture gray-white, anterior part of columella and inner
edge of outer lip slightly tinged with orange.
Type Material: Holotype MNHN 22816 (=IM-2009-
7392), 49.0x 18.5 mm (Figures 3G5); paratype I, IM-2009-
9489, Stn CP3139, 1195 m, 45.7x17.5 mm (Figures 6-7);
paratype II IM-2009-9488, Stn CP3139, 1195 m,
44.6x16.3 mm (Figures 8-9); paratype III, stn CC3158,
1220-1248 m, 43.3x17.7 mm (Figures 10-11).
Other Material Examined: Station CP3139, 23°35' S,
36°06' E, 1 100- 1200m, 8 specimens: 49.5x17.4 mm,
48.0x17.8 mm, 48.0x17.3 mm, 47.2x18.2 mm, 44.5x
19.8 mm, 42.9 x 18.3 mm, 42.7.0 x 16.0 mm, 43.0 x 17.2 mm.
Station CC3158, 21°46' S, 36° 12' E, 1220-1248 m, 2 spec-
imens: 43.3x17.6 mm (live), 39.6x16.7 mm. Station
CC3156, 21°40/ S, 36°35' E, 1810-1820 m, 1 specimen:
56.3x21.3 mm. Station CP3145, 21°47' S, 36°24' E, 1408-
1421 m, 2 specimens: 55.5x40.0 mm, 50. 8x 18.0 mm.
Type Locality: 23°35/ S, 36°06' E, 1100-1200 m, sta-
tion CP3139.
Etymology: Specific epithet refers to the deep-sea hab-
itat of the new species.
Distribution: Endemic in waters off Mozambique. A
distribution between latitudes 21-23° S, in 1100-1500 m
deep is revealed by the trawling operations, but the southern
and northern limits are still imprecise because of lack of
exploration beyond the explored range (see map. Figure 48).
Table 2. Summary of conchological differences between the five comparable Fusivoluta.
P. Bail and N. Puillandre, 2012
Page 131
Figures 3-21. Fusivoluta profundorum 3-5. Holotype. MNHN 22816, Stn CP 3139, 1194 m, 49.0 mm. 6-7: Paratype I, IM-
2009-9489, Stn CP3139, 45.7 mm. 8-9: Paratype II, IM-2009-9488, Stn CP3139, 44.6 mm. 10-1 1. Paratype III, stn CC3158. 1220-
1248m, 43.3 mm. 12-13. Stn CP3139, 42.9 mm. 14-15. Stn CP3139, 43.0 mm. 16-17. Stn CP3139, 44,5 mm. 18-19. Stn CP3145,
1408-1421 m, 55.5 mm. 20-21. Stn CC3158, 1220-1248 m, 43.3 mm (live taken).
Page 132
THE NAUTILUS, Vol. 126, No. 4
Figures 22-39. Living species of Fusivoluta (except for F. decussata , all the shells illustrated here are from the first author's
collection). 22-23. F. bamardi, offTugela Bank, Natal, RSA, 320 m, 135 mm. 24-25. F. decussata , off Mashbe River, Transkei, 700 m,
47 mm. Natal Museum (Photo D. Herbert). 26-27. F. clarkei , off Maputo, Mozambique, 450-510 m, 122.3 mm. 28-29. F. aff. anomala,
Zanzibar Channel, 850 m, 47.5 mm. 30-31. F anomala. South of Ras Afun, Somalia, 420-450 m, 61.8 mm. 32-33. F pyrrhostoma,
Agulhas Bank, RSA 400 m, 78.1mm. 34-35. F sculpturata, Capetown, no depth given, 27.5 mm. 36-37. F blaizei , off Cape
St Blaize, RSA, circa 250 m, 48.2 mm. 38-39. F wesselsi Tegula Bank, Natal, RSA, 50 m, 24.6 mm.
P. Bail and N. Puillandre, 2012
Page 133
/' i
v‘ ■'
Figure 40. Comparison between the early spires of
Fusivoluta profundorum (left) and F. clarkei (right).
Discussion: Fusivoluta profundorum shows very few
variants, differing only by the strength of the axial ribs,
knobbed specimens occurring mainly below 1400 m.
The genus Fusivoluta comprises ten East African spe-
cies, of which five species, rising in successive ranges all
along the east coast of Africa, must be compared with
F. profundorum-.
Fusivoluta bamardi Rehder, 1969 (Figures 22-23) from
the coast off Natal, is a large shell (up to 145 mm long)
with 7-8 teleoconeh whorls, a high spire, axial ribs
forming small nodules on the three early whorls, crossed
by fine spiral lirae, becoming obsolete on the subsequent
whorls giving a smooth surface. Background colour and
aperture deep beige-orange.
Fusivoluta decussata Barnard, 1959 (Figures 24—25),
from Transkei, is a rare species, with only 5 specimens
known. It is a medium-sized, solid species (approx. 50 mm)
bearing a strong cancellate sculpture, the intersection
between spiral and axial cords forming nodules. The
thickness of this shell, the lack of true axial ribs and its
notched suture contrast with the other members of the
genus. Barnard himself placed this distinctive shell pro-
visionally in Fusivoluta , pending studies of its anatomy.
Fusivoluta clarkei Rehder, 1969 (Figures 26-27) from off
Mozambique, is a parapatrie species occurring all along
the same range as F profundorum, but at lesser depth,
from 450 to 900 m. It is a highly polymorphic species
varying in size, shape and sculpture, according to depth.
The type locality (holotype DNMN 12833) is off Joao
Belo, Gaza district in 440 m. This typical form
(Figures 41-43) the closest to F profundorum , thrives
between 450-650 m. It differs from the latter by a large
size, 75-125 mm long, an elongate-fusiform shape with
high spire and relatively long anterior canal. The 2-3 first
post-nuclear whorls are made angulate by nodulose axial
libs, reduced to weak axial plicae on tire rest of the spire,
absent on the last whorl. Numerous strong spiral cords
extending onto the whole teleoconeh. Columella straight
or weakly concave. Background color yellowish pink, the
subsutural zone slightly darker.
Between 600 to 900 m, F. clarkei tends to be broader,
with more pronounced spiral cords (Figures 44-45). Some
shells are very squat, with reduced axial ribs, and very prom-
inent spiral sculpture, which imparts the shell a rough
Figures 41-47. Variations of Fusivoluta clarkei. 41. North Natal, RSA, 350-400 m, P. Bail collection (ex Meyer collection) 70.0 mm.
42. Mozambique, IM-2009-9495, Stn CC3154, 19°36' S, 36°47' E, 636 m, MNHN, 58. 2mm. 43. Mozambique, IM-2009-7340,
Stn CP3141, 23°33' S, 35°55' E, 684-698 m, MNHN, 72.8 mm. 44. Mozambique, 750-800 m, P. Bail collection, 84.6 mm. 45. Mozambique,
Stn CP3140, 23° 33 S, 36°07S, 890-900m, MNHN, 77.1 mm. 46. Mozambique, Stn CP3148, 21°32' S, 35°47' E, 768-787 m, MNHN,
75.3 mm. 47. South Mozambique, 750-800 m, P Bail collection (ex Meyer collection), 86.8 mm.
Page 134
THE NAUTILUS, Vol. 126, No. 4
Figure 48. Map ol the estimated range of Fusivoluta
profundorum.
cancellate surface, very similar to that of F decussata
(Figures 46-47). No live-taken specimens of this form have
been found until now. From 900 m to 1200 m, no Fusivoluta
have been brought up by tire few dredgings operated by
Mainbaza expedition, suggesting a possible depth gap
between the habitat of F. clarkei and F. profundorum , sub-
ject to confirmation by future expeditions.
Fusivoluta aff. anomala (Figures 28-29), from off
Zanzibar, is a rare species discovered during the 1 980s by
Russian boats operating along the Zanzibar Channel in
deep water (600-800 m). Sharing the same medium size
( circa 45 mm) and light stucture with F. profundorum, it
differs by a turreted protoconch of 3 whorls and a very
tapered spire. Sculpture of strong axial ribs, adapieally
attenuated, numbering 14-15 on the penultimate whorl,
crossed by spiral cords divided into two parts: one sub-
sutural, the other on the anterior third part of the body
whorl, delimiting a smooth central part. Its relationship to
the species below is problematic.
Fusivoluta anomala (Martens, 1902) (Figures 30-31) from
off Somalia, is also a very rare species, known only from the
holotvpe. It differs from F. profundorum by its solid, heavy
structure, a very elongate-fusiform shape with a high nar-
row spire, made angulate by axial nodules (13 on the pen-
ultimate whorl), crossed by raised spiral cords. Both spiral
and axial sculptures extend onto tire whole teleoconch.
Aperture large with the outer lip very flared. Base color
yellowish-beige.
Four species are too different to warrant extensive com-
parison: Fusivoluta pyrrhostoma (Watson, 1882) (Fig-
ures 32-33) from Western Cape differs by a rough
surface and its tilted raised protoconch; Fusivoluta
sculpturata (Tomlin, 1945) (Figures 34-35) from West-
ern Cape is a small species, under 30 mm long, with
heavily sculptured surface; Fusivoluta blaizei (Barnard,
1 959) (Figures 36-37) from Western Cape is a rather small,
elongate, bicoloured shell with very high spire; Fusivoluta
wesselsi Kilburn, 1980 (Figures 38-39) from Natal is a nar-
row, fusiform, small-sized specie, under 25 mm when adult.
ACKNOWLEDGMENTS
Material used in the present paper was collected in
Mozambique in 2009 during the MAINBAZA expedi-
tion, a joint project between MNHN (principal inves-
tigator Philippe Bouchet) and Instituto Espanol de
Oeeanografia (principal investigator Ana Ramos). The
project was part of a group of Mozambique-Madagascar
expeditions (La Planete Revisitee/Our Planet Revisited
http://laplaneterevisitee.org/en) funded by the Total
Foundation, Prince Albert II of Monaco Foundation,
and Stavros Niarchos Foundation, and conducted by
MNHN and Pro-Natura International. The authors are
grateful to Roy Aiken (RSA), for his efficient help and
for giving us the opportunity to study specimens from
his collection; Barbara Buge, Jose Utge, and Audrey
Rivasseau from the MNHN, Paris for processing and
eviration of the molecular collection; Dr. Dai Herbert
from the Natal Museum (RSA) for the photographs
of rare species from the Museum collection; Virginie
Heros, for continued assistance; Allan Limpus from
Bundaberg, Australia, for giving us the opportunity to
study specimens from his collection; and Philippe
Bouchet for helpful comments on the manuscript.
LITERATURE CITED
Barnard, K.H. 1959. Contribution to the Knowledge of South
African Marine Mollusca. Fulgoraria blaizei. Annals of the
South African Museum 45: 28-31, text-fig. 8.
Fraussen K. and J. Rosado. 2011. The Cantharus group
(Gastropoda: Bucc-inidae) on Ahnirante Leite Bank
(Mozambique) with description of two new species and
one new genus. Novapex 12: 73-79.
Huelsenbeck, J.P., F. Ronquist, and B. Hall. 2001. MrBayes:
bayesian inference of phylogeny. Bioinformatics 17:
754-755.
P. Bail and N. Puillandre, 2012
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Jovelin, R. and |. L. |ustine. 2001. Phylogenetic relationships within
the Polyopisthoeotylean monogeneans (Plathyhelminthes)
inferred from partial 28S rDNA sequences. Internationa]
Journal for Parasitology 31: 393-401
Puillandre N., Yu. Kantor, A. Sysoev, A. Couloux, C. Meyer, T.
Rawlings, J.A. Todd, and P. Bouehet. 2011. The dragon
tamed? A molecular phylogeny of the Conoidea (Mollusca,
Gastropoda). Journal of Molluscan Studies 77: 259-272.
Rehder A H. 1969. Two Genera of Deep Water Volutidae from
South Africa. The Veliger 11: 200-209, pis 40-43, figs 1-43.
Tomlin J.R. 1945. Two South African specieds renamed Jour-
nal of Conehology 22: 135.
THE NAUTILUS 126(4): 136-142, 2012
Page 136
A new species of Mysella from Patagonia (Bivalvia: Galeommatoidea)
Cristian Ituarte
Museo Argentino de Ciencias Naturales
Av. Angel Gallardo 470
C1405DJR Buenos Aires, ARGENTINA
[email protected]
Diego G. Zelaya
Departamento Biodiversidad y Biologia Experimental
Facultad de Ciencias Exactas y Naturales
Universidad de Buenos Aires
and
Museo de La Plata
Paseo del Bosque s/n
1900 La Plata
Buenos Aires, ARGENTINA
Juan Pablo Martin
Unidad Academica San Julian
Universidad National de la Patagonia Austral
Colon y Sargento Cabral
9310 Puerto San |ulian
Santa Cruz, ARGENTINA
ABSTRACT
The poorly known diversity ol the genus Mysella in the south-
western Atlantic is improved with the description of a new
species from San Julian Bay, Santa Cruz Province, Argentina.
Mysella patagona new species is characterized by the relative
large, strikingly subquadrate, slightly inequilateral and flat
shell; a cleft hinge plate, with two widely diverging teeth in
the right valve, and a cylindrical internal ligament located
in a deep ligamental pit. From the anatomical point of view,
M. patagona new species is characterized by the presence of
inner and outer demibranchs, with very few interlamellar
junctions. The marked subquadrate shell outline easily distin-
guishes M patagona from any other Mysella species currently
known from the Magellan Region.
Additional keywords: Southwestern Atlantic, Southern
Hemisphere, bivalves
INTRODUCTION
Recent contributions have described and re-described
several small-sized Magellanic bivalves, helping
improve the scanty attention given to this fauna in the
past (Zelaya and Ituarte, 2002, 2004, 2009, 2012). Mem-
bers of the genus Mysella are not an exception. At
present, four species of Mysella are known from
Magellanic waters: Mysella mabillei (Dali, 1908),
Mysella rochebrunei (Dali, 1908) and Mysella sculpta
Soot-Ryen, 1957, all from the Magellan Strait, and
Mysella arthuri (Cooper and Preston, 1910) from
Malvinas (Falkland) Islands.
In the past few years, mysellids from other parts of
the world have been covered in several contributions
(Boyko and Mikkelsen, 2002; Coan and Valentich-Scott,
2012; Passos and Domaneschi, 2006; Passos et al.,
2005). Gofas and Salas (2008), after studying the type
species of Mysella Angas, 1877, Rochefortia Velain,
1878, Rochefortula Finlay, 1927, and Altenaeum
Spaink, 1972, considered Mysella and Rochefortia as
synonyms, instead of two separate genera as done by
other authors (Coan et ah, 2000). Furthermore, Gofas
and Salas (2008) concluded that “Mysella” bident ata
(Montagu, 1803), as well as other European species
previously placed in Mysella, are not congeneric with
Mysella anomala Angas, 1877, the type species of the
genus, and introduced the genus Kurtiella to include
them. Coan and Valentich-Scott (2012) also considered
the Pacific “mysellids” from tropical West America
under Kurtiella.
In the present paper, a new species of Mysella from
southern Patagonia is described and fully illustrated.
MATERIALS AND METHODS
Specimens collected during low tides by sieving (1 mm
mesh) portions of substratum from a tidal flat at San
Julian Bay were immediately fixed in 70% ethanol. Spec-
imens for histology were decalcified by immersion in
Bouin’s fixative for 8 h, rinsed in tap water, dehydrated
in an ethanol series, and embedded in epoxy resin
(Historesin Leiea®). Sections (3.5 pm thick) were stained
with hematoxylin-eosin. Voucher specimens are depos-
ited at Museo Argentino de Ciencias Naturales (MACN)
and Museo de La Plata (MLP). Linear measurements
(shell length [L], shell height [H], and shell width [W])
were taken widi a Zeiss Stemi 2000-C stereomicroscope
with ocular micrometer.
C. Ituarte et al, 2012
Page 137
Figures 1-7. Mysella patagona new species: Shell morphology. 1. Holotype, MACN-In 38865, lateral view of left valve. 2-7.
Paratypes MACN-In 38866. 2. Posterior view. 3. Shell surface with periostracum folds. 4. Inner view of right valve. 5. Inner view of
left valve. 6. Right valve, detail of hinge. 7. Lett valve, detail of hinge. Scale bars: Figure 1 = 1 mm; Figures 2, 4, 5 = 0.5 mm;
Figure 3 = 20 pm; Figures 6, 7 = 300 pm.
Page 138
THE NAUTILUS, Vol. 126, No. 4
SYSTEMATICS
Mysella patagona new species
(Figures 1-19)
Description: Shell thin, relatively large for genus
(maximum observed L = 7 mm), compressed (W/H
ratio = 0.50 ± 0.05, n = 15), nearly equilateral, anterior
half longer (Figures 1, 2). Valves subequal. Shell outline
subquadrate (H/L ratio = 0.70 ± 0.02, n = 15); beaks
small, low, subcentral, opisthogyrate, slightly projecting
above dorsal margin (figures 1, 4, 5). Dorsal margin
about 55% of shell length, slightly flaring anteriorly, with
anterior anti posterior portions similar in length; anterior
half straight or slightly concave, posterior half slightly
curved or straight. Anterior and posterior margins wide
and evenly curved, anterior higher, forming weak angles
at junction with dorsal margin (figures 1, 4, 5). Ventral
margin nearly straight (Figure 1 ), slightly curved at ante-
rior end. Shell surface whitish, sometimes brownish in
larger specimens, sculptured with very low, irregularly
spaced periostracal folds, and slightly marked growth
lines (Figure 3). Prodissoeonch oval, length about 650 pm
(n = 3), smooth.
Hinge plate not solid, cleft just beneath beaks, leaving
a passage for internal ligament (resilium) (Figures 6, 7, 8,
10). Right valve with two small, subequal, peg-like teeth,
anterior longer and slender, posterior somewhat colum-
nar, located on either side of resilium (Figures 4, 6, 8).
Anteriorly to anterior tooth and posteriorly to posterior
tooth, hinge plate margin forms low ridge delimiting with
dorsal shell margin two grooves (Figures 4, 6) that accom-
modate margin of opposite valve (Figures 5, 7). Left valve
edentulous, with dorsal margin moderately thickened on
each side of resilium, forming short lamellae that interlock
with grooves in right valve (Figures 5, 7). Ligamental pit
Figures 8-11. Mysella patagona new species: Ligament and ligamental pit. 8. Detail of right valve hinge showing teeth, ligament
and deep ligamental pit. 9. Ventral view of ligament and ligamental pit. 10. Detail of ligamental pit in left valve. 11. Detail of ligament
with lithodesma. Abbreviations: li, lithodesma; lig, ligament; lp, ligamental pit; lr, ridge of ligamental pit. Scale bars: Figure 8 = 200 pm;
Figures 9, 10 = 100 pm; Figure 11 =50 pm.
C. Ituarte et al, 2012
Page 139
(resilifer) deep. In both valves, two strong, vertical pre-
and post-ligamental ridges delimiting ligamental pit, are
connected laterally with hinge plate (Figures 8, 10). Lig-
ament strong, cylindrical (Figure 9), with a wide, some-
what trapezoidal, calcified area, forming very thin ventral
shield, lithodesma (Figures 11, 12, 19), that extends to
cover partially anterior and posterior portions of liga-
ment (Figures 13, 19).
Anatomy: Mantle border thin, nearly smooth; widely
open in a long inhalant-pedal aperture extending for
about 3/4 of mantle margin length, separated from small
posterior exhalant aperture by short presiphonal suture.
Short, slightly pointed papillae present and restricted to
anterior part of inhalant-pedal aperture and posterior to
presiphonal suture (Figures 14, 15). Ctenidia non-plicate,
inner and outer demibranchs present, outer one reflected
upward (Figure 14), with few interlamellar junctions,
mainly restricted to lower portion of ctenidia (Figure 16).
Ctenidial axis almost parallel to dorsoventral axis. Outer
demibranch small, less than half lengtii of inner
demihranch, with up to 40 filaments approximately paral-
lel to antero-posterior axis (Figure 14). Ascending and
descending lamellae of outer demibranch equally devel-
oped. Inner demibranch with 50 obliquely directed fila-
ments (in larger studied specimen); ascending lamella
approximately half length of descending lamella. Tips of
filaments of ascending lamellae attached to visceral mass
tegument by tissue junctions (Figure 16). Left and right
inner demibranchs fused at posterior end at level of
presiphonal suture (Figure 15). Brooding in gills was
Figures 12-13. Mysella patagona new species: Ligament and
lithodesma. 12. Hinge and ligament from ventral view. 13. Liga-
ment and lithodesma: anteroventral view (upper), ventral view
(centre), posteroventral view (bottom). Abbreviations: at, ante-
rior right tooth; li, lithodesma; lig, ligament; pt, posterior right
tooth. Scale bars: Figure 13 = 0.5 mm; Figure 14 = 0.2 mm.
observed in one specimen (ca. 7 mm length) collected
in April (Austral Autumn). Labial palps with 10 sorting
ridges. Ventral surface of presiphonal suture ciliated,
forming a groove when valves are closed (Figures 15, 18).
Foot large, compressed laterally, with well developed
posterior heel (Figure 14). Anterior part of foot heavily
ciliated. Transverse section of anterior and posterior
adductor muscles ovate, subequal, anterior adductor with
posterior indentation (Figure 14). Anterior pedal retractor
strong. Prominent byssal gland opening in a ciliated byssal
groove (Figure 17).
Habitat: Mysella patagona new species lives in lower
intertidal and shallow subtidal mud-sandy beaches:
46-68% mud (silt + clay), 31-53% fine sand, and 2.5-
4.0% organic matter content. Mysella patagona is a
conspicuous species of the infaunal assemblage domi-
nated by the bivalve Darina solenoides (King, 1832),
the amphipod Ampelisca sp., and the polychaetes
Scolecolepides uncinatus Blake, 1983 and Eteone sculpta
Ehlers, 1897. Mysella patagona showed abundances of
up to 30,000 ind.m ~, representing in some areas the
dominant species of the infaunal assemblage (J.P Martin,
unpublished data).
Type Locality: 49°17'30" S, 67°43'22" W, northwest
of San Julian Bay, Santa Cruz Province, Argentina.
Material Examined: Holotype (MACN-In 38865)
and 15 paratypes from the type locality (4 MACN-In
38866, mounted for SEM; 5 MACN-In 38866, ethanol
preserved specimens; 5 MLP 13635, dry specimens).
Etymology: The species name, a noun in apposition,
honors to Patagones, the name given by Spaniard
explorers to the ancient people who inhabited vast
regions of the Argentine southern littoral.
Distribution: Only known from the type locality.
Remarks: Mysella patagona new species is similar to
Mt/sella rochebrunei (Dali, 1908), differing by its larger
size and by having a markedly subquadrangular shell
outline with straight, nearly horizontal (not sloping)
anterior and posterior parts of dorsal margin, ventral
margin nearly straight, slightly curve at anterior end, and
more prominent beaks. The larger size and subquadrate
shell outline easily separate Mysella patagona new spe-
cies from the other mysellid species reported from the
Magellan Region: Mysella mabillei (Dali, 1908) from
eastern Magellan Strait, Mysella artkuri (Cooper and
Preston, 1910) from the Malvinas (Falkland) Islands,
and Mysella scidpta Soot-Ryen, 1957, from the Magellan
Strait, as well as from the Antarctic species Mysella
charcoti (Lamy, 1906) and Mysella narchii Passos
and Domaneschi, 2006; all of these have a strikingly
trigonal-ovate and markedly inequilateral shells. Fur-
thermore, Mysella arthuri has only one, the anterior,
cardinal tooth in the right valve.
Kurtiella Gofas and Salas, 2008, was based on “the com-
plete regression of the liinge plate beneath the umbones”,
Page 140
THE NAUTILUS, Vol. 126, No. 4
id
r
pam
Figure 14. Mysella patagona new species: Gross anatomy, left lateral view, left shell and mantle removed. Abbreviations:
aam, anterior adductor muscle; ap, anterior pedal retractor; ex, exhalant aperture; f, foot; id, inner demibanch; lp, labial palp;
mb, mantle border; od, outer demibranch; p, papillae; pam, posterior adductor muscle; r, rectum.
a condition not present in Mysella anomala , the type of
the genus in which the solid hinge plate underlies a not
outstanding ligamental pit. In Kurtiella species the inter-
nal ligament is connected to the shell. The latter condi-
tion is more similar to that found in Mysella patagona,
in which, however, the space below the beaks is occupied
by a strong resilifer which is flanked by two strong
lateral ridges. This condition is not seen in the type of
Mysella nor in Kurtiella (see for example the hinge of
K bidentata (Montagu, 1803) illustrated by Gofas and
Salas (2008; fig. 7 E, F). Anterior and posterior teeth of
right valve are subequal in M. patagona as in Kurtiella
species, but while in Kurtiella only the inner
demibranchs are present, M. patagona new species has
both inner and outer demibranchs. For the above rea-
sons, the generic location of the new species is difficult,
and we prefer to be conservative, placing Mysella
patagona provisionally under Mysella.
The presence of a lithodesma associated with the
internal ligament, a structure well known for many,
but not all anomalodesmatans (Harper et ah, 2006),
and reported by Morton (1980) in some montacutids
as Mysella (Montacutona) compacta (Gould, 1861)
and Mysella (Montacutona) olivacea (Habe, 1959), was
also reported by Gofas and Salas (2008) as one of
the diagnostic characters of Kurtiella (originally placed
in Montacutidae).
As recently discussed by Boyko and Mikkelsen (2008),
the family-level relationships in the Galeommatoidea are
not clear, nor diey have been revised for relationships at
the family and genus levels. Montacutidae is the family to
which Mysella was traditionally assigned. Mysella patagona
new species agrees in general terms with montacutid
anatomy, differing in having both inner and outer
demibranchs, an important character at the family level.
Coan et al. (2000) and Coan and Valentich- Scott (2012),
not finding consistent characters to separate the
Erycinidae, Kellidae and Montacutidae, considered them
under a single family, Lasaeidae, a criterion also followed
by Bieler et al. (2010).
C. Ituarte et al, 2012
Page 141
Figures 15-19. Mysella patagona new species. Histology. 15. Transverse section showing posterior fusion of descending lamellae
of inner demibranch and presiphonal suture. 16. Detail of inner demibranch. 17. Sagittal section of foot. IS. Detail of posterior
mantle fusion. 19. Sagittal section with detail of ligament. Abbreviations: aam, anterior adductor muscle; al, ascending lamella
of inner demibranch; ap, anterior pedal retractor; bg, byssus gland; eg, ciliated groove; dg, digestive gland; dl, descending lamella
of inner demibranch; f, foot; g, gonad ilj, interlamellar junction; li, organic matrix of lithodesma; lig, fibrous portion of ligament;
lp, labial palp; m, mantle; mb, mantle border; p, periostraeum pg, pedal ganglion; pss, presiphonal suture; rmb, retractor muscle
of mantle border; vg, visceral ganglia. Scale bars: Figure 15 = 200 pm; Figures 16, 18 = 50 pm; Figures 17, 19 = 100pm.
ACKNOWLEDGMENTS
This work was partly supported by grant PICT20 10-0730
from ANPCyT - FONCyTand PIP1640 from CONICET
to C.I. C.l and D. Z. are members of CONICET.
LITERATURE CITED
Boyko, C.B. and P. M. Mikkelsen. 2002. Anatomy and biology of
Mijsella pedroana (Mollusca; Bivalvia: Galeommatoidea) and
it commensal relationship with Blepharipoda occidentalis
(Crustracea: Anomura; Albuneidae). Zoologiseher Anzeiger
241; 149-160.
Bieler, R., J.G. Carter and E.V. Coan. 2010. Classification
of Bivalve families. Pp. 113-133, in: Bouehet, P and
J.P Rocroi, (2010). Nomenclator of Bivalve Families.
Malacologia 52: 1-184.
Coan, E.V, P.V. Scott and F. R. Bernard. 2000. Bivalve seashells
of western North America: marine bivalve mollusks from
Arctic Alaska to Baja California. Santa Barbara Museum
of Natural History Monographs 2: 1-764.
Coan E.V. and P.V. Valentieh Scott. 2012. Bivalve seashells
of tropical West America: marine bivalve mollusks from
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THE NAUTILUS, Vol. 126, No. 4
Baja California to Northern Peru. Santa Barbara Museum
of Natural History Monographs, 6, 1258 pp.
Passos, F. D. and O. Domaneschi. 2006. A new species of
Mysella Angas, 1877 (Bivalvia: Galeommatoidea) from
Admiralty Bay, King George Island, South Shetlands,
Antarctica, with data on its biology and functional anatomy.
Polar Biology 29: 389-398.
Passos F.D., O. Domaneschi and A.F. Sartori. 2005. Biology and
functional morphology of the pallia! organs of the Antarctic
bivalve Mysella charcoti (Lamy, 1906) (Galeommatoidea:
Lasaeidae). Polar Biology 28: 372-380.
Gofas, S. and C. Salas. 2008. A review of European " Mysella ”
species (Bivalvia, Montaeutidae), with description of Kiirtiella
new genus. Journal of Molluscan Studies 74: 119-135.
Harper, E.M., H Dreyer, and G. Steiner. 2006. Reconstructing
the Anomalodesmata (Mollusca: Bivalvia): moqrhology
and molecules. Zoological Journal of the Linnean Society
148: 395-420.
Morton B. 1980. Some aspects of the biology and functional
morphology (including the presence of a ligamental
lithodesma) of Montacutona compacta and M. olivacea
(Bivalvia: Leptonacea) associated with coelenterates in
Hong Kong. Journal of Zoology 192: 431-455.
Zelaya, D.G. and C. Ituarte. 2002. The identity of Waldo
parasiticus (Dali, 1876) and description of Waldo trapezialis
new species (Bivalvia, Galeommatoidea). The Nautilus
116: 109-117.
Zelaya, D.G. and C. Ituarte. 2004. The genus Neolepton
Monterosato, 1875 in southern South America (Bivalvia:
Neolepton idae). Journal of Molluscan Studies 70: 123-137.
Zelaya, D.G. and C. Ituarte. 2009. A new species of
Pseudokellya Pelseneer, 1903 from the Southern Ocean
(Bivalvia: Cyamiidae). The Nautilus 123: 1-8.
Zelaya, D.G. and C. Ituarte. 2012. Tellimya new species: First
record of Tellimya Brown, 1827 in South America
(Bivalvia: Montaeutidae). Malaeologia 55: 173-182.
THE NAUTILUS 126(4):143-147, 2012
Page 143
The dating of modern books and their included taxa
Richard E. Petit
806 Saint Charles Road
North Myrtle Beach, SC 29582 USA
[email protected]
ABSTRACT
Disparities in dating the molluscan taxa introduced in three books
by E | Petuch are discussed and reconciled. Also addressed is
the potential problem in using the copyright date or the Library
of Congress Control Number for dating nomenclatural acts
that are published in books.
INTRODUCTION
The publication dates of three books written by E. ].
Petuch, either as sole or senior author, have been vari-
ously treated in the literature. As more than one hundred
new molluscan taxa are introduced in the three works,
these have been investigated in an attempt to determine
their correct date ol publication. These books are listed
and treated individually herein.
Information developed about the dating of modern
books is included as it affects numerous works. It may
surprise some people that the copyright year date in a
book is not necessarily the year of publication. An attempt
is made herein to describe the various ways through
which publication dates are determined, correctly and
incorrectly. Unfortunately, there is not a simple way to
precisely determine publication dates.
A fourth book by the same author, containing no
new nomina, is also treated as it has a misleading copy-
right date.
Abbreviations used in the text are: CIP - Cataloging-
in-Publication Data; ICZN - International Code of Zoo-
logical Nomenclature; ISBN - International Standard
Book Number; LCC - Library of Congress Classifica-
tion; LCCN - Library of Congress Control Number;
LOC - Library of Congress; MARC - Machine-Readable
Cataloging record.
BOOK PRINTING AND PUBLICATION DATA
Information on the production and publication dates of
books usually appears on what was once called a colo-
phon page located at the end of the book. I terns normally
listed are the International Standard Book Number
(ISBN), Library of Congress Control Number (LCCN),
Library of Congress Classification (LCC), and other
items comprising Cataloging-in-Publieation Data (CIP).
In modern works such data is ou the verso of the title
page. In some books this information appears in several
areas of the page, usually headed by the copyright date
which is not part of the CIP. In most books, especially
those published by large publishing firms, the CIP appears
in a block headed “Library of Congress Cataloging in
Publication Data” while in others it appears without a
heading. This LOC data is utilized by most librarians as
the basis for their library records and catalogs.
It is difficult to believe that cataloguing a book could
be so involved or esoteric until one makes an effort to
understand the terms and numbers involved. Only the
barest essentials will be mentioned here. In-depth
knowledge of the system and its ramifications would
require years of study.
Usually the first number to appear in the LOC block is
the ISBN. The ISBN consists of 10 or 13 digits, with
some books having both. These numbers are sold to
publishers who then assign them to their books. This is
a monopoly, with single numbers now costing $125.
However, blocks of 10 can be purchased for $250 and
100 numbers cost only $575. The ISBN is intended for the
benefit of booksellers, not to establish the date of publica-
tion. In order for a number to be properly placed into use
after it has been assigned to a book, the publisher must
report such use to R. R. Bowker, the database of record
for the ISBN Agency. Bowker’s Books in Print can be
searched only by subscription and should not be confused
with an online ISBN search which can be made at http://
www.isbnsearch.org. Such search will yield a description
of the book and links to booksellers who have it available.
The LCC (what many would refer to as the “Call
Number”) starts with a letter or letters (for science books
it is Q) and does not involve date. Following the LCC,
usually separated by a wide space or on a separate line, is
the LCCN. This is a number of seven to ten digits, the
first two or four indicating the year cataloging began and
a number assigned, which may, or may not, coincide with
the year of publication. This data is taken neither from
the book nor from the ISBN form, but from a separate
CIP form submitted by the publisher. Not all books
Page 144
THE NAUTILUS, Vol. 126, No. 4
having a ISBN also have a LCC or a LCCN (e.g., Petuch
and Sargent 2012).
A search for a hook in the LOC on-line catalog returns
a detailed listing similar to most library listings. At the
top there are several additional options including “Full
Record” and “MARC Tags.” Clicking on the latter brings
up a detailed history of the cataloging of the item but
the tag numbers are not explained. However, they are
explained in large part on the internet. These tags are
sometimes important as one tag indicates the last date a
catalog entry was accessed and checked or emended
(although the changes are not noted) and which is usu-
ally the date that an actual book was inspected and not
just a form. However, due to back logs a book might wait
a considerable time before being checked, if at all. More
about MARC can be found at http://www.loc.gov/marc/
umb/umOl to06.html.
The LOC data block also contains a date, usually pre-
ceded by a “c” in the form cl988. The “c” is used in lieu
of the copyright symbol [©] and is taken from the CIP
form. It might as well be considered an abbreviation for
the Latin circa which would be appropriate, especially
considering that it is sometimes used for books that are
not registered at the Copyright Office (e.g. Petuch
2007). The LOC cannot place an absolute publication
date in their data unless the book is actually imprinted
“Published (date).” As a LOC specialist wrote me, “Pub-
lishers do not care about a publication date per se.”
Not included in the LOC block of data is the copyright
date, usually in the form of: © 2010 Publishers Name.
This date is unfortunately meaningless for determining
the date of publication. In practice a book with a © 2010
date could have been published in 2009 or 2010, or possi-
bly even in some other year. There will be more about this
under the discussion of the dating of Cenozoic Seas.
Usually on the colophon page, but not in the LOC
data block, are one or two strings of numbers, placed
there by the printer, which are referred to as the Printer’s
Key. An example is: 93 92 91 90 89 88 5 4 3 2 1. The
lowest number in the double digit series is the year of
printing and the lowest number of the single digits is
the printing number. In this example it is a 1988 1st print-
ing. There is no “rule” as to how many digits are used in
each series, or whether they should be in ascending or
descending order. These data, inserted by the printer,
have no real bearing on publication date.
In the following section book titles are in bold type
and the block of text immediately below the book title is
the LOC Catalog Data.
BOOKS DATED
Field guide to the ecphoras. [E ] Petuch]
LOC catalog data (in part): LCCN 88035283; Personal name:
Petuch, Edward ].; Published/Created: Charlottesville,
Va., U.S.A.: Coastal Education & Research Foundation,
cl988; Description: 140 p.: ill.: 22 cm.; ISBN 0938415034;
LCC QE809.M85 P48 1988.
Not included in the pagination are the title and con-
tents pages [iv] and the front and rear covers which bear
figures with captions within [iv].
In addition to the LOC data tins book bears the
imprint “Copyright ' 1988 by the Coastal Education &
Research Foundation.” However, the book is not regis-
tered with die U.S. Copyright Office as shown by a
search at http://www.copyright.gov/records/.
The Printer's Key in this book is 94 93 92 91 89 88 5 4
3 2 1. The lowest of the first six numbers indicates the
year of printing as 1988. Tins indication is invalid for
dating as the printing process may not have actually been
completed in 1988 as scheduled. The other series of
numbers shows that this is the 1st edition.
The MARC tags for this book show that the record
was started on 19 December 1988 (MARC tag line
008) from CIP data (tag line 906). MARC tag line 005
shows that the last transaction was on 20 June 1989,
considered to be the date that a book was checked
against the record.
At the bottom of the verso of the Contents page is
the printed statement “Species validation date: 15 Feb-
ruary 1989." There is no provision in any edition of the
International Code of Zoological Nomenclature (ICZN)
for a “validation date” to be determined in such fashion
as all of the new taxa in any work must date from the
actual publication date. This is a matter of concern as
three genera and 22 species names appear as new in
the Field Guide.
The ISBN is of no value in establishing a date as will
he explained in a Discussion at the end of this paper. In a
futile search for definitive dating it has been noted that
at least two book dealers list the publication date as
January 1, 1989 with one even adding “Sun.” to that date.
The origin of this date has not been established.
The third line of the Books in Print entry for Field
guide to the ecphoras is “Publication Date: United States:
01 Oct 1989". This cannot be correct as a copy at hand
was received on 27 March 1989. Further down in the
listing is a line for "Date last updated”, stated to be
“16 Aug 2008”. A search of Amazon.com by ISBN
returns an entry for die book listing it as “(Jan 1989)"
but it is not in stock. The source of this date cannot be
determined. A search for this book at www.isbnseareh.
org by ISBN returns ‘"Warning: Invalid argument sup-
plied ...” with what appears to be a link but is non-
functional even when copied and pasted.
The publication date in most library catalogues mimics
the LOC listing, with date as el988. It is dated 1988 by
Donovan (1991: 52), Ward (1992: 143; 2008: 393), and
on the U. S. Geological Survey National Geologic Map
Database (http://ngmdb.usgs.gov/Prodese/proddesc_9 1675
.htm). It was also dated as 1988 by Petuch (1988: 80) in
Literature Cited. However, the date 1989 has been con-
sistently used by Petuch in his later works.
This is the last of four books, all by Petuch, published
by Coastal Education & Research Foundation, Inc. (CERF).
Advertisements for their books were placed in 1988 and
1989 issues of the quarterly American Conchologist.
R.E. Petit, 2012
Page 145
As late as the June 1989 issue the Field Guide was not
among those listed. It was first advertised in the Sep-
tember 1989 issue where it was included with the other
three. Dr. Charles W. Finkl, President of CERF, advises
that records that would show publication dates no longer
exist. The Journal of Coastal Research, published by
CERF, did not list it as a publication until Volume 5
Number 2 [Spring 1989] where it is listed as “(1989)”.
As no record of this work being available in 1988 has
been located, and the only printed date appearing therein
aside from the copyright date is the “validation date” of 15
February 1989, that date must be accepted as the publi-
cation date of this book. The copy at hand was received on
27 March 1989 but it is not known if it was ordered
prepublication or after it was known to be available. No
prepublication advertisements have been located.
It should be noted that “ecphoras” in the title is nei-
ther italicized nor capitalized, being used as a vernacular
as explained by Petueh in his Introduction.
Cenozoic Seas. The view from eastern
North America. [E. J. Petueh]
LOC catalog data (in part): LCCN 2003065367; Personal
name: Petueh, Edward J.; Published/Created: Boca
Raton, Fla.: CRC Press, c2004; Description 308 p.: ill.:
26 cm.; ISBN 0849316324; LCC QE39.5.P25 P45 2004,
The above LOC data is taken from the LOC website.
The LCC printed in the LOC data block in the book
differs by ending in 2003.
Not included are introductory pages [xii].
Although the book bears an imprinted copyright date of
2004, the publisher’s web site showed it as having been
published on 29 December 2003. Contact with the pub-
lisher brought a communication, dated 21 January 2004,
from John Sulzyeld, Senior Editor of CRC Press, advising
that “12/29/03 was the actual date of publication.”
An explanation of the difference in dates was offered
to Dr. M. G. Harasewych by Mr. Sulzycki in June 2012
when he confirmed the 2003 publication date. He wrote
that: “This copyright issue has come to be because of the
modern day book publishing convention of giving books
the following year copyright date if they publish July 1 or
after. In the case of Cenozoic Seas, it was an unfortunate
oversight that the copyright date was not the year of
publication. We do that for books where we are aware
that they include new species and higher taxa. It is our
fault as a publisher and we bear final responsibility for
this mistake with Cenozoic Seas. The copyright date
should have been 2003 since the book published in
December, 2003.” The Copyright Date is listed as 29
December 2003 by the U.S. Copyright Office.
Notice of the 2003 publication date was sent by email
to Dr. Petueh by this author on the same day that it was
received, 21 January 2004, suggesting that he have a
small notice printed in several journals noting that his
new taxa (12 genera, 38 species) should be dated 2003
instead of 2004. Such action was never taken and the
new taxa have been consistently dated as 2004 by all later
workers, including Petueh.
Under ICZN Article 21.4 the new taxa in this work
must date from 29 December 2003.
Molluscan paleontology of the Chesapeake Miocene
[E. J. Petueh and M. Drolshagen]
LOC catalog data (in part): LCCN 200901 1500; Personal
name: Petueh, Edward J.; Published/Created: Boca
Raton: CRC Press, c2010; Related names: Drolshagen,
Mardie, 1952-; Description: xvii, 160 p.: ill, maps; 27 cm.
+ I CD-ROM (4 3/4 in.); ISBN 9781439811597 and
1439811598; LCC QE801.P439 2010.
The only date printed in this book is on the copyright/
colophon page where the copyright date is 2010. However,
the LCCN 2009011500 indicates that cataloging began in
2009. There are no other dates. The publisher’s web site
lists the publication date as 19 August 2009. The copy at
hand was sliipped from Amazon on 22 August 2009.
Other evidence for publication in 2009 exists in a news-
paper article (Chowning, 2009) that describes the book,
mentions people for whom species were named, and
includes a photograph of the book on a bench with fossils.
This book is not registered with the U.S. Copy-
right Office.
Despite having been published in 2009, there is no
internal evidence in the book for that date. All American
libraries checked in a survey of on-line library catalogues
have the publication date listed as “c2010”. The only
library catalogue giving a date of 2009 is that of the
Natural History Museum, London.
David Williamson of the Library of Congress has
kindly provided (personal communication, 3 August
2012) an explanation of the post-dated copyright date:
“Regarding the c2010 thing, towards the fall publishers
start putting the copyright date for the next year. It’s a tax
tiling as I understand it. . . . Since there is nothing that
actually says published 2009, we have to use c2010 as the
inferred date of publication.”
The only citation of this recently published work
found is in a review by Allmon (201 1) where it is dated
as 2010. No record of any of the taxa in this recently
published book being cited has been found.
It is indisputable that the publication date is 19 August
2009 and the two new genera and 28 new species must
be dated 2009.
The Geology of the Florida Keys and Everglades
[E. J. Petueh]
Not listed by LOC or Copyright Office. Published by
Thomson, Mason, Ohio, viii + 84 p.
The copyright date of this book is 2008 but the
Printer’s Key indicates a 2007 printing date. This is con-
firmed by an ISBN search that returns December 2007
as the date of publication. The book bears ISBN
9781426630613 and 1426630611.
Page 146
THE NAUTILUS, Vol. 126, No. 4
This book is not registered with the U.S. Copy-
right Office.
This attractive small book has no new taxa but has
excellent photographs of lower Florida habitats and envi-
ronments as well as dozens of full page color plates of
fossil mollusks. It is unusual to depict fossil mollusks in
color but the result is startling when compared with
previously published black and white figures of the same
specimens. The color and shading create a pleasing
effect that is almost three-dimensional.
A request to Thomson for information about the
actual publication date remains unanswered. As no new
taxa are involved the date is rather immaterial, but for
citation purposes it is here considered that the publica-
tion date is December 2007.
DISCUSSION
No mention has been found in the malaeologieal litera-
ture of the fact that the copyright date of a book is not
always the date of publication.
It would be reasonable to assume that a publication
date used by a bookseller in combination with an ISBN is
the date reported by the publisher. It must be further
assumed that the publisher reported the true date of
publication. However, it should be remembered that
notification to Bowkers of assignment of a number to a
book, submission of a CIP form to the LOC, and the
submission of a form to the Copyright Office, all often
occur prior to actual publication. It is the publishers
data, as reported on a CIP form, which is normally used
by the LOC to generate the LOC data used by librarians.
As demonstrated by the confusing listings cited above,
these assumptions should be made only when no other
evidence is available.
The Copyright Dates listed by the U.S. Copyright
Office would intuitively be the same as the publication
dates. Unfortunately, they are no more accurate than
the © or e dates imprinted within the books. In fact,
the U.S. Copyright Office data may include errors of a
different type. While checking the Copyright Catalog
for data on the books discussed herein, another Petuch
work came to attention. Published in 1997 with the title
Coastal Paleoceanography of eastern North America
(Miocene - Pleistocene), it is registered, with a copyright
date of 1996 from a document dated “27Sep95”, under
the title Paleoceanography of eastern North America
( Miocene to Holocene). It would appear that the title
was changed and the publisher did not notify the Copy-
right Office.
The practice of describing new taxa and other nomen-
clatural acts in commercial books has long been frowned
upon by many in the malaeologieal community, espe-
cially when new taxa appear in an appendix to a basically
non-taxonomie work. There are numerous reasons for
such objection, among them being lesser (or absent)
peer review in the book business for systematic portions
of a book and the danger of reissuing later printings
without proper understanding on the part of readers of
potential nomenclatural implications.
This note is not intended to address the problems
with the definition of various terms (“date of publication
specified”; “existence as a published work”; etc.) within
Chapter 5 of the International Code of Zoological
Nomenclature. The works discussed herein must be
dealt with under Article 21.4 which treats cases where
“the date of publication in a work is found to be incor-
rect." The ambiguous portion of that Article, “in exis-
tence as a published work”, is much too subjective for
comment here as “published work” does not mean the
same to everyone. A publisher’s statement of publication
date, by letter or on web site, when available, must be
interpreted as being equal to that date being printed in
the book.
The problem of misdating or non-dating in malaeolog-
ieal literature is not new as many dating problems are
found in the older literature. One example usually men-
tioned in this context is that of Anton ( 1838), where the
title page is imprinted 1839 but a printer’s date in the
back is 1838. This was clarified when an 1838 review of
the work was located (Cernohorsky, 1978). In the mod-
ern literature, this problem usually exists only in books
published by firms that are not accustomed to publishing
books that include new zoological taxa. In such cases,
there is no proper understanding of the implications of
nomenclatural priority and the constraints presented by
an incorrect or nonexistent publication date. This prob-
lem is not restricted to small publishers but afflicts such
larger firms as CBC and Princeton University Press. The
latter firm published the important work “Seas hells of
southern Florida . . . Bivalves ” by Mikkelsen and Bieler,
with “Copyright © 2008” and complete Library of Con-
gress Cataloging-in-Publication Data. That data included
a LCCN indicating a 2007 date. The actual publication
date was November 2007, as was pointed out by the
authors (who have a policy of not introducing new taxa
in such book publications).
The book industry, including printers, publishers, dis-
tributors and sellers have no real interest in accurate
publication dates. David Williamson of the LOC wrote
(personal communication, 3 August 2012) that “Pub-
lishers do not care about a publication date per se. They
are interested in the date it is released to the public for
sale (which is not given in the book) and the date of the
copyright in case of litigation. Other than that there is
very little interest in a true publication date.” Even pub-
lishers can’t agree on what that means, I work with them
on a data committee and they go around and around on
what the definition is — date of printing? Date of ship-
ping to the store? Date it goes on sale?” This question is
incompletely addressed in ICZN Chapter 5.
An entirely different problem with determining date
of publication is found in a work by Petuch and Sargent
(2012), where the only printed date is demonstrably a
year earlier than the actual date of publication, and has
neither a LCCN nor ISBN. That dating problem was
addressed by Petit (2012).
R.E. Petit, 2012
Page 147
For works already published, we will have to continue
to do the best we can to establish publication dates where
none is stated. For the future, it is incumbent upon those
introducing new taxa or nomenclatural acts to be sure
that they are publishing in a journal or series that is
known to place the publication date in the work itself. If
an author’s work is to be published by a firm not accus-
tomed to handling scientific books and papers involving
time-sensitive items, the author should ensure that the
publication date is shown as such in the work.
CONCLUSIONS
It is here considered that the correct dates of publication
for the first three works discussed are: Field guide to the
ecphoras , 1989, and not 1988 as sometimes stated; Ceno-
zoic Seas , 2003, and not 2004 as shown in the CIP and in
subsequent literature; Molluscan paleontology of the
Chesapeake Miocene , 2009, and not the copyright date
of 2010. It follows, of course, that these dates must apply
to the considerable number of new names introduced in
them. The fourth work. The geology of the Florida Keys
and Everglades , without new taxa, dates from 2007 and
not the copyright date of 2008.
ACKNOWLEDGMENTS
Numerous people rendered assistance and advice in
making this paper possible. Chief among them are
Charles W. Einkl, CERF, West Palm Reach, Florida;
Jennifer J. Harbster and David Williamson, Library of
Congress, Washington, DC; John Sulzyeld, CRC Press,
Roca Raton, Florida; M.G. Harasewych, Smithsonian
Institution, Washington, DC; Gary Rosenberg, Academy
of Natural Sciences, Philadelphia; Rudiger Rieler and
Christine Giannoni, The Field Museum, Chicago; Bruce
D. Neville, College Station, Texas; L. S. Chowning,
Urbanna, Virginia.
Although this paper could not have been written
without the assistance and input of those acknowledged,
any errors, misstatements or misinterpretations of the
various numbering and cataloging systems must remain
this author’s responsibility.
LITERATURE CITED
Allmon, W.D. 201 1. Review of: Molluscan Paleontology of the
Chesapeake Miocene, by Edward | Petuch and Mardie
Drolshagen, 2010. Palaios, DOI: 10.21 10/palo.2011.BR64.
Anton, H.E. 1838. Verzeichniss der Conehylien welche sich in
der Sammlung von Hermann Eduard Anton befinden.
Eduard Anton, Halle, xvi + 110 pp. [Imprinted 1839 but
published in 1838. See Cernohorsky 1978.]
Books in Print. 2012. Books in print. [http://www
.booksinprint2.com] As this is a subscription database,
only the “root” URL is shown.
Cernohorsky, W.O. 1978. The date of publication of Anton’s
“Verzeichniss der Conehylien". The Veliger 20: 299.
Chowning, L.S. 2009. Local shells could be over 20 million
years old. Southside Sentinel (Urbanna, VA) for Novem-
ber 19, 2009, p. 2.
Donovan, S.K. 1991 Review: “New Caribbean Molluscan
Faunas”, “Field Guide to the Ecphoras,” and “Neogene
History' of Tropical American Mollusks,” by Edward J.
Petuch. Journal of the Geological Society of Jamaica 28:
52-54.
International Commission on Zoological Nomenclature 1999.
International Code of Zoological Nomenclature. Fourth
edition. I.T.Z.N., London, xxix 4- 306 pp.
Mikkelsen, PM. and R. Bieler 2007 Seashells of southern
Florida: living marine mollusks ol the Florida Keys and
adjacent regions. Bivalves, viii 4- 503 pp. Princeton Uni-
versity Press, Princeton and Oxford. [Copyright date 2008;
published November 2007]
Petit, R.E. 2012. A review of Rare and Unusual Shells of
Southern Florida (Mainland, Florida Keys, Dry Tortugas)
by Edward J. Petuch and Dennis M. Sargent. Conchologia
Ingrata 8: 1-9.
Petuch, E.J. 1988. New gastropods from the Maryland Mio-
cene. Bulletin of Paleomalacology 1: 69-80.
Petuch, E.J. 1989. Field guide to the ecphoras. The Coastal
Education & Research Foundation, Inc., Charlottesville,
Virginia. 140 pp. [imprinted “Species validation date: 15
February 1989”] [listed in 1988: 80 as “ 1988" and “144 pp.]
[Review: Donovan, 1991]
Petuch, E | 1997. Coastal Paleoceanography of eastern North
America (Miocene - Pleistocene). Kendall/Hunt Publishing
Company, Dubuque, Iowa, vii 4- 373 pp.
Petuch, E.|. 2003. Cenozoic Seas. The view from eastern North
America. CRC Press, Boca Raton, Florida, [xii] 4- 308 pp.
(98 plates included in pagination) [Review: Allmon, 2005]
Petuch, E.J. 2007. The Geology of the Florida Keys and Ever-
glades. Thomson, Mason, Ohio, viii + 84 p
Petuch, E.|. and M. Drolshagen. 2009. Molluscan paleontology
of the Chesapeake Miocene. CRC Press, Boca Raton,
Florida, xviii + 160 p. (+ CD). [Review: Allmon, 2011]
Petuch. E.J. and D M. Sargent. 2012. Rare and unusual shells
of Southern Florida (Mainland, Florida Keys, Diy
Tortugas). Conch Republic Books, Mount Dora, Florida.
189 pp. (9 February)
Ward, L.W. 1992. Molluscan biostratigraphy of the Miocene -
Middle Atlantic Coastal Plain of North America. Virginia
Museum of Natural History, Memoir 2, 220 pp.
Ward, L.W. 2008. Synthesis of paleontological and stratigraphic
investigations at the Lee Creek Mine, Aurora, N.C. (1958-
2007). Virginia Museum of Natural History, Special Publi-
cation 14, pp. 325-432.
THE NAUTILUS 126(4): 148-149, 2012 Page 148
Research Note
New and confirmed fish hosts for the
threatened freshwater mussel Lampsilis
bracteata (Gould, 1855), the Texas Fatmueket
(Bivalvia: Unionidae)
The widespread decline of the freshwater mussel fauna
endemic to Texas indicates fundamental changes to the
ecology of the river systems inhabited by these species. In
order to appropriately manage these declining species, it
is imperative to understand their life history characteris-
tics. Lampsilis bracteata (Gould, 1855) has experienced a
precipitous decline in recent decades (Howells, 2010).
Historically, the species was dispersed throughout the
headwaters of the Edwards Plateau region and was found
in the Colorado and Guadalupe-San Antonio drainages
(Howells, 2010). However, as only several small popu-
lations are currently known to exist within these basins,
the species is listed as state threatened by the Texas Parks
and Wildlife Department (Texas Register 35, 2010) and as
a candidate species for federal protection by the U.S. Fish
and Wildlife Sendee (USFWS) (USFWS, 2011).
The purpose of this study was to confirm previously
identified host fishes (Howells, 1997) and identify addi-
tional hosts on which L bracteata glochidia may success-
fully transform. Although the reproductive biology of
tliis species has been studied to some degree in the past
(Howells, 1997), the quality of glochidia transformation on
previously identified host fishes and their suitability for
large-scale artificial propagation efforts remain unknown.
Two trials of gloehidial infestations were performed on
potential hosts, including Lepomis cyanellus (Rafinesque,
1819), Lepomis macrochirus (Rafinesque, 1819), Micropterus
salmoides (Laeepede, 1802), and Micropterus treculii
(Vaillant and Bocourt, 1883) (Table 1). Our methods fol-
lowed standard host-identification protocols (Jones et ah,
2004) and utilized a passive infestation technique. Each
trial was conducted independently using new batches of
fish and glochidia from different females. At the time of
the study, the only fish available were species already
housed at the USFWS San Marcos National Fish Hatchery
and Technology Center (Table 1 ).
Displaying gravid L. bracteata females were observed
in the San Saba River in Menard County, Texas, between
J uly and October 2011. Females were found by search-
ing bedrock crevices in the river bottom while diving
with the aid of a mask and snorkel. We checked
displaying females for gravidity' by carefully opening the
valves and inspecting for inflated marsupial gills.
Glochidia were collected from gravid mussels in July of
2011 by flushing well water through the marsupial gills
using a hypodermic needle. Potential host fishes were
placed in a plastic container holding 7.5 L of water.
Glochidia from two females also were placed in the con-
tainer. Airstones were used to agitate the water in the
container for 45 min. After infestation, fish were sepa-
rated by species and placed in individual aquaria. We
performed infestations on the same days that glochidia
were collected. Shell length, height, and hinge length
were measured for glochidia and newly transformed
juveniles (to the nearest 0.01 mm). Infested fish were
maintained in aquaria between 21 and 24 °C, which
were siphoned every 2 d for the first 14 d of the trial
and daily thereafter. We counted the total number of
juvenile mussels produced per fish (Table 1).
We were able to successfully transform a moderate to
high number of glochidia on all four of the tested fish
species. We confirmed two previously identified hosts
and identified two new hosts (Table 1). Transformation
time among the four fish species ranged from 17 to
26 d. Lepomis cyanellus produced the greatest number
of juveniles per fish (Table 1). Glochidia of L. bracteata
had a mean height of 0.21 mm, mean length of 0.18 mm,
and mean hinge length of 0.10 (SE < 0.01 and n = 10).
Glochidia grew only slightly during transformation on the
hosts. Newly transformed juveniles had a mean height of
0.25 mm, mean lengdi of 0.22 mm, and mean hinge
length of 0.10 mm (SE < 0.01 and n = 10).
The use of a mantle flap lure (Figure 1) and centrarchid
1 losts by Lampsilis bracteata parallels what researchers have
recorded for other Lampsilis spp. throughout die United
States (Zale and Neves, 1982, Haag et ah, 1999). The suc-
cessful transformation of L. bracteata on fish species that
are readily available to state and federal fish hatcheries is
Table 1. Results of fish host trials for Lampsilis bracteata conducted in 2011. (*) previously identified by Howells (1997)
M.S. Johnson et al., 2012,
Page 149
Figure 1. Mantle display of a gravid female Lampsilis
bracteata observed in the San Saba River, Menard Co., Texas
on July 22, 2011.
promising for future artificial propagation. Furthermore,
the successful propagation and advancement of juvenile
grow-out procedures for other Lampsilis spp. provides a
guide for the production of juveniles that may be used to
augment wiki populations or reintroduce the species into
areas where it has been extirpated. Although such a strategy
has already been developed, researchers within Texas still
need to invest time and resources into elucidating the details
of this species’ life history and ecological requirements.
ACKNOWLEDGMENTS
We thank E. Tsakiris, L. DiGiaeomo, A. Fabis, J. Fuller,
and S. McDonald for their assistance in the field, Julie
Groce and Neal Wilkins for supporting various aspects
of this study, the Institute of Renewable Natural
Resources, and the Texas Department of Transportation
(TxDOT) for funding this study.
and Villosa vibex (Bivalvia: Unionidae). American Midland
Naturalist 141:149-157.
Howells, R.G. 1997. New fish hosts for nine freshwater mussels
(Bivalvia: Unionidae) in Texas. Texas Journal of Science
49: 255-258.
Howells, R.G. 2010. Rare mussels: Summary of selected
biological and ecological data for Texas. Report submit-
ted to U.S. Fish & Wildlife. Report on file with Save Our
Springs Alliance.
Jones, J.W., R.J. Neves, S.A. Ahlstedt, and R.A. Mair. 2004. Life
history and propagation of the endangered dromedary
pearlymussel ( Dramas dramas ) (Bivalvia: Unionidae).
Journal of the North American Benthological Society
23: 515-525.
Texas Register 35. 2010. Threatened and endangered nongame
species. Chapter 65. Wildlife Subchapter G. 31 TAC
§65.175. Adopted rules. January 8, 2010: 249-251. Texas
Secretary of State.
U.S. Fish and Wildlife Service (USFWS). 2011. Endangered
and threatened wildlife and plants: 90-day finding on peti-
tions to list nine species of mussels from Texas as threat-
ened or endangered with critical habitat. Federal Register
74: 66260-66271.
Matthew S. Johnson1
Texas A&M Institute of Renewable Natural Resources,
College Station, TX 77843 USA
[email protected]
Patricia D. Caccavale
National Fish Hatchery and Technology Center,
U.S. Fish and Wildlife Service, San Marcos, TX 78666 USA
Charles R. Randklev
Texas A&M Institute of Renewable Natural Resources,
College Station, TX 77843 USA
James R. Gibson
National Fish Hatchery and Technology Center,
U.S. Fish and Wildlife Service, San Marcos, TX 78666 USA
LITERATURE CITED
Haag, W.R., M.L. Warren Jr., and M. Shillingsford 1999. Host 1 Current Address: Department of Fish and Wildlife Conserva-
fishes and host-attracting behavior of Lampsilis altilis tion, Virginia Tech, Blacksburg, VA 24060 USA
THE0NAUTILU S
Volume 126
2012
Aim, J.W.
Alewood, P.F.
Amano, K
Bail, P.
Bogan, A.E
Caccavale, P.D. ...
Chaytor, J.D
Costa, PM.S
Dutertre, S
Fraussen, K
Geiger, D.L
Gibson, J.R
Harasewych, M.G.
Hickman, C.S
Huber, M
Ituarte, C
Johnson, M.S
Kiel, S
Landman, N.H
Leal, J.H
Lee, H.G
Lewis, R.|
Lima, S.F.B
AUTHOR INDEX
47 Mapes, R.H
47 Martin, J.P
79 McLean, J.H
127 Mulvihill, R.S
105 Muttenthaler, M.
148 Ortigosa, D
86 Pastorino, G
25 Pearce, T.A
47 Petit, R.E
33 Porter, K.A
43 Puillandre, N
148 Randklev, C.R
119 Rex, M.A
57 Sellanes, |
15 Stahlschmidt, P. ..
136 Stuart, C.T.
148 Urteaga, D
79 Valdes, A
113 Walton, H
41 Watters, G.T.
15 Weaver, P.G
47 WlNGERD, J.S
41 Zelaya, D.G
113
136
89
38
47
98
. 25,68
38
41, 143
38
127
148
86
33
33
86
68
98
47
1
105
47
136
NEW TAXA PROPOSED IN VOLUME 126
GASTROPODA
Abbottella (Abbotella) urbana Watters, 2012, new species (Annulariidae) 1
Anomphalogaza Hickman, 2012, new genus (Gazidae) 62
Anomphalogaza moluccensis Hickman, 2012, new species (Gazidae) 62
Articulipoma rhodei Watters, 2012, new species (Annulariidae) 8
Calogaza colmani Hickman, 2012, new species (Gazidae) 58
Cerion petuchi Harasewych, 2012, new species (Cerionidae, fossil) 120
Chondropoma (Chondroporna) crystallinurn Watters, 2012, new species (Annulariidae) 3
Chondroporna (Chondropoma) duffijhooksonnn Watters, 2012, new species (Annulariidae) 4
Chondropoma (Chondropoma) vanattae polijchroma Watters, 2012, new subspecies (Annulariidae) 5
Chondropoma (Wetmorepoma) morsecodex Watters, 2012, new species (Annulariidae) 6
Chondropomium blaineorum Watters, 2012, new species (Annulariidae) 9
Chondropomium eusarcum saonense Watters, 2012, new subspecies (Annulariidae) 10
Chondropomium lynx Watters, 2012, new species (Annulariidae) 10
Eosipho zephyrus Fraussen, Sellanes, and Stahlschmidt, 2012, new species (Buccinidae) 34
Felimare sisalensis Ortigosa and Valdes, 2012, new species (Chromodorididae) 101
Fusivoluta profundorum Bail and Puillandre, 2012, new species (Volutidae) 130
Licina bartschi Watters, 2012, new species (Annulariidae) 13
Notocochlis laurae Costa and Pastorino, 2012, new species (Naticidae) 25
Natica juani Costa and Pastorino, 2012, new species (Naticidae) 25
Parachondria (Parachondria) gettlemani Watters, 2012, new species (Annulariidae) 6
Xeniostoma McLean, 2012, new genus (Calliostomatidae) 90
Xeniostoma inexpectans McLean, 2012, new species (Calliostomatidae) 91
BIVALVIA
Americardia lightboumi Lee and Huber, 2012, new species (Cardiidae) 17
Americardia columbella Lee and Huber, 2012, new species (Cardiidae) 19
Calyptogena veneriformis Amano and Kiel, 2012, new species (Vesicomyidae) 82
Mysella patagona Martin and Zelaya, 2012, new species (Galeommatoidea) 138
Triaslacus Bogan and Weaver, 2012, new genus (cf. Unionidae, fossil) 108
Triaslacus carolinensis Bogan and Weaver, 2012, new species (cf. Unionidae, fossil) 108
Biidiger Bieler
David Campbell
Stephanie Clark
Eugene V. Coan
Carlo M. Cunha
David Dockery
Tom Duda
Jeffrey T. Garner
Lindsey Groves
Carole S Hickman
Rebecca Johnson
Alan Kabat
REVIEWERS FOR VOLUME 126
Christian Klug
Alan Kohn
Elena Krylova
M .G. Harasewyeh
Joseph Hartman
Michael Hollman
Bruce A. Marshall
Gary McDonald
Paula M. Mikkelsen
Jeff Nekola
Sven Nielsen
Vinicius Padula
Gary Rosenberg
Aydin Orstan
John Slapcinsky
Richard Squires
John Stanisic
Paul Valentieh-Scott
Claude Vilvens
Peter Ward
G. Thomas Watters
Fred G. Thompson
Francisco Welter-Schultes
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Division of Cultural Affairs and the
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INSTRUCTIONS TO AUTHORS
THE NAUTILUS publishes articles on all aspects of the
biology, paleontology, and systematics of mollusks.
Manuscripts describing original, unpublished research
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THEft'NAUTILU S
Volume 127, Number 1
April 30, 2013
ISSN 0028-1344
CONTENTS
Research Note
Blake R. Hossack New distribution record for the rare limpet Acroloxus coloradensis
Robert L. Newell (Henderson, 1930) (Gastropoda: Aeroloxidae) from Montana 40
Notiee
42
THE NAUTILUS 127(1):1— 18. 2013
Page 1
Taxonomic review of Triphorinae (Gastropoda: Triphoridae)
from the Vitoria-Trindade Seamount Chain, southeastern Brazil
Mauricio R. Fernandes
Departamento de Invertebrados
Museu Nacional, Universidade Federal do Rio de Janeiro
Quinta da Boa Vista, Sao Cristovao
29040-040, Rio de Janeiro, BRAZIL
[email protected]
Jose H. Leal
The Bailey- Matt hews Shell Museum
3075 Sanibel-Captiva Road
Sanibel, FL 33957 USA
Alexandre D. Pimenta
Departamento de Invertebrados
Museu Nacional, Universidade Federal do Rio de Janeiro
Quinta da Boa Vista, Sao Cristovao
29040-040, Rio de Janeiro, BRAZIL
ABSTRACT
In the present work we identified the shells of Triphorinae col-
lected at the seamounts of the Vitoria-Trindade Chain off south-
eastern Brazil by the expeditions MD55 and REVIZEE-Central,
the main sources of mollusks Irom the study site in the last
decades. Of the 13 species found in this study, Cosmotriphora
melanura and Nototriphora decorata were previously reported
from the Vitoria-Trindade Chain. Cosmotriphora amoldoi,
Iniforis cannelae , Iniforis pseudothomae, Latitriphora alhida ,
Coriophora novem, Triphora elhjae and Triphora elvirae are
reported for the first time from Brazil. The known ranges in
Brazil of Monophorus olivaceus and Triphora atlantica are
extended to the Vitoria-Trindade Chain. Isotriphora tigrina
new species, diagnosed by its teleoconch with two main spiral
cords and three brown and smooth basal cords, is currently
restricted to the Vitoria-Trindade Chain and Bahia state.
Isotriphora onca new species, diagnosed by its teleoconch with
two main spiral cords and creamy to light brown color, is cur-
rently restricted to the Vitoria-Trindade Chain. The number of
species of Recent Triphoridae in the Vitoria-Trindade Chain
increased from six to 15, and in Brazil from 17 to 26.
Additional Keywords: Western Atlantic, MD55, REVIZEE,
Mollusca, Triphoroidea, RV Marion-Dufresne
INTRODUCTION
Triphoridae is a large group of marine microgastropods
that feed on sponges, mainly distributed in tropical and
temperate seas (Marshall, 1983). Triphorids are usually
recognized by their left-coiled shell, although species of
Metaxiinae are dextral. Bouchet and Rocroi (2005) con-
sidered the existence of three subfamilies: Triphorinae,
Metaxiinae, and Iniforinae, although the validity of die
latter is contested by Marshall (1983). We follow Niitzel
(1998), recognizing only Triphorinae and Metaxiinae.
The maximum diversity of Triphoridae is found in the
tropical Indo-Pacific (Rolan and Fernandez-Garces, 2008).
In the western Atlantic, there are 93 recent species
(Simone, 2006; Rolan and Fernandez-Garces, 2008, 2009;
Faber, 2010; Fernandes and Pimenta, 2011). In Brazil,
17 species are reported (Absalao, 1989; Simone, 2006;
Rolan and Fernandez-Garces, 2008; Lee, 2009; Rios, 2009;
Fernandes and Pimenta, 2011), although some of them
need confirmation.
The Vitoria-Trindade Seamount Chain (Vitoria-Trindade
Ridge; Columbia Fracture Zone) comprises a series of
seamounts and oceanic islands between 20°-21° S and
28°50' - 38°30' W, off Espfrito Santo state, southeastern
Brazil. The sequence of seamounts initiates near the
base of the continental slope (Champlain and Vitoria
seamounts), running eastward to Trindade Island and
Martin Vaz Archipelago (Figure 1), about 1167 km from
the continent (Leal, 1991; O'Hara et ah, 2010). Only
Trindade Island and Martin Vaz Archipelago rise above
sea level, while the seamount summits rise to between 50
and 150 m below the surface. The Vitoria-Trindade Chain
is under the influence of the warm, saline, southward-
flowing Brazil Current (O'Hara et ah, 2010).
Two scientific expeditions provided the majority of
mollusks collected at the Vitoria-Trindade Chain in the
last decades: the Freneh-Brazilian Expedition MD55,
conducted in 1987, and REVIZEE-Central (“Programa
de Avaliayao do Potencial Sustentavel de Recursos Vivos
da Zona Economica Exclusiva”, SCORE Central; “Pro-
gram of Evaluation of the Sustainable Potential of Living
Resources in the Exclusive Economic Zone”, SCORE
Central), conducted between 1996 and 2002.
The prosobranchs collected at the Vitoria-Trindade
Chain by the Expedition MD55 were studied by Leal
(1991) as the basis for a biogeographic analysis of the
oceanic islands of Brazil. Leal (1991) recognized 10 taxa
Page 2
THE NAUTILUS, Vol. 127, No. 1
Figure 1. Stations in which shells of Triphoridae were collected at the Vitoria-Trindade Chain by Expedition MD55 (DC) and
REVIZE E-Central (C1-C5).
for Triphoridae, of which seven were identified only at
generic level, although that author admitted that “some
of the study species do not fit these classifications at the
generic level”. Absalao et al. (2006) listed six species of
triphorids from the Vitoria-Trindade Chain, collected
by REVIZEE. Fernandes and Pimenta (2011) reported
Metaxia excelsa Faber and Moolenbeek, 1991 and
Metaxia prompta Rolan and Fernandez-Garces, 2008
at the study site.
Within these works, the species reported at the Vitoria-
Trindade Chain are Metaxia excelsa , Metaxia prompta,
Cosmotriphora melanura ( C . B. Adams, 1850), Nototriphora
decorata (C. B. Adams, 1850), Iniforis turristhomae
(Holten, 1S02) and Marshallora nigrocincta (C. B. Adams,
1839) in addition to seven morphotypes that remained
identified only at generic level.
As the Metaxiinae were already revised for Brazil
(Fernandes and Pimenta, 2011), the present work is
focused on the taxonomy of Triphorinae from the Vitoria-
Trindade Chain based on the material collected by the
expeditions MD55 and REVIZEE-Central.
MATERIALS AND METHODS
In May/] une 1987, the Research Vessel Marion-Dufresne
made a series of dredgings and trawlings in southern
Bahia state (Abrolhos bank) and in the Southeast region
of Brazil, during the Expedition MD55 (Tavares, 1999).
There were 66 benthic collecting stations, in depths
between 15 and 5100 m, including the top of the sea-
mounts that form the Vitoria-Trindade Chain. The sta-
tions in which shells of Triphoridae were collected at the
Vitoria-Trindade Chain by the Expedition MD55 are
listed in Table 1 .
The program REVIZEE was divided in four regions
(one for each SCORE), representing the different ocean-
ographic characteristics of the Brazilian coast (Lavrado,
2006). The SCORE Central comprised the area between
Salvador (Bahia state) to Cabo de Sao Tome (Rio de
Janeiro state), including the Vitoria-Trindade Chain.
REVIZEE-Central consisted of six campaigns (Cl to C6),
of which the first one (Cl) was conducted by the Ocean-
ographic Ship Antares and the remainder (C2 to C6)
by the supply boat N/RB Astro Garoupa. There were
227 collecting stations, in depths between 12 to 2076 m.
The stations in which shells of Triphoridae were collected
at the Vitoria-Trindade Chain by REVIZEE-Central are
listed in Table 2.
All material available consisted of empty shells. The
taxonomic identifications were based on eonehological
comparisons under a stereo microscope. Each species was
also observed in scanning electron microscope (SEM)
images. The whorl-counting procedure follows Leal
(1991). The embryonic shell is here considered the
beginning of the protoconch, typically sculptured on
Triphorinae with spherical granules, cruciform granules
or reticulated pattern . The remainder whorls of the
protoconch constitute the larval shell, usually sculptured
on Triphorinae with spiral and axial cords.
In order to obtain complete sinonimic lists, some cita-
tions were included even without direct or indirect
examination of the respectively material. In these cases,
the expression “not illustrated” follows the citation.
In the section of “material examined”, the number
inside brackets indicates the quantity of shells in each lot.
Abbreviations used: (ABC Islands) Aruba, Bonaire and
Curasao; (AMNH) American Museum of Natural History,
New York, USA; (ANSP) Academy of Natural Sciences,
Philadelphia, USA; (CEB) E. Rolan, Vigo, Spain; (CFG)
B. Fernandez-Garces, Cienfuegos, Cuba; (FLMNII)
Florida Museum of Natural History, Gainesville, USA;
(IBUFBJ) Instituto de Biologia, Universidade Federal
do Bio de Janeiro, Rio de Janeiro, Brazil; (IES) Instituto
de Ecologia y Sistematica, Havana, Cuba; (MCZ) Museum
of Comparative Zoology, Cambridge, USA; (MHNS) Museo
de Historia Natural, Santiago de Compostela, Spain;
(MNCN) Museo Nacional de Ciencias Naturales, Madrid,
Spain; (MNHN) Museum national d’Histoire naturelle,
Paris, France; (MNBJ) Museu Nacional, Universidade
Federal do Rio de Janeiro, Rio de Janeiro, Brazil; (MORG)
Museu Oceanografico do Rio Grande, Rio Grande, Brazil;
( MZSP) Museu de Zoologia da Universidade de Sao Paulo,
Sao Paulo, Brazil; (NHM) Natural History Museum,
Fondon, England; (ZMA) Zoologisch Museum Amsterdam,
Amsterdam, The Netherlands.
RESULTS
Family Triphoridae Gray, 1847
Subfamily Triphorinae Gray, 1 847
Diagnosis: Sinistral shells, with a tubular or subtubular
anterior canal, the posterior canal forming a notch, a hole
or a tube (based on Wilson, 1993).
Genus Cosmotriphora Olsson and Harbison, 1953
Type Species: Cerithium melanurci C. B. Adams, 1850,
original designation; Recent, Atlantic Ocean.
Diagnosis: Embryonic shell with granules on abapieal
region, larval shell with axial riblets crossed by one spiral
cord at beginning, two cords latter; teleoconch with a
later development of median spiral cord; radular formula
9- 1-1-1 -9, with lour cusps on central and lateral teeth.
Page 4
THE NAUTILUS, Vol. 127, No. 1
three cusps on internal marginal teeth, two cusps on
external marginal teeth (based on the description of the
type species of the genus in Bouchet, 1985).
Cosmotriphora amoldoi Faber and M<x>Ienbeek, 1991
(Figures 2, 18, 29)
Cosmotriphora amoldoi Faber and Moolenbeek (1991:
81, figs. 1-2); Rolan and Fernandez-Garces (1994:
20, figs. 12-15; 2007: 20, pi. I, figs. 19-21); Lee
(2009: 88, text-fig.); Garcia and Lee (201 1 ).
Triphora melanura : Absalao et al. (2006: 238, in part)
non C. B. Adams, 1850.
Triphora sp.: Absalao et al. (2006: 238, in part).
Type Material: Holotype: ZMA 391001 . One paratvpe
ZMA 391002.
Type Locality: Playa Lechi, Bonaire.
Material Examined: IBUFRJ 13568, REVIZEE C5-
24 R [1]; IBUFRJ 19447, REVIZEE C1-C62 [2]; MORG
51900, REVIZEE C I -C62 [3]; MNHN, MD5535-DC59 [1],
Remarks: In addition to C. arnoldoi , the other species
of Cosmotriphora in the West Atlantic is C. melanura
(C. B. Adams), which has an earlier development of the
median spiral cord (Figure 19) and a white teleoeonch
(Figure 3), instead of the reddish-brown blotches in the
teleoeonch of C. amoldoi (Figure 2). This pattern of col-
oration is similar to Nototriphora decorata (C. B. Adams,
1850), a very common species in the West Atlantic.
Cosmotriphora amoldoi has a more curvilinear profile;
its median spiral cord of the teleoeonch has the same size
as the adapieal and abapical spiral cords on the eighth/
ninth whorl (although appearing weakly on the fifth
whorl) (Figure 18), while in N. decorata it reaches the
same size on the fifth/sixth whorl (Figure 25); its axial
blotches are more or less parallel, not irregular as in
N. decorata-, the three spiral cords are covered by
blotches, except in the body whorl, where the adapieal
spiral cord is usually totally white (Figure 2).
The generic placement of this species by Faber and
Moolenbeek (1991) was made without knowledge of
radula and operculum, possibly only using protoconch
morphology (Rolan and Fernandez-Garces, 1994). The
presence of granules on the entire embryonic whorl of
Figures 2-17. Triphoridae species. 2. Cosmotriphora amoldoi , MNHN. 3. Cosmotriphora melanura , MNRJ 25019. 4. Iniforis
cannelae , MNRJ 25014. 5 Iniforis pseudothomae , MNRJ 25033. 6. Iniforis sp., IBUFRJ 10563. 7. Isotriphora tigrina new species,
MNRJ 25992, holotype. 8. Isotriphora tigrina new species, MNRJ 16227, paratype. 9. Isotriphora onca new species, MNRJ 16236,
holotype. 10. Isotriphora onca new species, MNHN IM-2012-21 11, paratype. 1 1. Latitriphora alhida, MNRJ 25016. 12. Coriophora
novem , MNRJ 25008. 13. Monophorus olivaceus, MNRJ 25027. 14. Nototriphora decorata, MNRJ 25002. 15. Triphora atlantica ,
IBUFRJ 13142. 16. Triphora elh/ae, MNRJ 18955. 17. Triphora elvirae, MNHN. Scale bar = 1 mm.
M.R. Fernandes, A.D. Pimenta and J.H. Leal, 2013
Page 5
Figures 18-28. Triphoridae species. 18. Cosmotriphora amoldoi, IBUFRJ 13568. 19. Cosmotriphora melanura , MNRJ 25024.
20. Iniforis carmelae, MNRJ 25541. 21. Iniforis pseudothomae , MNRJ 25033. 22. Latitriphora alb id a , MNRJ 25016. 23. Coriophora
novem , MNRJ 25011. 24. Monophorus olivaceus, MNHN. 25. Nototriphora decorata, MNRJ 25033. 26. Triphora atlantica , IBUFRJ
13142. 27. Triphora elhjae , MNRJ 18955. 28. Triphora elvirae , MNRJ 25032. Scale bar = 1 mm.
C. amoldoi (Figure 29), not only on its abapical region as
in C. melanura (Figure 30), suggests an affinity witli the
genus Nototriphora Marshall, 1983, pending confirma-
tion about the importance of this difference between
Cosmotriphora and Nototriphora. Marshall (1983) men-
tioned only differences in operculum and radula about
the two genera.
Distribution: USA: Florida (Lee, 2009), Louisiana
(Garcia and Lee, 2002); Gulf of Mexico (Rosenberg
et ah, 2009); Bahamas (Faber and Moolenbeek, 1991);
Cuba (Rolan and Fernandez-Garces, 1994); Puerto Rico
(Faber and Moolenbeek, 1991); ABC Islands (type
locality); Brazil: Vitoria-Trindade Chain (this study).
Cosmotriphora melanura (C. B. Adams, 1850)
(Figures 3, 19, 30)
Cerithium melanura C. B. Adams (1850: 117); Clench
and Turner (1950: 307, pi. 38, fig. 10).
Triforis melanura : Dali and Simpson (1901: 423, pi. 58,
fig. 7).
Triforis grimaldii Dautzenberg and Fischer (1906: 41,
pi. Ill, figs. 9-10).
Triphora melanura : Rios (1970: 45, not illustrated; 1975:
50, pi. 13, fig. 187; 1985: 161, pi. 53, fig. 761; 1994: 94,
pi. 31, fig. 374; 2009: 172, text-fig.); Abbott (1974: 111,
fig. 1 134); Vokes and Vokes (1983: 18, pi. 27, fig. 14);
Jong and Coomans (1988: 49, not illustrated); Absalao
Page 6
THE NAUTILUS, Vol. 127, No. 1
Figures 29-38. Protoconehs. 29. Cosmotriphora amoldoi, IBUFRJ 13568. 30. Cosmotriphora melanura, MNRJ 25024. 31. Iniforis
cannelae, MNRJ 25541. 32. Iniforis pseudothomae , MNRJ 25033. 33. Coriophora novem, MNIIN. 34. Monophorus olivaceus,
MNIIN. 35. Nototriphora decorata, MNRJ 25033. 36. Triphora atlantica, IBUFRJ 13142. 37. Triphora elli/ae, MNRJ 18955.
38. Triphora elvirae , MNIIN. Seale bar = 100 pm.
(1989: 3, not illustrated); Merlano and Hegedus
(1994: 148, pi. XLVI, fig. 524); Absalao et al. (2006:
238, fig. 9, in part); Gomes et al. (2006: 188, not
illustrated); Santos et al. (2007: 226, not illustrated).
Cosmotriphora melanura: Marshall (1983: 110, fig. 27 D-F);
Bouchet (1985: 35-37, fig. 27); Fernandes and Rolan
(1988: 22, pi. 1, fig. 1, pi. 2, fig. 2); Leal (1991: 120, pi. 16,
figs. F-G); Rolan and Femandez-Garces (1994:
19, figs. 11, 25, 26, 30 CM); Redfem (2001: 65, pi. 32,
fig. 274); Espinosa and Ortea (2001: 20, not illustrated);
Ardovini and Cossignani (2004: 134, text-fig.); Rolan
(2005: 106, pi. 30, fig. 438); Rolan and Femandez-Garces
(2007: 20, pi. 1, figs. 14-16); Jensen and Pearce (2009:
128, not illustrated); Lee (2009: 88, text-fig.); Tunnell
et al. (2010: 204, text-fig.); Garcia and Lee (201 1).
Triphora (Cosmotriphora) melanura: Ode (1989: 109, fig. 3).
Triphora sp.: Absalao et al. (2006: 238, in part).
Type Material: Lectotype: MCZ 186159.
Type Locality: Jamaica.
Material Examined: IBUFRJ 9309, REVIZEE Cl-
C61 [52]; IBUFRJ 10222, REVIZEE C1-C62 [11];
IBUFRJ 11952, REVIZEE C5-23R [1]; IBUFRJ 11993,
REVIZEE C5-42R [1]; IBUFRJ 12010, REVIZEE C5-
45 R [1]; IBUFRJ 12018, REVIZEE C5-48R [1];
IBUFRJ 13567, REVIZEE C5-24R [5]; IBUFRJ 13581,
REVIZEE C5-23R [6]; IBUFRJ 13594, REVIZEE C5-
45 R [3]; IBUFRJ 13749, REVIZEE C5-24R [8];
IBUFRJ 14372, REVIZEE C5-45R [1]; IBUFRJ 14577,
REVIZEE C5-30R [6]; IBUFRJ 14649, REVIZEE
C5-49R [2]; IBUFRJ 14674, REVIZEE C5-44R [14];
IBUFRJ 14694, REVIZEE C5-48R [3]; IBUFRJ 14713,
REVIZEE C5-45R [2]; IBUFRJ 14717, REVIZEE
C5-42R [3]; IBUFRJ 19457, REVIZEE C1-C61 [14];
IBUFRJ 19458, REVIZEE C1-C62 [8]; MNRJ 12740,
REVIZEE C5-42R [5]; MNRJ 12768, REVIZEE C5-
21R [1]; MNRJ 25017, MD55 24-DC42 [3]; MNRJ
25018, MD55 23-DC4I [5]; MNRJ 25019, MD55 25-
DC43 [5]; MNRJ 25020, MD55 10-DC24 [3]; MNRJ
25021, MD55 14-DC27 [1]; MNRJ 25022, MD55 21-
DC35 [28]; MNRJ 25023, MD55 23-DC40 [1]; MNRJ
25024, MD55 9-DC22 [6]; MORG 51899, REVIZEE
C1-C62 [3]; MNHN, MD55 20-DC34 [23]; MNHN,
MD55 25-DC43 [3]; MNHN, Enseada dos Portugueses,
Trindade Is., v/1987 [2],
Remarks: This species is clearly distinguished by its
white teleoconch and brown protoconch (Figure 3), with
a mammilliform shape (Figure 30). This was by far the
most common species on the material studied, and it has
an amphi-Atlantic distribution (Rolan and Fernandez-
Garces, 1994). However, the shells of C. melanura of
the East Atlantic seem to have some differences with
the Caribbean shells, especially the later development
of the median spiral cord in the teleoconch (Bouchet,
1985). In the material examined by Bouchet (1985) and
M.R. Fernandes, A.D. Pimenta and J.H. Leal, 2013
Page?
Rolan and Fernandez-Garces (1994) of the west coast ol
Africa, the median spiral cord appears between the sixth
and eighth whorl, while in the shells herein studied and
the Caribbean ones (e.g., Rolan and Fernandez-Garces,
1994) this cord usually begins in the third whorl
(Figure 19). Following Rolan and Fernandez-Garces
(1994), the protoconchs and radulae of the two populations
seem to be similar, and we accept the small divergence in
the development of the median spiral cord as an intra-
specific variation. This morph is the basis for the well-
established synonym Triforis grimaldii Dautzenberg and
Fischer, 1906, from the west coast of Africa.
Triphora dealbata (C. R. Adams, 1850), from Jamaica,
is considered by some authors (e.g. Rolan and Fernandez-
Garces, 2008) as a possible synonym for C. melanura.
In addition to a wide geographic distribution,
Cosmotriphora melanura is also considered by Dali (1892)
to be present in tire Miocene of Florida.
Distribution: Europe (Bouchet, 1985); West coast of
Africa (Rolan, 2005); Bermuda (Jensen and Pearce,
2009); USA: North Carolina (Abbott, 1974), Florida
(Lee, 2009), Louisiana (Garcia and Lee, 2002), Texas
(Tunnell et al., 2010); Mexico (Vokes and Vokes, 1983);
Bahamas (Redfern, 2001); Cuba (Rolan and Fernandez-
Garces, 1994); Belize (Miloslavieh et ah, 2010); Jamaica
(type locality); Puerto Rico (Dali and Simpson, 1901);
Virgin Islands; Costa Rica (Espinosa and Ortea, 2001);
Colombia (Merlano and Hegedus, 1994); ABC Islands
(Jong and Coomans, 1988); Brazil: Amapa to Bahia
(Rios, 1985), Sao Pedro-Sao Paulo Is. (Rios, 2009),
Fernando de Noronha and Vitoria-Trindade Chain (Leal,
1991; Gomes et ah, 2006), Espfrito Santo (Absalao, 1989;
Absalao et ah, 2006), Rio de Janeiro (Absalao, 1989;
Absalao et ah, 2006; Santos et ah, 2007).
Genus Iniforis Jousseaume, 1884
Type Species: Iniforis malvaceus Jousseaume, 1884,
original designation; Recent, New Caledonia.
Diagnosis: Paueispiral or multispiral protoconch;
teleoconch whorls with two rows of large nodules and
granulose interspaces; suture indistinguishable; aperture
round; posterior and anterior canals short and tubular
(based on Wilson, 1993).
Iniforis carmelae Rolan and Fernandez-Garces, 1993
(Figures 4, 20, 31)
Triphora sp. 3: Leal (1991: 123, pi. 17, figs. A-B).
Iniforis carmelae Rolan and Fernandez-Garces (1993:
102, figs. 12-15, 28-30; 2007: 21, pi. II, figs. 18-22).
Type Material: Holotype: MNCN 15.05/6822. Paratypes:
One in ZMA 3.93.006, AMNH 226457, IES, NHM
1992134, MNHN; five in the CFG; 15 in the CER.
Type Locality: Cienfuegos, in the South of Cuba.
Material Examined: MNRJ 25013, MD55 35-DC59
[2]; MNRJ 25014, MD55 21-DC35 [1]; MNRJ 25541
MD55 9-DC22 [1],
Remarks: The material here studied agrees with the
original description in having the white and relatively
broad protoconch with approximately 2.75 whorls
(Figure 31) and the later development of a very small
median spiral cord on the teleoconch (ninth/tenth whorl,
very close to the adapical spiral cord; Figure 20) in addi-
tion to the discontinuous brown coloration on the
abapical spiral cord of the teleoconch (Figure 4). How-
ever, the shells from Vitoria-Trindade Chain display
some slight differences from the original description: a
more tuberculated subperipheral cord (Figure 20, if
compared to the figure 28 in Rolan and Fernandez-
Garces, 1993), the continuous brown coloration of the
abapical spiral cord of tire teleoconch on its initial whorls
(Figure 4), and the spiral cord on the second whorl of
the protoconch is positioned in its middle (Figure 31)
although the original description shows this spiral cord
situated barely under the middle of the whorl (figs. 29-30
in Rolan and Fernandez-Garces, 1993).
Iniforis carmelae can be best differentiated from
Iniforis immaculata Rolan and Fernandez-Garces, 1993,
a species from Cuba, by the shell color, translucent white
in the later. Iniforis gudeliae Rolan and Fernandez-
Garces, 2009 and Iniforis pelorcei Rolan and Fernandez-
Garces, 2009, two species from the Caribbean, have
the second whorl of protoconch with two spiral cords
(I. carmelae has only one) in addition to a more contin-
uous brown coloration on the abapical spiral cord of
the teleoconch.
Distribution: Gulf of Mexico (Rosenberg et al., 2009);
Cuba (type locality); Brazil: Vitoria-Trindade Chain
(this study).
Iniforis pseudothomae Rohm and Fernandez-
Garces, 1993
(Figures 5, 21, 32)
Triphora sp. 2: Leal (1991: 123, pi. 16, figs. l^M).
Iniforis pseudothomae Rolan and Fernandez-Garces
(1993: 100, figs. 5-8, 22-23; 2007: 21, pi. II,
figs. 9-12).
Triphora decorata : Absalao et al. (2006: 238, in part) non
C. B. Adams, 1850.
Triphora melanura-. Absalao et al. (2006: 238, in part)
non C. B. Adams, 1850.
Triphora sp.: Absalao et al. (2006: 238, in part).
Triphora turristhomae : Rios (2009: 173, text-fig.) non
Holten, 1802.
Type Material: Holotype: MNCN 15.05/6820. Paratypes:
One in AMNH 226459, MNHN, NHM 1992133, ZMA
3.93.005; five in the IES; ten in the CER and CFG.
Type Locality: Cienfuegos, in the South of Cuba.
Page 8
THE NAUTILUS, Vol. 127, No. 1
Material Examined: IBUFRJ 19443, REVIZEE
C5-30R [1]; IBUFRJ 19455, REVIZEE C.1-C61 [1];
MNRJ 25033, MD55 9-DC22 [8]; MNRJ 25034, MD55
25-DC43 [2]; MNHN, MD55 20-DC34 [4],
Remarks: As suspected by Rolan and Fernandez-
Garces (1993), Triphora sp. 2 in Leal (1991) represents
Iniforis pseudothomae . This species has a teleoconch
very similar to I. carmelae, with discontinuous brown
color on the abapical spiral cord (Figure 5); shells of
I pseudothomae never develop a median spiral cord as
1 carmelae. Also, the brown and elongated protoconch
of 7 pseudothomae has around 5-5.5 whorls and two
spiral cords on the larval shell, except on the last whorl
(Figure 32), instead of the ~2.75 white whorls of
7. carmelae , with one spiral cord on the larval shell
(Figure 31). The brown and mutispiral protoconch of
I pseudothomae also differentiates it from 7. g udeliae
and 7. pelorcei.
Tl le illustration provided by Rios (2009) as
Triphora turristhomae (Holten, 1802) actually depicts
7 pseudothomae. Possibly die records of Iniforis
turristhomae (Holten, 1802) from Brazil are erroneous
(Leal, 1991), and they can be actually referred to
7 pseudothomae , but this hypothesis demands further
investigation. Iniforis turristhomae has a continuous
brown band on the abapical spiral cord of the teleoconch
and two spiral cords on the protoconch in contrast to
7 pseudothomae , which has a discontinuous brown band
on the abapical spiral cord of the teleoconch (Figure 5)
and only one spiral cord on the last whorl of the
protoconch (Figure 32).
The single shell of Iniforis turristhomae (Holten, 1802)
from the Vitoria-Trindade Chain, reported by Absalao
et al. (2006) (IBUFRJ 10563, station REVIZEE C2-
22F), is very eroded, making a correct identification
difficult. Actually, this shell has a continuous brown
band on the abapical spiral cord (Figure 6), ruling out
7. pseudothomae. The presence of brown color between
the two spiral cords and the development of a very small
median cord on the body whorl adjacent to the adapical
spiral cord are not consistent with 7. turristhomae. We
prefer to maintain this shell as Iniforis sp., invalidating
the previous record of 7. turristhomae for the Vitoria-
Trindade Chain.
Distribution: Gulf of Mexico (Rosenberg et al., 2009);
Cuba (type locality); ABC Islands (Miloslavich et al., 2010);
Brazil: Vitoria-Trindade Chain (diis study).
Genus Isotriphora Cotton and Godfrey, 1931
Type Species: Triforis tasmanica Tenison- Woods, 1875,
original designation; Recent, southern Australia.
Diagnosis: Paucispiral protoconch, blunt-tipped, with
two nodulose spiral cords; teleoconch whorls heavily
nodulose; anterior canal short to moderately long, usually
subtubular (based on Wilson, 1993).
Isotriphora tigrina new species
(Figures 7-8, 39-44)
Triphora sp. 4: Leal (1991: 124, pi. 17, figs. C-D).
Triphora sp.: Absalao et al. (2006: 238, in part).
Type Material: Holotype: MNR( 25992. Paratypes:
REVIZEE C5-49R: MNRJ 16227 [2], IBUFRJ 17051
[2]; type locality: MZSP 105154 [3], MNRJ 25993 [7],
MNHN IM-2012-2110 [8],
Type Locality: Trindade Island, Vitoria-Trindade
Chain, Brazil (20°30' S, 29°19' W, 52-60 m). Station 35-
DC59 of the Expedition MD55.
Other Material Examined: IBUFRJ 14667, REVIZEE
C5-49R [3]; IBUFRJ 14718, REVIZEE C5-48R [3];
MNRJ 25029, MD55 21-DC35 [2]; MNRJ 25030,
MD55 35-DC59 [1]; MNHN, Enseada dos Portugueses,
Trindade Is., v/1987 [1]; MZSP 100957, Cairu, Morro
de Sao Paulo, Bahia, Petronio Coelho-Filho coll.,
2011 [1].
Diagnosis: Shell with a truncated apex; teleoconch
with two main spiral cords, the abapical one with brown
inter-nodular spaces during most of tire shell; subperipheral
cord and two basal cords brown and smooth.
Description: Shell elongated, conical, reaching 5.2 mm
in length, 1 .4 mm in width, wnth up to 13 whorls.
Paucispiral protoconch with truncated apex and without
clear differentiation from teleoconch. After the very
narrow nucleus, two white spiral cords (one adapical and
one small median) and a brown abapical spiral cord
emerge; after approximately one whorl the small median
spiral cord disappears, almost simultaneously with a
change in color of abapical spiral cord (brown to white)
and adapical spiral cord (white to brown) on next whorl;
adapical spiral cord tinted with brown for ~ two whorls,
where it gradually fades, assuming color of shell
background, which varies from creamy to light brown;
simultaneously, abapical spiral cord becomes brown in
inter-nodular spaces, until shell ends. Nodules of abapical
spiral cord more spaced than adapical one; at the 11th
whorl, a very small and wavy median spiral cord appears
very close to adapical one; suture barely distinct, with a
small cord more visible on last whorls; subperipheral cord
and two basal cords, all smooth and brown; aperture
ovoid, with a small posterior canal; outer lip well projected
to front of shell; long and nearly closed siphonal canal,
curving downward/backward. Animal unknown.
Etymology: This species is named for its brown
subperipheral and basal cords, resembling the stripes of
a tiger.
Remarks: A similar species to 7. tigrina in the west-
ern Atlantic is Isotriphora guanahacabibes Rolan and
Fernandez-Garces, 2008, a species from Caribbean.
However 7. tigrina has a unique color pattern of brown
bands, in the adapical spiral cord of earlier whorls and
M.R. Fernandes, A.D. Pimenta and J.H. Leal, 2013
Page 9
mainly in the inter-nodular spaces of the abapical spiral
cord of the remaining whorls (instead of the whitish shells
of I. guanahacabibes) (Figures 7-8); two distinct basal
cords (while I guanahacabibes has only one) (Figure 40),
and a later development ol the median spiral cord (in
the 11th whorl of the shell in I tigrina , ninth in
I. guanahacabibes , as figured in Rolan and Fernandez-
Garces, 2008). Isotriphora peetersae (Moolenbeek and
Faber, 1989), another species from Caribbean, has a
totally different color than I. tigrina , with about five initial
whorls of die shell being white and the remainder dark
brown, in addition to a median spiral cord appearing on
the fourth/fifth whorl of the teleoconch. Isotriphora
taenialba Rolan and Espinosa, 1994, a species from Cuba,
has the adapical spiral cord of the teleoconch and suture
dark brown, and the median spiral cord becomes nodulous
much earlier on the fourth/fifth whorl of die teleoconch.
Isotriphora tigrina shares with Iniforis carmelae and
Iniforis pseudothomae the discontinuous brown color
on the abapical spiral cord of the teleoconch, but in
I. tigrina this color is mainly restricted to the inter-nodular
spaces. The protoconch typical of the genus Isotriphora
(widi nodular spiral cords) (Figure 44), the presence of a
smooth subperipheral cord (Figure 40) and a small poste-
rior canal above the aperture (not deflected to the other
side of the shell, as in Iniforis ) are other features that
distinguish /. tigrina from these two species. Also, a small
and non-tuberculated median spiral cord appears on
die teleoconch of I. tigrina (Figures 39-40), while
/ pseudothomae never develops it.
Figures 39-44. Isotriphora tigrina new species. 39, 43. Entire shell. 40, 42. Last whorl anti base. 41. Adapical view of
protoconch. 44. Protoconch and beginning of teleoconch. 39-40, 42-44. MNRJ 25993, paratype. 41. IBUFRJ 14718. Seale bar =
1 mm to figures 39-40, 42-43, 100 pm to figures 41, 44.
Page 10
THE NAUTILUS, Vol. 127, No. 1
Distribution: Brazil: Bahia state and Vitoria-Trindade
Chain (this study).
Isotriphora onca new species
(Figures 9-10, 45-50)
Triphora sp.: Absalao et al. (2006: 238, in part).
Type Material: Holotype: MNBJ 16236. Paratypes:
MD55 27-DC47: MNHN IM -2012-21 11 [1]; REVIZEE C5-
42R: MZSP 1 12068 [2], IBUFRJ 14367 [7], MNRJ 16230 [8],
Type Locality: Trindade Island, Vitoria-Trindade
Chain, Brazil (20°30'S, 29°16'W, 360 m). Station C5-41F
of REVIZEE-Central.
Other Material Examined: IBUFRJ 12123, REVIZEE
C5-42R [1]; IBUFRJ 14411, REVIZEE C5-42R [22];
MNRJ 16232, REVIZEE C5-42R [4]; MNRJ 17918,
REVIZEE C5-42R [3],
Diagnosis: Shell with a truncated apex; color creamy
to light brown, except the whitish apex; teleoconch with
two main spiral cords.
Description: Shell elongated, conical, reaching 5.6 mm
in length, 1.4 mm in width, with up to 14 whorls.
Paucispiral protoconch with truncated apex and without
clear differentiation from teleoconch. After the very nar-
row nucleus two spiral cords emerge; after approxi-
mately one whorl the adapical spiral cord disappears,
and a small cord starts to develop above that, becoming
a new adapical spiral cord. Creamy to light brown back-
ground color, nodules slightly lighter, inter-nodular
spaces slightly darker than background color, especially
on abapical spiral cord; two-three initial whorls whitish,
with a faded brown abapical spiral cord. Nodules ot the
abapical spiral cord may be slightly more spaced than
adapical one; few shells have a very small median spiral
cord after the 13th whorl, very close to adapical spiral
cord; suture barely distinct, with a small cord more visi-
ble on last whorls; subperipheral cord and two basal cords,
all smooth and of same color as background; adapical
basal cord, closer to subperipheral cord, smaller and
absent in a few shells; aperture ovoid, with small posterior
canal; outer lip well projected to front of shell; long and
nearly closed siphonal canal, curving downward/backward.
Animal unknown.
Etymology: The specific name alludes to the jaguar
( Panthera onca ), due to the similarity with the previously
described species, Isotriphora tigrina.
Remarks: This species resembles I. tigrina , including
the general shape of the shell (although I onca lias a
more slender profile), the truncated apex with similar
development of the spiral cords, the suture barely dis-
tinct and the presence of two main spiral cords. How-
ever, I. onca has a different color pattern (Figure 9-10),
being almost entirely creamy to light brown, with a whit-
ish apex, and the subperipheral-basal cords are not
tinted with brown. Isotriphora onca has a smaller (some-
times absent) adapical basal cord (Figure 46), in addition
to narrower whorls and closer nodules on the abapical
spiral cord of the teleoconch than I. tigrina. Also, the
very small median spiral cord appears later on I. onca.
Isotriphora onca is distinguished from I. guanahacabibes
by significant differences in color (whitish in I.
guanahacabibes, or with small areas of light brown;
creamy to light brown in I. onca, with the apex whitish)
and the later development of the small median spiral
cord on 7 onca. As mentioned before, I peetersae has
about five whitish whorls on the beginning of the shell,
instead of the two/three initial whorls on 7. onca. Fur-
thermore, the dark color on the remainder whorls on
the teleoconch of 7 peetersae (creamy to light brown in
7. onca) and the earlier development of a strong median
spiral cord clearly distinguishes both species. Isotriphora
taenialba has a totally different color pattern and an
earlier development of a strong median spiral cord.
Material from the type locality (including the holo-
type) was most likely deposited by turbidity currents,
as all other shells were collected at shallower depths
(~ 85-105 m).
Distribution: Brazil: Vitoria-Trindade Chain (this study).
Genus Latitriphora Marshall, 1983
Type Species: Triphora latilirata Vereo, 1909, original
designation; Recent, southern Australia.
Diagnosis: Multispiral protoconch, embryonic shell
with hemispherical granules, larval shell with two spiral
threads and uninterrupted axial riblets; teleoconch with
three spirals cords commencing simultaneously; nodules
strongly flattened, with sharp edges that overhang sides
of spirals (based on Marshall, 1983).
Latitriphora albida (A. Adams, 1854)
(Figures 11, 22)
Triphoris albidus A. Adams (1854: 278).
Triforis (Sychar) samanae Dali (1889: 248); Dali and
Simpson (1901: 423, pi. 54, fig. 18).
Triphora samanae-. Jong and Coomans (1988: 51, pi. 34,
fig. 245).
Triphora sp. 5: Leal (1991: 124).
Latitriphora albida: Rolan and Femandez-Garces (1995:
14, figs. 29-32; 2007: 22, pi. Ill, figs. 8-9); Redfern
(2001: 67, pi. 33, fig. 280); Lee (2009: 89, text-fig.).
Triphora decorata: Absalao et al. (2006: 238, in part) non
C. B. Adams, 1850.
Type Material: Leetotype: NHM 196563. Paralectotype:
MIM 196654.
Type Locality: Honduras.
Material Examined: MNRJ 17922, Praia do Andrada,
Trindade Is., J. H. Leal, C. B. Castro and W. Zwink
coll., 17/xii/1982 [1]; MNRJ 25005, Enseada dos
M.R. Fernandes, A.D. Pimenta and J.H. Leal, 2013
Page 1 1
Figures 45-50. Isotriphora onca new species. 45, 49. Entire shell. 46, 48. Last whorl and base. 47. Adapieal view ot protoconch.
50. protoconch and beginning of teleoconch. 45-50. MNR| 16230, paratype. Scale bar = 1 mm to figures 45-46, 48-49, 100 pin to
figures 47, 50.
Portugueses, Trindade Is., 10 m depth, P. Bouehet and
J. H. Leal coll., 22/v/1987 [1]; MNRJ 25015, MD55
25-DC43 [1]; MNRJ 25016, MD55 9-DC22 [1].
Remarks: This is the only species of the genus
Latitriphora at the West Atlantic. The shell of L. albida
is very similar to that of Nototriphora decorata , hut the
teleoconch of L. albida has a more flattened profile of its
axially elongated and very close-set beads (Figure 22).
Also, die beads of the adapieal spiral cord almost reach
the beads of the abapical spiral cord of the adjacent
whorl, and smaller and paler brown blotches are present
on L. albida (Figure 1 1). The median spiral cord
of the teleoconch of L. albida begins earlier — in
the third whorl, following Lee (2009), instead of the
transition between fourth and fifth whorl on N. decorata
(Figure 25) — but it could not be determined in which
whorl it develops in the material examined of L. albida,
as all the shells had the apex broken. In the comments on
L. albida , Lee (2009) cited that this species develops the
median spiral cord later than N. decorata, when the
opposite actually occurs.
Distribution: Bermuda (Jensen and Pearce, 2009);
USA: Georgia (Rosenberg, 2009), Florida (Lee, 2009);
Gulf of Mexico (Rosenberg et al., 2009); Bahamas
(Redfern, 2001); Cuba (Rolan and Fernandez-Garces,
1995); Belize (Miloslavich et ah, 2010); Honduras (type
locality); Hispaniola (Miloslavich et ah, 2010); Puerto Rico
(Dali and Simpson, 1901, cited as Triforis samanae);
Page 12
THE NAUTILUS, Vol. 127, No. I
Colombia (Lee, 2009); ABC Islands (Jong and Coomans,
1988, cited as Triphora sanianae ); Brazil: Vitoria-Trindade
Chain (this study).
Genus Coriophora Laseron, 1958
Type Species: Coriophora negrita Laseron, 1958, orig-
inal designation; Recent, western Pacific.
Diagnosis: Paucispiral or multispiral protoconch; on
the latter, embryonic shell with hemispherical granules,
larval shell with axial riblets crossed by a central spiral
thread; teleoconch with a later development of median
spiral cord; radular formula 24-1-1-1-24, central and
marginal teeth with three cusps, lateral teeth with four
to five cusps (based on the diagnosis of Mesophora on
Marshall, 1983 and Wilson, 1993).
Synonym: Mesophora Laseron, 1958 non Borgmeier,
1937; see Ozdikmen (2013).
Coriophora novem (Nowell-Ustieke, 1969)
(Ligures 12, 23, 33)
Triphora novem Nowell-Ustieke (1969; 12, pi. 2, fig. 20);
Jong and Coomans (1988: 49, pi. 34, fig. 236).
Triphora sp. indet. A: Ode (1989: 111, fig. 6).
Marshallora sp. 1: Leal (1991: 121, pi. 16, figs. H-I).
Mesophora aff. novem: Rolan and Fernandez-Garces
(1995: 11, figs. 12-16).
Mesophora novem: Rolan and Lernandez-Garces (1995:
11, figs. 8-11; 2007: 23, pi. IV, figs. 1-5); Redfem (2001:
67, pi. 33, fig. 282); Lee (2009: 90, text-fig.); Tunnell
et al. (2010: 205, text-fig.); Garcia and Lee (2011).
Triphora decorata: Absalao et al. (2006: 238, in part) non
C. B. Adams, 1850.
Triphora melanura: Absalao et al. (2006: 238, in part)
non C. B. Adams, 1850.
Triphora nigrocincta: Absalao et al. (2006: 238); Gomes
et al. (2006: 187) non C. B. Adams, 1839.
Triphora sp.: Absalao et al. (2006: 238, in part).
Type Material: Holotype: AMNH 195419.
Type Locality: Virgin Islands.
Material Examined: IBULRJ 11992, REVIZEE C5-
42R [1]; IBUFRJ 19442, REVIZEE C5-30R [1]; IBUFRJ
19444, REVIZEE C1-C61 [4]; IBUFRJ 19452, REVIZEE
C5-49R [1]; IBUFRJ 19453, REVIZEE C1-C61 [1];
MNRJ 25006, MD55 16-DC29 [1]; MNRJ 25007, MD55
9-DC22 [1]; MNRJ 25008, MD55 35-DC59 [1]; MNRJ
25009, MD55 25-DC43 [2]; MNRJ 25010, MD55 23-
DC41 [1]; MNRJ 25011, MD55 21-DC35 [1]; MZSP
105164, Enseada dos Portugueses, Trindade Is., v/1987
[1]; MNHN, MD55 35-DC59 [1]; MNHN, MD55 20-
DC34 [1],
Other Material Examined: Sulphur Bank: IBUFRJ
13321, REVIZEE C5-13R, 16°47' S, 37°41' W, 50 m,
30A4/200 1 [1],
Remarks: This is the only species of the genus
Coriophora at the West Atlantic. The shells here identi-
fied as C. novem exhibit a unique color pattern, with the
three initial whorls of the teleoconch white and the
others with a brown background and violet/greyish nod-
ules (Figure 12). The median spiral cord appears around
the eighth/ninth whorl of the teleoconch, or even at the
end ot the seventh whorl, but always closer to adapieal
spiral cord (Figure 23).
Rolan and Fernandez-Garces (1995) distinguished
two morphs, M. novem and M. aff. novem ; later the two
morphs were named as M. novem only (Rolan and
Fernandez-Garces, 2007). The shells herein examined
are much more similar to M. aff. novem sensu Rolan
and Fernandez-Garces (1995), especially with respect to
the darker teleoconch color (except for the initial white
whorls), the same color between the spiral cords (and
also lacking a darker adapieal spiral cord) and the appar-
ent shorter length of the siphonal canal when compared
to the figures 3 and 4 (plate IV) of Rolan and Fernandez-
Garces (2007). The few shells with protoconch in the
material here examined initially present two spiral cords
in the larval shell with the subsequent disappearance of
the adapieal one later on the first whorl of larval shell,
strengthening of this cord at the end of the second whorl,
and later disappearance of this cord on the last whorl
(Figure 33), again similar to M. aff. novem of Rolan and
Fernandez-Garces (1995). The shells of C. novem from
Cuba have two spiral cords throughout the larval shell
(Rolan and Fernandez-Garces, 1995). Conservatively, we
prefer to regard this darker morph also as C. novem.
The shells illustrated by Tunnell et al. (2010) and
Garcia and Lee (2011) as M. novem appear to be its
darker morph, just like the brief description of this spe-
cies in Lee (2009). In contrast, Redfern (2001) describes
the paler morph.
Although Marshall (1983) and Wilson (1983) reported
that the protoconch of Mesophora (synonym of
Coriophora) bears only one spiral cord, Laseron (1958)
considered that the protoconch in this genus “may be
singly or doubly keeled,” which supports fire allocation
of this species to Mesophora by Rolan and Fernandez-
Garces (1995). The radular formula 24-1-1-1-24 of
Mesophora (Marshall, 1983) is different from that of
M. aff. novem (actually C. novem) in Rolan and
Fernandez-Garces (1995), 12-1-1-1-12, although the
number of cusps in each tooth cohort is the same.
Marshallora gutta Fernandes and Rolan, 1988, a spe-
cies from Cape Verde, has some similarities with the
shells herein studied. It includes protoconch features
(high density of granules on the first whorl; presence of
two spiral cords, except the initial whorls and the very
end of the protoconch), the initial whorls of the
teleoconch being white, large size of the rounded nod-
ules, suture indistinct and later development of the
median spiral cord. The main differences between the
two species include some anatomical (like the radular
formula) and shell features (C. novem has more nodulose
subperipheral and basal cords, a homogeneous color on
M.R. Fernandes, A.D. Pimenta and J.H. Leal, 2013
Page 13
base and a major extension of the white color on the
beginning of the teleoconch).
The material that Absalao et al. (2006) named as
Triphora nigrocincta (C. B. Adams, 1839) to REVIZEE-
Central (IBUFRJ 11992 to the Vitoria-Trindade Chain;
IBUFRJ 13321, to the Sulphur Bank, near Abrolhos),
and Gomes et al. (2006) reproduced, is actually formed
by two shells of C. novern.
The single shell ol C. novem collected at die station
MD55 16-DC29 (depth of 310-350 m) is probably a taph-
onomie anomaly (possibly resulting from deposition by
turbidity currents), as the remaining shells from Vitoria-
Trindade Chain and records in the literature indicate
that this species is observed only in shallower depths
(~ 20-105 m).
Distribution: USA: Florida (Lee, 2009), Louisiana
(Garcia and Lee, 2002), Texas (Tunnel] et al., 2010); Gulf
of Mexico (Rosenberg et al., 2009); Bahamas (Redfem,
2001); Cuba (Rolan and Fernandez-Garees, 1995);
Jamaica (Rosenberg, 2009); Puerto Rico (Lee, 2009);
Virgin Islands (type locality); ABC Islands (Jong and
Coomans, 1988); Brazil: Sulphur Bank and Vitoria-
Trindade Chain (this study).
Genus Monophorus Grillo, 1877
Type Species: Trochus perversus Linnaeus, 1758, by
monotypy; Recent, Europe and northern Africa.
Diagnosis: Paucispiral or multispiral protoconch; on
the latter, embryonic shell with cruciform tubercles,
larval shell with axial riblets crossed by two median spiral
cords, with or without a smooth spiral zone; teleoconch
with a later development of median spiral cord; radular
formula (8-12)-l-l-l-(8-12), central tooth with five cusps,
lateral teeth with five to seven cusps, most of marginal
teeth with four to live cusps (based on Marshall, 1983).
Synonyms: Biforina Buequoy, Dautzenberg and Dollfus,
1884; Notosinister Finlay, 1927; according to Marshall
(1983).
Monophorus olivaceus (Dali, 1889)
(Figures 13, 24, 34)
Triforis decorata var. olivacea Dali (1889: 244).
Triphora omata auct. non Deshayes, 1832: Warmke and
Abbott (1962: 76, pi. 13, fig. i); Rios (1970: 45, not
illustrated; 1975: 51, pi. 13, fig. 189; 1985: 161, pi. 53,
fig. 763; 1994: 95, pi. 31, fig. 376; 2009: 173, text-fig.);
Jong and Coomans (1988: 50, not illustrated); Absalao
(1989: 3, not illustrated); Merlano and Hegedus
(1994: 149, pi. XLVI, fig. 527); Absalao and Pimenta
(2005: 29, fig. 64); Absalao et al. (2006: 238, in part);
Gomes et al. (2006: 187;not illustrated); Santos et al.
(2007: 226; not illustrated).
Triphora (Cosmotriphora) omata auct. non Deshayes,
1832: Ode (1989: 110).
Triphora sp. 1: Leal (1991: 122).
Monophorus olivaceus : Rolan and Fernandez-Garees
(1994: 17, figs. 1-3, 6, 8, 30 MO; 2007: 23, pi. IV, figs. 23-
27; 2008: 87, figs. 4 B-F, H-K); Redfem (2001: 67,
pi. 33, fig. 284); Espinosa and Ortea (2001: 20, not illus-
trated); Lee (2009: 90, text-fig.); Garcia and Lee (201 1).
Triphora sp.: Absalao et al. (2006: 238, in part).
Cosmotriphora omata auct. non Deshayes, 1832:
Tunnell et al. (2010: 204, text-fig.).
Type Material: Lectotype: MCZ 7379.
Type Locality: Key West, west ol Florida, Gulf of
Mexico, 91 m.
Material Examined: MNRJ 25025, MD55 23-DC4I
[1]; MNRJ 25026, MD55 20-DC34 [2]; MNRJ 25027,
MD55 13-DC26 [2]; MNRJ 25028, MD55 10-DC24 [1];
MNHN, MD55 20-DC34 [2],
Remarks: In this species, the adapical and median
(appearing later) spiral cords of the teleoconch have the
color pattern of one white bead usually intercalary
between two or three brown beads, although not being
a rule, and the abapieal spiral cord is white (Figure 13).
Monophorus olivaceus has a large shell size for the family,
up to 14.77 mm and 14 whorls of teleoconch in the shells
of this study (apex missing).
Monophorus ateralhus Rolan and Fernandez-Garees,
1994 is the other species of Monophoms from the West
Atlantic. The shell of M. ateralhus has continuous dark
brown color on adapical and median spiral cords, while
M olivaceus has a discontinuous brown and white color
on these cords.
According to Faber and Moolenbeek (1991) and
Rolan and Fernandez-Garees (2008), die previous records
of Triphora omata from the western Atlantic (e.g. Abbott,
1974; Jong and Coomans, 1988; Rios, 1994) actually rep-
resent M. olivaceus , since the type material of T omata
does not match the western Atlantic species (Rolan and
Fernandez-Garees, 2008).
Triphora sp. 1 in Leal (1991) is herein considered to
be M olivaceus. However, the record of Triphora omata
by Leal (1991), based on material from Fernando de
Noronha Archipelago (off Brazil), is pending examina-
tion, since it was not illustrated.
Monophoms olivaceus is reported to show some varia-
tion in size and in hue of its brown color, as pointed in
the original description and in Rolan and Fernandez-
Garees (2007; 2008). Also, Rolan and Fernandez-Garees
(2008) included a shell from Florida with a predomi-
nant white coloration on the median spiral cord of the
teleoconch as belonging to this species.
Distribution: USA: Florida (type locality), Louisiana
(Lee, 2009), Texas (Tunnell et al., 2010, cited as
Cosmotriphora omata ); Gulf of Mexico (Rosenberg
et al., 2009); Bahamas (Redfem, 2001); Cuba (Rolan
and Fernandez-Garees, 1994); Belize (Miloslavieh et al.,
2010); Virgin Islands; St. Vincent; Grenada (Rolan and
Fernandez-Garees, 2008); Costa Rica (Espinosa and Ortea,
Page 14
THE NAUTILUS, Vol. 127, No. 1
2001); Colombia (Merlano and Hegedus, 1994, cited as
Triphora omata): Venezuela (Rolan and Fernandez-
Garees, 2008); ABC Islands (Jong and Coomans, 1988,
cited as T omata): Brazil: Amapa to Bahia (Rios, 1985,
cited as T. omata), Fernando de Noronha and Abrolhos
(Rios, 1985; Gomes et al., 2006, cited as T. omata), Vitoria-
Trindade Chain (this study), Espirito Santo (Absalao,
1989, cited as T. omata), Rio de Janeiro (Absalao, 1989;
Absalao and Pimenta, 2005; Santos et al., 2007; cited as
T. omata).
Genus Nototriphora Marshall, 1983
Type Species: Notosinister aupouria Powell, 1937,
original designation. Recent, New Zealand.
Diagnosis: Paucispiral or multispiral protoconch; on
the latter, embryonic shell with hemispherical granules,
larval shell with axial riblets crossed by two median spiral
cords, the adapieal one initially strong, but soon weaken-
ing and reappearing later; teleoconch with a later devel-
opment of median spiral cord; radular formula 9-1 -1-1-9,
central tooth with three cusps, lateral teeth with four
cusps, marginal teeth with short outer cusps and long
median cusps (based on Marshall, 1983).
Nototriphora decorata (C. B. Adams, 1850)
(Figures 14, 25, 35)
Cerithium decoration C. B. Adams (1850: 117); Clench
and Turner (1950: 272, pi. 38, fig. 2).
Triphoris variegatus A. Adams (1854: 277).
Triforis arthiiri Jousseaume (1884: 221) [nornen nudum.].
Triphora decorata : Rios (1970: 45, not illustrated; 1975:
50, pi. 13, fig. 186; 1985: 160, pi. 53, fig. 760; 1994:
94, pi. 31, fig. 373; 2009: 172, text-fig.); Abbott
(1974: 111, fig. 1133); Jong and Coomans (1988: 51,
not illustrated); Absalao (1989: 3, not illustrated); Leal
(1991: 122, pi. 16, figs. [— K); Merlano and Hegedus
(1994: 148, pi. XLVI, fig. 521); Absalao et al. (2006:
238, in part); Gomes et al. (2006: 187; not illustrated);
Santos et al. (2007: 226, not illustrated).
Triphora ( Cosmotriphora ) decorata : Ode (1989: 110,
fig. 5).
Nototriphora decorata : Rolan and Fernandez-Garees
(1994: 19, figs. 10, 14, 16, 30 ND; 2007: 24, pi. V,
figs. 1-5); Redfem (2001: 68, pi. 33, fig. 285);
Espinosa and Ortea (2001: 20, not illustrated); Lee
(2009: 91, text-fig.); Tunnell et al. (2010: 206, text-fig.).
Triphora sp.: Absalao et al. (2006: 238, in part).
Type Material: Leetotype: MCZ 186178.
Type Locality: Jamaica.
Material Examined: IBUFRJ 9313, REVIZEE Cl-
C61 [2]: IBUFRJ 19445, REVIZEE C5-44R [1];
IBUFRJ 19450, REVIZEE C5-48R [1]; IBUFRJ 19454,
REVIZEE C1-C62 [4]; IBUFRJ 19456, REVIZEE Cl-
C62 [6]; MNRJ 17924, REVIZEE C5-42R [1]; MNRJ
25001, MD55 35-DC59 [3]; MNRJ 25002, MD55 9-
DC22 [1]; MNRJ 25003, MD55 13-DC26 [1]; MNHN,
MD55 20-DC34 [3]; MNHN, Enseada dos Portugueses,
Trindade Is., v/1987 [5],
Remarks: This is a very common species in Brazil and
the only representative of the genus Nototriphora on
the West Atlantic. Its teleoconch has brown spots over a
creamy-white background (Figure 14), and the median
spiral cord appears around the fourth/fifth whorl (Fig-
ure 25). This pattern of coloration on N. decorata can
cause problems in separating this species from other
triphorids. In addition to significant differences on the
protoconch (e.g., number of spiral cords on larval shell),
the shell of Nototriphora decorata has a more axial and
irregular pattern of brown spots than Monophorus
olivaceus, which has a more spiral pattern (and its
abapical spiral cord never shows beads tinted with brown,
contrary to N. decorata). Also, the median spiral cord
appears later in the teleoconch of M. olivaceus, only
around the fifth/sixth whorl (Rolan and Fernandez-
Garees, 1994) or after the seventh whorl (Lee, 2009).
Nototriphora decorata has more numerous and smaller
nodules than M. olivaceus, contrary to Abbott (1974), as
pointed out by Ode (1989) and Lee (2009). The dif-
ferences between N. decorata and Latitriphora albida /
Cosmotriphora amoldoi were mentioned in the remarks
under the last two species.
Lee (2009) recognized intraspecific variation in
N. decorata, with a morph delicately beaded and with
blotches present at any of the three spiral cords, and
another coarsely beaded and with dark blotches restricted
to the adapieal and median spiral cords. The shells
here examined correspond to Lee’s (2009) first morph
(Figure 14). Beyond that, Rolan and Fernandez-Garees
(2008) commented on a violet coloration (instead of
brown) in some shells of N. decorata from Cuba, possibly
resultant of an ecological variation.
Nototriphora decorata is reported to have an amphi-
Atlantie distribution in some works (e.g. Leal, 1991; Rios,
1994, 2009), but it is actually due to a confusion with the
eastern Atlantic distribution of Nototriphora canarica
(Nordsieck and Talavera, 1979), initially described as a
subspecies of N. decorata and later elevated to species
by Bouchet (1985).
According to Rolan and Fernandez-Garees (2008),
Triphoris variegatus A. Adams, 1854 is a synonym of
N. decorata. Triforis arthuri Jousseaume, 1884, nomen
nudum, was proposed as a replacement name for Triphoris
variegatus A. Adams, 1854 non Cerithium variegatum
C. B. Adams. However, Cerithium variegatum C. B. Adams
does not exist, being created as result of a lapsus calami by
Jousseaume (1884) while copying die list of names of bodi
C. B. Adams and A. Adams presented by Moreh (1875).
Distribution: Bermuda (Jensen and Pearce, 2009);
USA: North Carolina (Rosenberg, 2009), Florida (Lee,
2009), Louisiana (Garcia and Lee, 2002), Texas (Tunnell
et al., 2010); Gulf of Mexico (Rosenberg et al., 2009);
M R. Fernandes, A.D. Pimenta and f.H. Leal, 2013
Page 15
Bahamas (Redfern, 2001); Cuba (Rolan and Fernandez-
Garces, 1994); |amaica (type locality); Belize; Puerto
Rico; Lesser Antilles (Miloslavich et al., 2010); Costa
Rica (Espinosa and Ortea, 2001); Panama; Colombia;
Venezuela (Miloslavich et ah, 2010); ABC Islands (Jong
ami Coomans, 1988); Brazil: Amapa to Rio de Janeiro
(Rios, 1985), Fernando de Noronha and Vitoria-Trindade
Chain (Leal, 1991; Gomes et ah, 2006).
Genus “ Triphora ” Blainville, 1828
Type Species: Triphora gemmatum Blainville, 1828,
by monotypy. Recent, Mauritius.
Remarks: Although Marshall (1983) tried to define a
diagnosis lor Triphora , this genus still has uncertain limits.
It is commonly used as a “catch-all " taxon when generic
assignment is not possible, as “ Triphora ” s.l. (Rolan and
Fernandez-Garces, 2008; Lee, 2009).
Synonyms: Tristoma Menke, 1830; Triphoris Deshayes,
1832 (orthographic variant), non Triforis Deshayes, 1834;
according to Marshall (1983).
Triphora atlantica (Smith, 1890)
(Figures 15, 26, 36)
Triforis atlantica Smith (1890: 292, pi. XXI, fig. 26).
Triphora decorat a: Absalao et al. (2006: 238, in part) non
C. B. Adams, 1850.
Triphora melanura: Absalao et al. (2006: 238, in part)
non C. B. Adams, 1850.
Triphora pulchella: Absalao et ah (2006: 238, in part) non
C. B. Adams, 1850.
Triphora turristhomae : Absalao et ah (2006: 238, in part)
non Holten, 1802.
Triphora sp.: Absalao et ah (2006: 238, in part).
Triphora atlantica : Rolan and Fernandez-Garces (2008:
146, figs. 25 A— I); Lee (2009: 92, text-fig.); Garcia
and Lee (2011).
Type Material: Lectotype and paralectotypes on
NHM 89.10.1.1874-93.
Type Locality: St. Helena Island.
Material Examined: IBUFR| 13142, REVIZEE C2-
22R [2]; IBUFR| 14422, REVIZEE C5-30R [3]; IBUFRJ
19446, REVIZEE C1-C62 [1]; MNRJ 25012, MD55
35-DC59 [1]; MZSP 105148, Enseada dos Portugueses,
Trindade Is., v/1987 [1],
Remarks: The shells examined agree with the lecto-
type designated by Rolan and Fernandez-Garces (2008), as
with the description given by the authors. The teleoconch
of T. atlantica has a brown abapieal spiral cord with
lighter nodules, while the adapical spiral cord is white
with slightly darker inter-nodular spaces (Figure 15). A
median spiral cord appears on the eight/ninth whorl
of the teleoconch (Figure 26). The larval shell begins
with one spiral cord, later developing an additional spiral
cord (tire adapical one), which fades in the last whorl
(Figure 36). Compared to a shell from Florida (figure 25-1
on Rolan and Fernandez-Garces, 2008), the develop-
ment of the adapical spiral cord of protoconch seems to
occur later on the shells of T. atlantica from the Vitoria-
Trindade Chain (Figure 36). Rolan and Fernandez-
Garces (2008) described the axial ribs as being prosocline,
but they are actually opisthocline.
Triphora atlantica has a wide distribution throughout
the Atlantic Ocean, and its type locality is an island in the
south-central Atlantic; however, few Caribbean records
are given for this species. The shells figured in Abbott
(1974) as Triphora lilacina (Dali, 1889) may be T atlantica ,
as pointed out by Rolan and Fernandez-Garces (2008)
and Lee (2009). The shell drawn in the original descrip-
tion (Smith, 1890) does not show the brown color on the
abapieal spiral cord of the body whorl, possibly due to an
oversight by the author.
Distribution: USA: Florida (Lee, 2009), Louisiana
(Lee, 2009); Puerto Rico (Rolan and Fernandez-Garces,
2008); St. Helena (type locality); Brazil: Espirito Santo
(Rolan and Fernandez-Garces, 2008), Vitoria-Trindade
Chain (this study).
Triphora ellyae Jong and Coomans, 1988
(Figures 16, 27, 37)
Triphora ellyae Jong and Coomans ( 1988: 50, pi. 34, fig. 242);
Rolan and Fernandez-Garces (1995: 13, figs. 23-25;
2007: pi. V, figs. 17-21); Lee (2009: 92, text-fig.).
Triphora orteai Espinosa (2001: 21, fig. 7).
Triphora sp.: Absalao et al. (2006: 238, in part).
Type Material: Holotype: ZMA 3.87.072.
Type Locality: Aruba/Curayao (ABC Islands).
Material Examined: IBUFRJ 14637, REVIZEE C5-
48R [2]; Baeia de Campos, Rio de Janeiro state: MNR|
15400, 22°42' S, 40°40' W, 5 m, iii/2007 (9]; MNR| 18753,
22°42' S, 40°40' W, 110-120 m, 2006 [5]; MNRJ 18955,
22°42' S, 40° 40' W, 5-10 m [2].
Remarks: As the two shells from Vitoria-Trindade
Chain were eroded, although allowing a taxonomic deter-
mination, we illustrated a shell from Bacia de Campos
(Campos Basin), deposited at the molluscan collection
of MNRJ (Figures 16, 27, 37). The shells examined have
an ovoid shape, with almost all the shell being brown,
except the white adapical spiral cord of the teleoconch
and the entire first whorl of teleoconch (Figure 16). Also,
die nodules of the median and abapieal spiral cords are
lighter than the background color. The median spiral
cord develops in the fifth/sixth whorl of the teleoconch,
and the base is short, with three smooth cords (including
the subperipheral cord) (Figure 27). The shells here
studied showed some color variation on the base, which
can be light brown to whitish, with the subperipheral
and basal cords slightly darker.
Page 16
THE NAUTILUS, Vol. 127, No. 1
Triphora ellyae can he best differentiated from
T. atlantica by the earlier development of the median
spiral cord (fifth/sixth whorl of teleoconch in T. ellyae,
eight/ninth whorl in T. atlantica), the darker brown
median spiral cord, and the smooth subperipheral cord
(very nodulose in T. atlantica). Further, the shell of
T ellyae is smaller than T. atlantica and it has a slight
ovoid shape.
The study of the animal described by Espinosa (2001)
as Triphora orteai (synonymized by Rolan and Fernandez-
Garces, 2008 with T. ellyae) may point to the generic
position of this species.
Distribution: USA: Florida, Louisiana (Lee, 2009);
Gulf of Mexico (Rosenberg et ah, 2009); Cuba (Rolan
and Fernandez-Garces, 1995); Costa Rica (Espinosa,
2001, cited as Triphora orteai)-, ABC Islands (type locality);
Brazil: Vitoria-Trindade Chain and Rio de Janeiro state
(this study).
Triphora elvirae Jong and Coomans, 1988
(Figures 17, 28, 38)
Triphora elvirae Jong and Coomans ( 1 988: 50, pi. 34, fig. 240);
Rolan and Fernandez-Garces (1995: 13, figs. 20-22;
2007: pi. V, figs. 22-23); Garcia and Lee (2011).
Cosmotriphora elvirae : Redfern (2001: 65, pi. 32,
fig. 273).
Triphora pulchella : Absalao et al. (2006: 238, in part) non
C. B. Adams, 1850.
Triphora sp.: Absalao et al. (2006: 238, in part) non C. B.
Adams, 1850.
Type Material: Holotype: ZMA 3.87.071.
Type Locality: Curasao.
Material Examined: IBUFRJ 12105, REVIZEE C5-
30 R [1]; MNRJ 12771, REVIZEE C5-30R [2]; MNRJ
25031, MD55 9-DC22 [1]; MNRJ 25032, MD55 35- DC
59 [1]; MORG 51901, REVIZEE C1-C62 [3]; MNIIN,
MD55 23-DC40 [1]; MNHN, MD55 20-DC34 [1],
Remarks: This species has a brown abapical spiral
cord, in contrast with the adapical and median white
spiral cords (Figure 17). In addition to the abapical spiral
cord, the teleoconch of T. elvirae also shows the sub-
peripheral cord and usually the final portion of the base
tinted with brown. In the shells examined, the median
spiral cord appears in die seventh whorl of the teleoconch,
showing the same size as the abapical spiral cord (the
adapical one is slightly bigger) two whorls later (Fig-
ure 28). The material here studied is more similar to the
shells of Cuba (Rolan and Fernandez-Garces, 1995) than
to the holotype, which presents the light brown color ol
the abapical spiral cord extending beyond the nodules
(Rolan and Fernandez-Garces, 1995).
Triphora elvirae is similar to T atlantica, especially in
the brown coloration on the abapical spiral cord; how-
ever, the coloration is continuous in T. elvirae (Fig-
ure 17) and restricted to the inter-nodular spaces on
T. atlantica (Figure 15). The shell of T. ellyae has an
ovoid shape and a brown median spiral cord on the
teleoconch (Figure 16), while the shell of T. elvirae is
conical and the median spiral cord of the teleoconch is
white (Figure 17).
Redfern (2001) placed this species in Cosmotriphora,
but dris generical allocation requires further investiga-
tion, as the animal and radula have not yet been seen
(Rolan and Fernandez-Garces, 1995). In addition to
the description of protoconch of this species by Rolan
and Fernandez-Garces (1995), the last whorl presents a
weakening of the adapical spiral cord (Figure 38). Actu-
ally, the sequence ol spiral cords on the protoconch of
T. elvirae is similar to that observed on Coriophora
novem (Figure 33).
The shell illustrated in Merlano and Hegedus (1994)
as Triphora intermedia (C. B. Adams, 1850) appears to
belong to T. elvirae.
Distribution: USA: Louisiana (Garcia and Lee, 2002);
Gulf of Mexico (Rosenberg et al., 2009); Bahamas
(Redfern, 2001); Cuba (Rolan and Fernandez-Garces,
1995); Belize (Miloslavich et al., 2010); Curasao (type
locality); Brazil: Vitoria-Trindade Chain (this study).
DISCUSSION
Ol the 13 species found in this study, Cosmotriphora
melanura and Nototriphora decorata were previously
reported from the Vitoria-Trindade Chain. Cosmotriphora
amoldoi, Iniforis carmelae, Iniforis pseudothomae ,
Latitriphora alhida, Coriophora novem, Triphora ellyae
and Triphora elvirae were previously restricted to the
Caribbean and Gulf of Mexico, although L. albida is also
present in Bermuda and the southeastern coast of USA.
These species are reported for the first time from Brazil.
The known ranges of Monophorus olivaoeus and Triphora
atlantica in Brazil are herein extended to the Vitoria-
Trindade Chain.
Absalao et al. (2006) and Gomes et al. (2006) listed the
occurrence of Marshallora nigrocincta (C. B. Adams,
1839), under the name Triphora nigrocincta (C. B. Adams,
1839), to the Vitoria-Trindade Chain. As mentioned
before, their taxonomic identification was erroneous,
actually corresponding to Coriophora novem. Thus, the
presence of M nigrocincta at the study site is here
invalidated. Furthermore, the occurrence of this species
in Brazil needs further investigation as it may represent a
complex of species (Bouehet, 1985).
The previous record of Iniforis turristhomae at the
Vitoria-Trindade Chain (Absalao et al., 2006) is here
invalidated, as the single shell studied is very eroded,
and it could not be positively identified.
Until now, only three species of Isotriphora were rec-
ognized in the western Atlantic (Rosenberg, 2009) and
If worldwide (Rosenberg, 2011). Isotriphora tigrina new
species is currently restricted to the Vitoria-Trindade
Chain and Bahia state and Isotriphora onca new species
to the Vitoria-Trindade Chain. The records of the two
M.R. Fernandes, A.D. Pimenta and J.H. Leal, 2013
Page 17
speeies above represent the entirety of occurrences of
lsotriphora for Brazil. The non-planktotrophic inode of
development suggested by the protoconch in this genus
indicates a greater possibility for endemism.
With the present study, the number of species of
Triphoridae in the Vitoria-Trindade Chain is increased
from six to 15 and in Brazil from 17 to 26. Many other
species still need to be described or otherwise reported
from Brazil. Further scrutiny of this and other relatively
understudied elements are certain to greatly improve
our knowledge of the western Atlantic malacofauna in
coming years.
ACKNOWLEDGMENTS
We are grateful to Harry Lee (Florida, USA), who
reviewed the manuscript and lent material of his private
collection; to Philippe Bouchet for the invitation that
allowed one of us (JHL) to participate in the cruise
MD55; to Dr. Guilherme Muricy (MNR|) and Juliana
Segadilha (IBUFRJ), for their criticisms and suggestions
on the manuscript; to Dr. Carlo Magenta (MZSP), who
photographed some shells; to Dr. Ricardo Absalao
(IBUFRJ), Cleo Oliveira (IBUFRJ) and Dr. Luiz Simone
(MZSP), for loan of material; to Amanda Veiga, for
operating the SEM at the Departamento de Inverteb-
rados (MNR|); and to CNPq (Conselho Nacional de
Desenvolvimento Cientifico e Tecnologico), for provid-
ing a fellowship to the first author.
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THE NAUTILUS 127(1): 19-28, 2013
Page 19
Diel variation affects estimates of biodiversity and abundance
of nudib ranch (Gastropoda) faunas
Yen-Wei Chang1
Institute of Marine Biology and
Asia-Pacific Ocean Research Center
National Sun Yat-sen University
Kaohsiung 80424, TAIWAN
[email protected]
Tsen-Chien Chen
Department of Leisure Management
Minghsin University of Science
and Technology
Hsinchu 30401, TAIWAN
Richard C. Willan
Museum and Art Gallery of the Northern Territory
Darwin, Northern Territory 0801, AUSTRALIA
Hin-Kiu Mok Ming-Hung Yu
Institute of Marine Biology and Flyfish Diving Ltd
Asia-Pacific Ocean Research Center Taitung 95141, TAIWAN
National Sun Yat-sen University
Kaohsiung 80424, TAIWAN
ABSTRACT
With approximately 3000 species worldwide, the Nudibranchia
is numerically die largest group of sea slugs (Gastropoda:
Heterobranchia). In most previous surveys on diversity and
abundance of nudibraneh faunas, researchers only undertook
daytime surveys and neglected nighttime surveys lor practical
reasons. In order to assess whether such estimates are accurate
if they are based only on daytime surveys, we undertook repli-
cated diel (i.e., day and night) surveys for 12 consecutive
months (from December 2009 to November 2010) at Shilang
Marine Reserve, Green Island (Lyudao), Taiwan. We analyzed
the species accumulation curves, diversity indices, and similar-
ity of species composition as statistical estimates of overall
biodiversity and species abundance. Fifty-three species of
nudibranchs were recorded during all our surveys, but the
cumulative number of species showed no sign of reaching a
plateau at the end of our survey. We recorded 34 species during
the daytime surveys and 42 species during the nighttime sur-
veys. The mean number of species observed during the day and
night was 6.3 and 8.9, respectively. Phi/llidiella pustulosa was
the species observed most often during the daytime surveys,
while Tritonia sp. 1 (an undescribed species) was the most
abundant species at night. Species composition was clearly
different between day and night. Our results for this particular
region of the tropical Indo-Pacific Ocean, which is adjacent to
the “Coral Triangle” region of global maximum biodiversity of
nudibranchs, lead us to conclude that estimates of faunal diver-
sity and species composition within local communities (rich-
ness) are strongly influenced by whether surveys are
conducted during the day or the night. Furthermore, regard-
less of when and how such surveys are conducted in tropical
seas, the estimates of biodiversity they produce will inevitably
hill far short of total biodiversity.
1 Current Address: Museum and Art Gallery of the Northern
Territory, Darwin, Northern Territory 0801, AUSTRALIA
Additional Keywords: nudibraneh, biodiversity, day-night quan-
titative surveys, Green Island (Lyudao)
INTRODUCTION
The Nudibranchia is the most morphologically com-
plex and ecologically diverse group of the marine
Heterobranchia. It comprises about 3000 species world-
wide, though probably less than half of them are pres-
ently formally named (Wiigele and Willan, 2000; Willan,
pers. obs.). All nudibranchs are carnivorous, being pred-
ators on a wide variety of organisms — sponges, hydro-
zoans, scleractinian corals, octocorals, scyphozoans,
bryozoans, kamptozoans, crustaceans, ophiuroids, ascid-
ians, fishes, and even other sea slugs. Most nudibranchs
are highly specialized for feeding on only one particular
species of prey item. Therefore, the presence and abun-
dance of nudibranchs may act as a useful indicator of the
overall diversity of marine organisms in an ecosystem
(Lock et ak, 2010). The best way to measure the pres-
ence and abundance of nudibranchs is to study their
biodiversity both systematically and quantitatively. Bio-
diversity is the full variety of genetic diversity, species
diversity, and ecosystem diversity, which are usually mea-
sured by species richness and evenness. Biodiversity is
relevant to conservation. Understanding spatial and tem-
poral variation of species composition are important for
effective biodiversity conservation strategies (Roberts
and Gilliam, 1995).
In terms of the geographical distribution of nudi-
branchs, the number of species is far greater in the trop-
ical zones of the world than in temperate zones (Table 1),
and this conclusion is generally consistent across the
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THE NAUTILUS, Vol. 127, No. 1
Table 1. The total number of nudibranch species recorded at different latitudes of the world (arranged from north to south). The
dashed lines separate the temperate and subtropicaPtropical faunas. The data for number of species includes both described and
undescribed species.
globe. However, latitude alone does not completely explain
global biodiversity as some regions in the tropical Indo-
Pacific Ocean (in particular the “Coral Triangle”) have
biodiversity of nudibranchs orders of magnitude higher
than other tropical regions such as the Caribbean Sea and
tropical western Africa (Gosliner et ah, 2008).
Tl lis information on large scale biodiversity and pres-
ence of species within local communities (richness) in
nudibranch faunas are accumulating gradually, particu-
larly so for the megadiverse “Coral Triange” region and
the regions immediately adjacent to it such as the South
China Sea, Melanesia, Micronesia, and tropical northern
Australia. What is emerging already is that estimates of
biodiversity and species richness are heavily reliant upon
the way the data are collected, both temporally and spa-
tially. Several factors affect the occurrence of species and
hence their chance of being recorded - sampling effort
and technique, experience of the investigators (in partic-
ular dreir “search image” for cryptic and microscopic
species), season, time of day, environmental conditions
(such as surge), and the heterogeneity of the habitat
itself. As one example of such a quantitative survey,
we cite Johnson’s (1983) pioneering study of two spe-
cies of nudibranchs (i.e., the species that he called
Chromolaichma youngbleuthi (Kay and Young, 1969)
(presently Glossodoris rufomarginata (Bergh, 1890))
and Hypselodoris sp. 1 (presently Hypselodoris peasei
(Bergh, 1880)) on the subtidal reef at Hawaii where he
found that the number of individuals was not stable
throughout the year. Furthermore, the actual duration
of the survey is very important to the final estimate of
biodiversity. The longer the sampling period, the more
the cumulative number of species will increase, but this
increase is only very gradual as nudibranchs are inher-
ently rare in time and space (Marshall and Willan, 1999).
For example, Gary Cobb (pers. obs.) and his colleagues
have presently recorded 381 species of nudibranchs in
the same area in southern coastal Queensland (Table 1),
yet even more importantly, even after eight years of con-
tinuous (i.e., weekly) surveying, they are still discovering
further species at an average of 1.18 additional species
every week.
Since nudibranchs are rather slow-moving animals, one
does not expect them to show strong daily patterns of
activity; one assumes all species are always present and
always visible, and this assumption underlies surveys for
biodiversity and richness. However, careful observations
made in situ underwater show that some species do, in
fact, possess daily patterns of (feeding and locomotor)
activity; Bertsch and Johnson (1981: 66, 107) and Johnson
(1983: 3; 1989) report that one species of Hypselodoris
(presently known as H. peasei) in Hawaii was observed
during the day crawling or feeding in groups of up to 30 or
more individuals (its colloquial name being “locust”) on its
specific sponge food ( Luffariella sp., family Thorectidae).
However, just before sunset all the individuals left their
food sponge and moved one after the other into a nearby
crevice where they hid for the night. This is an example
of a diurnal (i.e., day-active) species. Conversely, some
species are reported as exclusively nocturnal (Johnson
1989; Behrens, 2005: 20, 142; Holland, 2008), for example
the tropical Hawaiian dorids Chromodoris sp., Aldisa
pikokai Bertsch and Johnson, 1982 and Sclerodoris
paliensis Bertsch and Johnson, 1982 (Johnson, 1989). In
our experience working with temperate and tropical
Pacific nudibranch faunas, the “classic” group of nocturnal
Y.-W. Chang et ah, 2013
Page 21
nudibranchs comprises members of the genus Janolus
(family Zephyrinidae), all of which are cryptic. During
the day, individuals hide deeply within the arborescent
Catenicellidae bryozoans, on which they feed. They are
truly impossible to find. But at night they emerge to
graze on the tips of these bryozoans and are quite
conspicuous (Miller and Willan, 1986).
Our study, conducted in southeastern Taiwan, investi-
gates such temporal variation quantitatively by compar-
ing the composition of the nudibranch fauna by day and
by night, and season by season, and it investigates quan-
titatively how predictive such a survey is for estimation of
overall nudibranch biodiversity at one particular locality.
In Taiwan, the only similarly rigorous precursor is that by
Su et al. (2009). These workers, all of whom were “ama-
teur” nudibranch researchers, undertook 17 consecutive
monthly surveys during the daytime at the Pescadores
Islands (Penghu) off the west coast of the mainland.
They concluded that species composition showed no
obvious pattern between seasons or months.
MATERIALS AND METHODS
Study Site: Taiwan is a large island located in the
tropical Western Pacific Ocean bathed by the Kuroshio
Current that originates southeast of the Philippine archi-
pelago and flows northward to Japan. The Kuroshio
branches south of Taiwan. The major branch goes through
the Philippine Sea past the eastern coast of Taiwan, and
the weaker branch goes through the Taiwan Strait past the
western coast of Taiwan. The Kuroshio Current is charac-
terized by high water temperature, high salinity and low
nutrients (Chen et al., 1994), physical parameters most
favorable for the development of coral reefs. Green Island
(Lyudao), a small remnant volcano located soudieast of
the Taiwanese mainland, lies in the path of the main
branch of the Kuroshio Current. Uniquely for Taiwan,
die study site at Shilang on the eastern coast of Green
Island (22° 39. 37' N, 121° 27.45' E) has numerous coral
reefs and coral patches (Figure 1), and since 2003 has
been declared a marine fishery reserve. Since then, all
types of fishing have been forbidden. Shilang is also an
ideal site for a full-year survey due to its fortuitous geo-
graphic location — in winter the strong northeastern mon-
soon is blocked by the central chain of mountains on
Green Island, and in summer Shilang is sheltered from
the southern monsoon because it is on the western coast
of Green Island. The seas are generally calm year round.
Field Survey Methods: During the period between
December 2009 and November 2010, we undertook
12 consecutive monthly diel surveys of nudibranchs by
scuba diving at Shilang. All of the surveys were conducted
using shore-entry and the divers had to walk about
30 meters across the rocky intertidal zone carrying their
scuba gear to access the water. Therefore, for safety
monthly surveys were always timed when the seas were
calmest. Most of the surveys were conducted during
periods of complete calm with low waves, but some months
there was a strong surge under water.
Figure 1. Map showing the location of Green Island off the southeastern coast of Taiwan. The enlargement on the right indicates
the study site at Shilang (22° 39.37' N, 121° 27.45' E).
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THE NAUTILUS, Vol. 127, No. 1
Figures 2-5. Techniques used for data collection at the study site: 2. One of the individually marked rocks (rock number 2).
3-4. Divers using a 2 m tape to measure a marked rock. 5. Detail of habitat recording using a 20 x 20 cm quadrat.
In order to study species composition systematically,
we marked eight rocks (Figure 2), within a total area of
600 irf (i.e. a rectangle 30 m in length from shore by
20 m width) prior to the commencement of our survey.
On our survey transect we always swam straight out to the
farthest rock (no. 8) and started the survey there allowing
approximately 70 minutes for each survey. On each
monthly dive, two divers searched thoroughly around the
marked rocks. One diver (the senior author of tins paper)
recorded all species and individuals and took photographs
of them all. His buddy diver (the fifth author of this
paper) was assigned to take photographs and search for
nudibranchs in caves. All recordings were written on
waterproof paper. All our surveys were conducted in
depths shallower than 12 m. On each occasion, seawater
temperatures were recorded in dive computers (Suunto
Mosquito). To avoid disturbance and to conform to the
conservation status of the Shilang Marine Reserve, neitiier
diver collected any specimens on these survey dives.
Waterproof torches (Saekodive AL-29) were used at night.
We delayed night dives until after 20 hr because of previ-
ous observations that nocturnal nudibranchs did not
emerge until then.
Habitat Analysis: In October 201 1, prior to the start of
the quantitative survey for nudibranchs, we measured the
height, length, anti width of each of the eight study rocks
by taking a photograph with a 2 m measuring tape as a
scale (Figures 3, 4). Measurements were computed by
linage J software (1.43u, National Institutes of Health,
USA). At the same time, we also undertook a quantitative
description of the habitat types on each rock. We placed
a 20 x 20 cm quadrat frame (Figure 5) randomly on 10
sites on each rock and took a photograph of each quadrat
using Vidana 1.0.1 beta software. It should be noted that
the cover of algae was particularly low at that time of year
because it was late autumn. We observed that the algae at
tliis site have an annual fluctuation, with a bloom during
spring to summer and a reduction in density and com-
position during winter. We analyzed 10 photographs on
each rock using Vidana L.0.1 beta software. Descriptive
information for the eight study rocks is given in Table 2.
Y.-W. Chang et al., 2013
Page 23
Table 2. Comparison of habitats sampled (eight separate rocks) and type of habitat. The proportion of each habitat type (and
percent cover) are given for each of the four habitat types.
Species Identification and Cumulative Curve: The
third author identified every individual from the photo-
graphs taken during the surveys. All individuals were
identified. After the identification process, we created the
data matrix with the sample time in rows and the species
in columns in an Excel spreadsheet (Appendix 1. Matri-
ces of species composition data for the day and night
surveys at the study site at Shilang, Green Island, Taiwan,
posted at http://nautilus.shellmuseum.org). The species
cumulative curves were plotted based on samples and
individuals for the diel surveys. Rarefaction curves, an
analytical method representing re-sampling of all individ-
uals, showed smoothed cumulative number of species
obtained per sample (Gotelli and Colwell, 2001). We
used Estimates version 8.2.0 (Colwell, 2009), which com-
puted rarefaction curves based on the samples.
Biodiversity Index and Species Similarity: In order
to compare the species richness and richness between the
day and night surveys, we used the Shannon-Wiener H'
Diversity^ Index (Pielou, 1975) for species richness:
H' = — (Pi In Pi), where Pi is the proportion of indi-
i=l
viduals of each species to the total number of individ-
uals. Pielou’s Evenness Index was used for assessing
species evenness (Pielou, 1966): J=H'/H'max, where H'
is Shannon-Wiener H' Diversity Index, H'maxAhe maxi-
mum of Shannon-Wiener IT diversity indexes. To com-
pare the species composition of the diel surveys, we log
transformed the number of individuals and used the
Bray-Curtis Coefficient to estimate the similarity. A den-
drogram was constructed using the unweighted pair-
group clustering method using arithmetic averages
(UPGMA clustering method) (Romesburg, 1984). All
the biodiversity and species similarity data were com-
puted by the Primer 6.0 software package (Clarke and
Gorley, 2006).
Statistical Analyses: To compare the number of spe-
cies, number of individuals, Shannon-Wiener H' Diver-
sity Index and Pielou’s Evenness Index between day and
night, we used the Wileoxon Signed Ranks Test to com-
pare the two sets of data that were arranged in the
Microsoft Excel spreadsheet. We also used the Kruskal-
Wallis Test to compare abundances between the seasons
for soft coral-feeding and sponge-feeding groups of nudi-
branchs, i.e., Tritoniidae and Chromodorididae plus
Phyllidiidae, respectively.
RESULTS
Habitat Analysis: The habitat types dominating the
eight rocks at our study site at Green Island (Lyudao)
were algae, (hard plus soft) corals, sponges and bare rock
surfaces. Detailed information about the habitats them-
selves is given in Table 2. Among the rocks, most had a
high percent cover of coral, ranging from 44.0 to 93.2 %.
The percentage cover of algae ranged from 1.5 to 28.6 %.
As mentioned above, it was possible that the lowest per-
centage of algae (1.5 %) was because the date chosen to
record the habitats was in late autumn (31 October 201 1)
and also because the particular rock (number 7) on which
this lowest algal cover recorded was located in relatively
deep water. For sponges, the percentage cover ranged
from 3.3 to 17.8 %. For bare substrates, the percentage
cover ranged from 2.0 to 21.7 %.
Species Identification and Cumulative Curve: Over
the year of the survey, we recorded a cumulative total of
53 species of nudibranchs (13 families) including six pos-
sible new species. Thirty-four species (10 families) were
recorded during the day and 42 species (13 families) at
night. The mean number of species recorded for each
2009 2010 2010 2010 2010 2010 2010 2010 2010 2010 2010 2010
Dale
Figure 6. Number of species for all surveys. □ represents
daytime surveys. ■ represents nighttime surveys.
Page 24
THE NAUTILUS, Vol. 127, No. 1
45 r
45
40
3 35
I 30
0
d 25
z
> 20
1 15
3
u
5 •
0I i 1 1 ' ' ' ' ' ' ' '
1 2 3 4 5 6 7 8 9 10 11 12
Samples
Figure 7. Relationship between the cumulative number
of species and monthly samples taken during all surveys.
□ represents daytime surveys. ■ represents nighttime
surveys represents daytime rarefaction. represents
nighttime rarefaction.
monthly survey was 6.3 during the day (range 2-12 spe-
cies) and 8.9 species during the night (range 2-17 species)
(Figure 6). There was no significant difference in the actual
number of species between the day and the night
(Wilcoxon Signed Ranks Test, df = 1 1, N = 1 2, P = 0. 1 16).
The mean number of individuals recorded for each
monthly survey was 8.9 individuals during the day (range
4-18 individuals) and 20.2 individuals during the night
(range 3-36 individuals). The number of individuals
observed at night was significantly higher than during the
day (Wilcoxon Signed Ranks Test, df = 11, N = 12, P =
0.016).
The highest number of species of nudibranehs we
recorded during the day was 12 species in April 2010.
The highest number of species we recorded during the
night was 17 species in November 2010. The lowest
number of species we recorded during the day was two
species in March and August 2010. The lowest number
of species we recorded during the night was two species
in January 2010.
The sample-based cumulative curves showed 34 spe-
cies were recorded during the day versus 42 species at
night (Figure 7). The figure for diversity had still not
reached a plateau after 12 months, which means that even
after twelve surveys (i.e., 24 collections of data) some
species had still not been observed. In the sample-based
rarefaction curves, the daytime curve was a good match
for its rarefaction curve. However, the nighttime curve
was far removed from its rarefaction curve (Figure 7).
Rarefaction curves were used to randomize all the species
in all the samples. If the cumulative curve was much lower
than the rarefaction curve, it represented the aggregation
of species during the samples. On the individual-based
cumulative curve, more individuals were recorded at
night and the slope obviously showed the species did not
reach a plateau (Figure 8).
Across all our surveys, the number of individuals
recorded at night was consistently greater than that for
the day except for December 2009, January 2010 and
Figure 8. Relationship between the cumulative number of
species and individuals sampled during all surveys. □ repre-
sents daytime surveys. ■ represents nighttime surveys.
April 2010. The highest number of individuals during
the day was 18 individuals in April 2010 and during the
night it was 36 individuals in July 2010. The lowest num-
ber of individuals during the day was three individuals in
August 2010 and during the night it was three individuals
in January 2010. We counted a total of 107 individuals
during the day and 249 individuals at night across all
surveys. The statistical analysis showed that the number
of individuals recorded at night was higher than during
the day. The main factor responsible for this skew that
affects the analysis was the appearance of Tritonia sp. 1
during the night. We recorded 111 individuals of this
species at night, however only two were recorded during
the day across all surveys.
Biodiversity Index and Species Similarity: The
Shannon-Wiener H' Diversity Index showed no clear
pattern between day and night surveys (Figure 9) and
the statistical results confirmed there was no significant
difference between day and night (Wilcoxon Signed Ranks
Test, df - 11, N =12, P = 0.433). Pielou’s Evenness Index
showed a lower value at night than during the day from
April to August (Figure 10). This is because large num-
bers of Tritonia sp. 1 appeared and aggregated at night,
which produced a lower value of evenness. There was no
significant difference in Pielous Evenness Index between
day and night samples (Wilcoxon Signed Ranks Test,
Figure 9. Shannon-Wiener H' Diversity Index for all surveys.
□ represents daytime surveys. ■ represents night time surveys.
Y.-W. Chang et al., 2013
Page 25
Figure 10. Pielou's Evenness Index for till surveys. □ repre-
sents daytime surveys. ■ represents nighttime surveys.
20 ■
F
^ 60 -
00000000000)0)000000000000
oooooooooooooooooooooooo
CSICMCNCMCMC\ICNJCslC\IC\ICMCNCNJCMCNJtNCM(SI(NC'JOJCNCNJfM
< O
(D 0} HI 1) ID O
O <
Figure 1 1. Dendrogram lor hierarchial clustering oi all sur-
veys (12 months with daytime and nighttime surveys), using
group-average linking of Bray-Curtis Coefficient. □ represents
daytime surveys ■ represents night time surveys.
df = 1 1, N = 12, P = 0.875). The analysis dendrogram for
hierarchial clustering showed the species compositions
between night and day were different (Figure 11). Most
of the species recorded during the day and the night
formed separated groups, except for two night surveys in
December 2009 and February 2010, during which the
most frequently recorded nudibranchs belonged to the
family Phyllidiidae. However, there was no clear pattern
between months and seasons (Figure 11). The most fre-
quently recorded species overall was Phyllidiella pustulosa
(Cuvier, 1804) (Figures 12-15). It appeared 17 times dur-
ing 24 surveys (12 months at daytime and 5 months at
nighttime). The most abundant species overall was Tritonia
sp. I (Figure 16), which was recorded 1 1 times during 24
surveys (one month at daytime and 10 months at night-
time). In terms of abundance. 111 individuals of Tritonia
sp. 1, were recorded at night and only two individuals were
recorded during the day. Most of them were recorded on
rock number eight on top of the octocoral Tubipora musica
Linnaeus, 1758 (family Tubiporidae) (which is their food
source) and the alga Caulerpa serrulata (Forsskal)
J.Agardh, 1837 (Figure 17). During the field surveys, we
were amazed how cryptic this species was despite it being
so common; we found that even on the same coral patch,
we could not find any Tritonia sp. 1 in the afternoon but a
few hours later at night, we recorded numerous individ-
uals. Some of them were mating on the top of the Tubipora
musica (Figure 18), while others were aggregated on the
adjacent bare rock (Figure 19).
As no nudibranchs feed on algae, we used soft coral
feeders (Tritoniidae) and sponge feeders (Chromodorididae
plus Phyllidiidae) to study the relationship between
seasons and abundances. The abundance of the sponge
feeders was not significantly different between the four
seasons (Kmskal- Wallis Test, df = 3, N = 12, P = 0.08).
The average number of individuals in winter and autumn
(10.7 and 12.7, respectively) was higher than spring and
summer (5.3 and 5.7, respectively). However, the abun-
dance of the soft coral feeders was highly significantly
different between the four seasons ( Kruskal-Wallis Test,
df = 3, N = 12, P = 0.03). The summer had the
highest average number of individuals (29.3), followed by
autumn (27.7), and spring (12.2). In winter, we recorded
the lowest average number of individuals (3.7). The
sponge feeders were more abundant in winter and
autumn. In contrast, the soft coral feeders were more
abundant in summer and autumn.
DISCUSSION
Comparing the number of species recorded during the
day versus during the night over the entire twelve
months of the survey, we found that those recorded
during the day were numerically greater than those at
night on only two occasions (January 2010 and April
2010), whereas the number of species recorded at night
was greater than during the day on ten occasions. How-
ever, the statistical analysis showed that the average
number of species was not significantly different
between the day and the night.
As explained above, we had marked eight rocks
from near the shore to some distance offshore. The most
visible difference to us between these rocks between the
day and the night was the appearance at night of larval
fish that aggregated in front of the torch, in addition to
nocturnally active crustaceans and polyehaetes. Fewer
nudibranchs were recorded on the surveys when there
was a strong surge (January 2010 night, March 2010 day,
and August 2010 day), confirming the observations noted
anecdotally at other locations (see Introduction). During
all our surveys, the visibility was always greater than 10
m because Green Island was bathed by the clear
Kuroshio Current and there are no rivers on the island
that would introduce terrigenous silt into the marine
environment and reduce visibility. The only occasions
when visibility dropped significantly were after typhoons.
The coral cover was stable during the whole year. The
algal density was higher in the spring and summer, and
lower during autumn and winter as noted above. During
Page 26
THE NAUTILUS, Vol. 127, No. 1
Figures 12-15. The most commonly recorded species during our survey, Phyllidiella pustulosa, and its habitat. All photographs
taken during the day. 12. Individual feeding on its food sponge. 13. Same turned over showing its oral tube everted (circled).
14. Individual then quickly retracted its oral tube; 15. The feeding scar left by this individual on the sponge (arrowed). All
photographs by the senior author.
the surveys, we recorded more phyllidiids during the day
(55 individuals cumulatively) than during the night (30
individuals cumulatively). However, some species were
recorded more frequently during the night, e.g. Ill indi-
viduals of Tritonia sp. 1 were recorded during the night
in comparison to two individuals during the day. This
species was obviously active during the night.
In a broader view, there are several factors that affect
the recording of the cumulative number of species of
nudibranchs at a particular locality. The first is latitude;
tropical regions support far higher numbers of species
than temperate regions (Table 1). The second is survey
effort; the longer the period of the survey, the greater the
cumulative number of species will be. Our study showed
that the number of species had still not reached a plateau
even after one year. In fact, records accumulated for
Taiwan over a decade reveal the biodiversity of nudi-
branchs to be over 580 species (pers. observ.), so we only
recorded less than 10% of the total fauna. This very
gradual accumulation of species is consistent with what
is known from nudibranchs elsewhere (e.g.. Heron
Island, Great Barrier Reef). The third is the time of the
survey; most of the previous surveys were conducted in the
day which may neglect the nocturnal species. Searches
conducted over a long period only in daytime may
never record some strictly nocturnal species. The last
factor is search experience (i.e. acquiring a “search
image”); with time, observers gradually improve their
recording skills as they become more adept at seeing
cryptic species, and at recognizing small juveniles and
species less than 5 mm long.
There was no obvious pattern in the Shannon-Wiener
H" Diversity Index during all our surveys. In Pielou’s
Evenness Index, most of the months showed that the
night data were higher than the day except for the period
from April to August 2010. During these five months, we
recorded at least nine individuals of Tritonia sp. 1. July
2010 had the lowest value because 27 individuals of
Y.-W. Chang et al., 2013
Page 27
Figures 16-19. In situ photographs of Tritonia sp. 1 and its habitat, all taken at night. 16. Individual showing its gill plumes and
finger-like oral tentacles. 17. Four individuals aggregated on the habitat. 18. Two individuals mating on top of the habitat 19. An
aggregation of eight adult individuals. Note the presence of an individual of the aeolid nudibranch Phijllodesmium briareum (circled)
on the bare rock some distance away from the closest Tritonia sp. 1. Photographs 16 and 19 by the fifth author. Photographs 17 and 18
by the senior author.
Tritonia sp. 1 were recorded. The species similarity
showed that there was no obvious pattern in temporal
change (month and season), which agrees with the
results of Su et al. (2009). However, when we compared
the species composition between day and night, we
observed that the composition at night was different to
that of the day; all the nocturnal species were clustered
together except for December 2009 and February 2010,
on which more phyllidiids were recorded than on other
night surveys.
We observed the number of individuals of Tritonia
sp. 1 during the night was different over the years; we
recorded one individual from December 2009 to Feb-
ruary 2010; 22 individuals from March to May 2010;
58 individuals from June to August 2010; and 30 individ-
uals from September to November 2010. These data
indicate this species is most abundant during summer
and least abundant during winter. However, percent
cover of its food source, Tubipora musica , was very stable
during all of our surveys. Therefore, further investiga-
tions are required to discover the factors that affect the
population cycle of this species of Tritonia.
ACKNOWLEDGMENTS
Our sincere thanks are extended to Ming-Hsueh Chen,
Tzu-Ming Cheng, Cheng-Lien Hou, and Shih-Wei Su for
help with scuba diving and site analysis at Green Island. We
thank colleagues for information about diversity of
nudibranchs in their regions of the tropical Pacific Ocean:
Scott Johnson (Marshall Islands), Jean-Franyis Herve
(New Caledonia), Gary Cobh, and David Mullins
Page 28
THE NAUTILUS, Vol. 127, No. 1
(Sunshine Coast, southern Queensland, Australia). This
study was funded by the Asia-Pacific Ocean Research
Centre, National Sun Yat-sen University, through a
Ministry of Education Grant aimed at the Top University
Plan (C030200). The first author was funded to visit the
Museum and Art Gallery of the Northern Territory
during 2011/2012 by the National Science Council of
Taiwan under its Study Abroad Program (NSC 100-2917-
1-1 10-005). Two reviewers offered constructive criticisms
for which we are most grateful.
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THE NAUTILUS 127(l):29-35, 2013
Page 29
Occupancy of strata of plant height and plant substrate by land snail
Gastropoda assemblages at Escaleras de Jaruco, Mayabeque, Cuba
Maike Hernandez Quinta
Division Zoologia, Invertehrados
Institute de Eeologia y Sisteinatica
Carretera de Varona km 3 lA Capdevila, Boyeros
11900, La Habana, CUBA
in aike@eeol ogia. cu
ABSTRACT
The study was carried out in three low hills (Beluea, La
Chirigota, and La Jaula) in the Escaleras de Jaruco-Tapaste-
Cheche, Mayabeque province, Cuba. On each low hill, ten
fixed quadrats of 3 x 3 m were selected and plant strata were
identified (strata I: 0-lm of height; II: 1.01-2 m; III: more than
2.01m) in addition to the substrate ol the plants used lor most
representative species of land snail assemblages. At the three
outcrops the lowest stratum (I) ol the vegetation presented
the greatest values ol absolute abundance. During the rainy
months, Chondropoma pictutti was the most abundant species
in stratum I followed by strata II and III respectively, while
during the less rainy months this species was only observed in
strata I and II. Helicina aspersa, C. auberianum , Jeanneretia
bicincta, and Zachrysia auricoma were observed in strata I
and II during both rainy and less rainy periods. The branches
and trunks contained higher species abundance and frequency
of individuals than leaves, which was true ol all the species with
the exception ol J. bicincta Individuals frequency on the upside
or downside surface ol leaves was very variable. Some species
showed small diflerences in the percentage found on both sides
of a leaf ( H . aspersa and Z. auricoma ) and other ones showed
larger differences (Z onitoides arboreus, C. pictum and J. bicincta).
Additional Keywords: Vegetation, low hills, tropical karstic
forest, Protected Area
INTRODUCTION
Land snails rank second in number of species in terres-
trial ecosystems, being outnumbered only by arthropods
(Russell-Hunter, 1983). Land snails are important not
only numerically, but also ecologically, because of their
role in nutrient cycling as detritivores, herbivores, and
carnivores (Purchon, 1977). Despite their diversity and
ecological importance, most of the published informa-
tion on terrestrial mollusks concerns systematics; rela-
tively little is known about terrestrial molluscan ecology
(Alvarez and Willig, 1993). This has improved in recent
times with an increase in ecological studies of land snail
communities since 2000 (Schilthuizen, 2011).
The majority of current papers dealing with habitat
preferences and distribution of terrestrial mollusks
concentrate on large-scale relationships, based on com-
parisons of molluscan communities from geographi-
cally distant regions (Stanisic, 1997; Hermida et al.,
2000; Muller et al., 2005; Cameron and Pokryszko,
2007; Stanisic et al., 2007; Correa-Sandoval et al , 2009;
Nekola 2002, 2009). Studies on small-scale snail distribu-
tion and microhabitat choice were scarce until fairly
recently. More recently, many field and experimental
studies have examined ecological differences in the
relative abundance and species composition of the com-
munities among and between microhabitats (Kozlowsld
and Kaluski, 2004; Kappes, 2005; Kappes et al., 2006,
2009; Perea et al., 2007; Solymos et ah, 2006, 2009;
Cejka and Hamerlik, 2009; Szybiak et ah, 2009).
What little knowledge exists on microhabitat choice
in Cuban land snail species is based on arboreal species.
The majority of studies have been on charismatic spe-
cies and populations; examples are Bidart and Espinosa
(1989) on Polymita picta nigrolimbata Torre, 1950,
Con/da alauda (Ferussac, 1821) and Caracolus sagenum
rostrata (Pfeiffer, 1847); Bidart et al. (1992) y Fernandez
et al. (2000) on Polymita muscarum (Lea, 1834); Alvarez
and Berovides (1989) and Bidart et al. (1992) on Liguus
fasciatus achatinus Clench 1934; Bidart et al. (1992) on
Hemitrochus lucipeta (Poey, 1854). The majority of these
works are focused on a only a few species. In contrast,
this study attempts to examine the entire land snail
community by identifying and analyzing the variations
of the use of plant height strata and substrate of the
plants in a natural Cuban habitat, namely the protected
area Escaleras de Jaruco-Tapaste-Cheche (Natural pro-
tected landscape), Mayabeque, Cuba.
MATERIALS AND METHODS
The study was conducted in the Escaleras de Jaruco-
Tapaste-Cheche Protected Area (23°00W N, 82°0T27" W;
23°03,27" N, 82°08/20" W) which is predominantly a
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THE NAUTILUS, Vol. 127, No. 1
tropical karstic forest. This park is situated in western
Cuba. The area comprises a series of raised limestone
outcrops and is a calcareous elevation of Miocene age
about 15 km in length, 3-7 km in width, and approxi-
mately 300 m a.s.h, where karst funnels, cones, and
domes are observed (Canas and Ysalgue 1978). The
study was carried out in three low hills: Beluca, La
Chirigota and La Jaula (Figure 1). Tropical karstic
forest is the vegetation type present in each elevation.
This includes evergreen and/or semi-deciduous for-
ests. At the highest elevations there are thickets with
weedy species.
The study was carried out monthly in the period of
abundant rains of August to November (November is
usually considered a less rainy month, but during this
study it behaved like a rainy month) and the less rainy
period of January to April 2010. In each locality, ten
fixed plots of 3x3 m separated by more than 20 m
were selected. Within each plot vegetation (branches-
trunk and leaves), the land snails were examined only
in the sub-canopy. The average of shrubs/trees was eight
for plot and 88 % of canopy.
The absolute abundance and the frequency of
occurrence (FO) of species were determined. Based
Figure 1. Geographic location of karstic low hills (limestone outcrops) selected for the study of terrestrial arboreal snails of the
protected area “Escaleras de Jaruco-Tapaste-Cheche”, Mayabeque, Cuba.
M.H. Quinta, 2013
Page 31
on occurrence, the species were classified into one of
four arbitrary categories: rare (FO < 25 %), occasional
(25% < FO > 50%), uncommon (50 % < FO > 75 %)
or common (75 % < FO > 100 %). We established
three strata (stratum I: 0-lm of height; II: 1.01-2 m;
III: more than 2.01m) of the vegetation to analyze
vertical distribution of the species. The absolute abun-
dance in the three strata in each elevation and in the
most representative species was compared by means of
a one-way AN OVA with Tukey HSD test as a post-hoc
test. GraphPad Prism 5 was used for these analyses.
RESULTS
Vertical Distribution of the Land Snail Community
in Vegetation: In both wet and dry periods, the
majority of species were rare (Table 1). In the rainy
months, Chondroponia pictum was uncommon in Beluca
and occasional in La Chirigota and La |aula, while in
the less rainy months it was rare or was not present at
La Chirigota. Jeanneretia bicincta showed a low fre-
quency in La Chirigota, while in La Jaula this species
was occasional in the rainy months. The rest of the spe-
cies was present in very low frequency but it is perti-
nent to highlight that Helicina aspersa in La Chirigota
and La |aula, and Zachrysia auricoma in La Jaula,
showed elevated values among the rare species (Table 1).
In the three elevations, the lowest stratum (stratum I)
of the vegetation was found to have the highest value of
absolute abundance of individuals (Ligure 2). In Beluca,
abundance did not show significant differences between
the three strata, in La Chirigota and in La Jaula strata I
and III were different (Ligure 2). The absolute abun-
dances of Chondroponia auberianuni ( F (2, 26) = 0.74,
P = 0.48, N = 30) and C. pictum ( F (2, 26) = 2.27,
P = 0.12, N = 30) in the three strata showed no
significant differences, while for Helicina aspersa
( F (2, 26) = 3.48, p = 0.04, N = 30), Jeanneretia
bicincta (F (2, 26) = 5.04, P = 0.01, N = 30) and
Zachrysia auricoma ( F (2, 26) = 3.68, P = 0.04, N =
30), the abundance in the three strata showed sig-
nificant differences. In H. aspersa , strata II and III
showed differences, while for the other two species
strata I an d III were different.
During the rainy months, Chondroponia pictum was
more abundant at stratum I with 110 individuals fol-
lowed by strata II and III with 76 and 34, respectively.
Table 1. Occurrence (%) and classification (arbitrary categories) ol the species present in the vegetation in three low hills
(limestone outcrops) of Escaleras de Jaruco, Mayabeque, Cuba. The general values (independently of the periods) are shown as well
as those for Rainy Months and Less Rainy Months.
Page 32
TIIE NAUTILUS, Vol. 127, No. 1
Strata
Figure 2. Absolute abundance of land snails in the three
strata of vegetation (one: I, two: II, three: III) ol three low
hills (limestone outcrops) in Escaleras de Jaruco during
the 2009-2010. The top of the bar shows the mean and
the vertical lines in the shape of “y” represent the standard
error. Letters represent significant differences of abundance
between stratums according to the analysis of multiple com-
parisons (Tukey HSD).
Table 2. Frequency of occurrence (FO) (%) and absolute
abundance of the species representative of the tropical karstie
forest in three low hills (limestone outcrops) of Escaleras de
Jaruco, Mayabeque, Cuba, during 2009-2010.
four individuals of this species were observed in strata I
and II, respectively. Helicina aspersa was observed with
greater abundance during the rainy months in strata I
and II, with seven individuals, while only one indi-
vidual was observed in stratum III; during the less rainy
months six individuals were observed in stratum II,
four in stratum I and one in stratum III. Chondropoma
auberianum, Jeanneretia bicincta , and Z achrysia auricoma
were only observed during the period of rainy months
in strata I and II.
Use of Vegetative Substrate: Absolute abundance
(25 ± 7 SD) and frequency of appearance (41%) of
the individuals present in the branches-trunk of vege-
tation, was higher than abundance (6 ± 1 SD) and fre-
quency of appearance (18 %) in the leaves. This was
true of most species (Table 2), with the exception of
Jeanneretia bicincta that was more abundant and fre-
quent in the leaves.
The appearance of the snail on either the upside or
downside surface of the leaves showed variation among
the different species. Four species were only observed
on the upsides of the leaves (. Eurycampta bonplandi,
Emoda submarginata, Chondropoma auberianum,
Liguus fasciatus) and two species only on the downside
( Alcadia hispida , Setipellis stigmatica ), each with only one
individual. Another six species were observed on both
sides of leaves, and Oleacina straminea presented one
only individual on each side. Helicina aspersa (down-
side, 42 % and upside 58 %) and also Zachrysia
auricoma (downside, 57 % and upside 43 %) showed
little difference in the percentage use of particular leaf
sides. In contrast, Z onitoides arboreus (downside, 67 %
and upside 33 %), C pictum (back, 17 % and bundle 83
%) and Jeanneretia bicincta (downside, 63 % and
upside 37 %) exhibited large percentage differences in
the use of particular leaf sides.
DISCUSSION
Determining the use of the environmental resources
by land snails is important in supplying information for
MIL Quinta, 2013
Page 33
a species survival. An essential resource for land snail
is food and protective refuges against dehydration
(Cook, 2001; Luchtel and Deyrup-Olsen, 2001). To
avoid dehydration, their survival depends in large part
not only on their behavior, but also on their remark-
able ability to tolerate dehydration through suites of
physiological mechanisms that help conserve and acquire
water (Luchtel and Deyrup-Olsen 2001).
The results of this study reveal perhaps behavioral
strategies to minimize dehydration given that the highest
abundance was observed in stratum I, primarily in shrubs
that are a part of the undergrowth, in which favorable
conditions of temperature and relative humidity exist.
Low abundance in stratum III of the vegetation may be
due to the presence of microclimatic conditions unfavor-
able for the survival of some of the snails observed in
this study. In some other species that inhabit different
environments, different behaviors has been described.
For example, in semiarid regions, species such as Cepaea
nemoralis climb up the vegetation, where temperature
is cooler, to escape the heat of the ground (Jaremovie
and Rollo, 1979). A similar behavior has been observed
in Theba pisana (Cowie, 1985) and Liguus fascial us
(Fernandez, 2005), which, during the summer when
temperatures increase, use the highest vegetation strata.
Chondropoma pictum and Helicina aspersa (representa-
tive species) do not exhibit similar seasonal behaviour
since they showed larger values of abundance in strata I
and II during the rainy and less rainy months.
Cook (2001) proposes that one such adaptation against
dehydration is the use of efficient microhabitats where
the risk of dehydration is lower. Nevertheless, the study
reveals some differences in species use of vertical strata
of vegetation particularly between Chondropoma pictum ,
Jeanneretia bicincta, and Helicina aspersa. The cause
of the low occurrence in this microhabitat is possibly
because these species also use rocks and litter as micro-
habitats, where they can persist with elevated frequency
and abundance, perhaps using microhabitats to rest,
find refuge, or forage (Hernandez, 2011).
Several authors have reported that Liguus fasciatus
uses strata higher in the vegetation (Fernandez, 2005;
Alvarez and Berovides, 1989) and this may explain the
low observed frequency of this species in the present
study, since very high strata may influence a species
detectability. Another aspect for consideration is an
individual's size; possibly due to the large size of Liguus
fasciatus, the species may be more susceptible to desic-
cation in the higher strata of the vegetation than the
smaller species. Perrott et al. (2007) demonstrated
experimentally that the individuals’ size is an important
factor in the desiccation stress in the juvenile snails of
Cantareus aspersus and adults that had smaller shells.
Helicina aspersa, known as an arboreal species
(Espinosa and Ortea, 2009), has also been reported in
other microhabitats such as rocks and litter (Hernandez,
2011). However, Perez (1999) observed this species
in the vegetation in the locality Hornos de Cal, Sancti
Spfritus, in very low frequency (10 %), even with a
greater sampling effort. Chondropoma pictum, despite
being well-represented in the vegetation, is reported
by Hernandez (2011) as a habitat generalist.
This study found that the majority of the species
use trunk-branches, except for jeanneretia bicincta ,
which is why it is necessary to determine experimen-
tally what induces the use of a determined vegetative
substrate. The fundamental use of trunk-branches has
been reported for other arboreal Cuban species like
Polymita muscarum (Fernandez et al., 2000), Polymita
venusta, Hemitrochus sp. (Santos, 2000) and Liguus
fasciatus (Alvarez and Berovides, 1989). Possibly the
presence of mollusks on the upside and downside
surfaces of the leaves represent a protective strategy
against adverse climatic condition, predators, or simply
a place for protected rest. In the present study, we
observed a similar frequency of use of the upside and
downside surfaces of leaves by some of the species
observed, but in species such as Chondropoma pictum
and /. bicincta , we observed differential use of only one
side of the leaf. In C. pictum the upside was used with
greater frequency, while, / bicincta used the back in
greater frequency.
Perhaps the variations of humidity between low and
high strata are the causes of the distribution of the
land snails in the vegetation, because the majority of
individuals occurred in the first two strata. This result
could be influenced by the abundance of operculated
gastropods ( Chondropoma pictum, C auberianum, Helicina
aspersa, Alcadia hispida, and Torrella inmersa ), which
are very susceptible to changes in relative humidity
(Russell Hunter, 1964). As a survival strategy, these spe-
cies frequent the lowermost zones of the vegetation
near the litter, where humidity is conserved. It is
possible that pulmonate species such as Jeanneretia
bicincta and Z achrysia auricoma do not display simi-
lar strategies (to the operculated gastropods), but
they frequented low shrubs. The presence of sources
of foods (lichens and mosses) in branches-trunks may
have conditioned a greater percentage of individuals
to these substrates in contrast to the leaves, or perhaps
branehes-leaves are used as refuges. This research show
that not all the species in the assembly of land snails
studied present similar strategies of survival, where the
micro-environmental conditions, physiological constraints,
and distribution of the food items influence their occur-
rence. These studies are not totally conclusive because
other parameters such as inter-specific competition may
also play a role in patterning the distribution of species
along the distinct vertical strata.
ACKNOWLEDGMENTS
To the staff of the protected area of Escaleras de Jaruco,
to afford the accomplishment of this investigation in the
area. To Betina Neyra, Annery Serrano, Anay Serrano,
Ormaily Madruga and Rayner Nunez for their help
in tlie fieldwork. To Kathryn E. Perez of the Department
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THE NAUTILUS, Vol. 127, No. 1
of Biology, University of Wisconsin - La Crosse, for the
critical reading and valuable comments and realized
suggestions. The fieldwork was partially funded by the
Cuban project “Creation, Conservation and Manage of
Zoological Collections” of the Ecology and Systematic
Institute, Havana, Cuba.
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THE NAUTILUS 127(l):36-39, 2013
Page 36
First occurrence of putative hybridization in the genus
Lobatus (Caenogastropoda: Strombidae)
Gijs C. Kronenberg
Mollusca Collection
Naturalis Biodiversity Center
P.O. Box 9517
NL-2300 RA Leiden
The Netherlands
ABSTRACT
A putative hybrid Lobatus gigas x L. gallus is reported. It is
compared with its putative parental species.
Additional Keyword: Caribbean
INTRODUCTION
Kronenberg (2008: 332, table) presented a review of
known putative hybrids within the gastropod family
Strombidae. A total of sixteen different combinations of
putative parental species, all of Indo-Pacific origin, were
indicated. Some of these were known from only one spec-
imen, but others from over 100 specimens. Presumed
cases of hybridization of Strombidae within the Caribbean
are anecdotal, see for instance Conch-L archives (www
listserv.uga.edu), and restricted to Strombus pugilis
Linnaeus, 1758 x S. alatus Gmelin, 1791. Images of
such putative hybrids are published on the website of
Bill Frank (www.jaxshells.org/strombsslax.htm).
Reed ( 1995a, b) observed frequent interspecific mating
between co-occurring species of Caribbean Strombidae
with a remarkably little discrimination in choice of part-
ner, i.e., males have frequently been observed attempting
to mate with non-eonspeeifie females. However, only
matings of S. pugilis x S. alatus produced fertile eggs.
In May 2012 I received an e-mail from Harry G. Lee,
from Jacksonville, Florida, who drew my attention to a
putative hybrid Lobatus gallus (Linnaeus, 1758) x L. gigas
(Linnaeus, 1758) in the private collection of Pete Stimpson,
in Loudon, Tennessee. Subsequently, Dr. Stimpson made
the specimen available for examination. The results are
presented herein.
SYSTEMATICS
Family Strombidae Rafinesque, 1815
Genus Lobatus Swainson, 1837
Type Species: Strombus bituberculatus Lamarck,
1822 (= Strombus raninus Gmelin, 1791) by monotypy.
Pliocene to Recent, Caribbean.
Discussion: The genus name Lobatus first appeared
in 1837 in an anonymously compiled catalogue. I redale
(1921: 208) concluded that authorship of that catalogue
should be attributed to Swainson. Therefore all newly
introduced names in this catalogue should be attributed
to Swainson. In his broad concept of the large, broad-
winged Strombidae, Abbott (1960) synonymized Lobatus
with Tricomis Jousseaume, 1886 (Type species: Tricomis
tricomis {— Strombus tricomis [Lightfoot], 1786) by
monotypy). Jung and Heitz (2001) argued that the intro-
duction of Lobatus should be considered as a historical
accident and considered the name unavailable.
The name Lobatus was further discussed by Kronenberg
and Lee (2007) and Landau et al. (2008), who concluded
that Lobatus was the first available name for the group
of large, broad winged Strombidae from the Caribbean
and Panamic Fauna province, rejecting the conclusions
reached by Jung and Heitz (2001). Based on molecular
(Latiolais et ah, 2006) and morphological (Kronenberg,
2008) data it was concluded that Lobatus could not be
considered a junior synonym of Tricomis.
Lobatus gallus (Linnaeus, 1758) x Lobatus gigas
(Linnaeus, 1758)
(Figures 1-5)
Description: Shell large, length from apex to tip of
anterior canal 131 mm, consisting of about 9 teleoconch
whorls and broad flaring outer lip (wing) with tip
reaching beyond apex. Protoconch broken off and first
teleoconch whorls badly eroded; of preserved teleoconch
whorls second with axial costae, irregularly interrupted
by broader costae, about three per whorl, here inter-
preted as varices. Costae gradually changing into
pointed knobs on shoulder with abapical part of base
adjacent to suture. Some knobs distinctly broader as a
G.C. Kronenberg, 2013
Page 37
Figures 1-5. Putative hybrid Lobatus gallus x L gigas. Puerto Rico, off Vlayagiiez at about 4.6 m, September 1972, Pete Stimpson
collection, unnumbered, height from apex to tip af anterior canal: 131 mm. 1. Apertural view. 2. Dorsal view. 3. Lateral view, note
relatively large shoulder knob. 4. Apical view, same scale as Figures 1-3. 5. Operculum, 40 mm length.
continuation of pattern of varices, last varieal knob on
antepenultimate whorl. Knobs gradually increasing in
number and size, penultimate whorl with eleven knobs.
On last whorl, knobs increasing in size at faster rate,
penultimate knob largest, slightly past mid dorsum, last
knob much smaller. Spiral sculpture much less promi-
nent, on first teleoconch whorls eroded, on other spire
whorls about seven discernible. Last whorl with promi-
nent spiral cords abapically to shoulder knobs. Outer lip
widely dilated, with adapieal broadly rounded point, rim
of lip slightly sinuous sloping down to point of attach-
ment with ventral side of last whorl. Lateral part of outer
lip slightly undulating on abapical half, corresponding
with spiral cords on dorsal side, stromboid notch well
developed. Adapertural side of outer lip smooth, colu-
mella smooth, apart from two narrow, low folds running
into aperture. Anterior canal slightly developed. Callus
on ventral side extended, thin, translucent, with gray-
metallic haze on central part. Color of shell cream with
irregular mottling of brown, rim of adapertural side of
outer lip cream, aperture light pink; base of columella
cream, adapieal part light pink. Operculum stromboid,
smooth edged, 40 mm in length. Soft parts unknown.
Material Examined: One specimen from Puerto
Rico, west coast, off Mayagtiez at about 4.6 m, Septem-
ber 1972, Pete Stimpson collection, unnumbered.
Distribution: So far only one specimen known, for
data see above.
Page 38
THE NAUTILUS, Vol. 127, No. 1
Remarks: In general the specimen is closest to L. gallus,
both in general shape as well as size. Although usually
smaller, the largest L. gallus measured attained a size of
134 mm from apex to tip of anterior canal. The hybrid
specimen differs from L. gallus by its shorter anterior
canal; the relative larger size of the penultimate shoul-
der knob on the last whorl; the shape of the posterior
lobe of the outer lip, which is much more narrow in
L. gallus- and the color of the aperture. In L. gallus the
rim of the adapertural side of the outer lip and the
columella are cream to pinkish light orange colored,
but deep within the aperture it is always white. In the
specimen here discussed, the rim of the outer lip is
white, but deep within the aperture as well as the colu-
mella, it is pink, a color never seen in L. gallus.
Lobatus gigas grows to a much larger size, although
dwarfs of about 150 mm are known. The hybrid speci-
men differs from L. gigas by its smaller size; slightly
longer anterior canal, the shape of the posterior lobe
of the outer lip; and the much larger knobs on the
penultimate whorl.
DISCUSSION
The occurrence of the traits in the specimen reported
herein could also be explained by developmental muta-
tions within a specimen of one the putative parental
species. However, in such a case one might expect that
not so many shifts in shell characters reported herein as
compared to one of the supposed parental species would
occur, or one might expect one or more unique traits, i.e.
shell characters that cannot be attributed to a character
of one of the parental species or being intermediate
between the supposed parental species. As this is not
the case, and the specimen exhibits characters that are
more or less intermediate between L. gallus and L. gigas ,
sometimes closer to either of these supposed parental
species, I conclude that at present we are dealing with a
putative hybrid L. gallus x L. gigas.
Several cases of putative hybridization within the
Strombidae were summarized by Kronenberg (2008).
All of these have been solely based on shell morphology
and this case is no exception. The remarks by Kronenberg
(1993), i.e., that the anatomy of the presumed hybrid was
never examined nor that mating between supposed paren-
tal species was ever observed, are in fact still valid for
all cases of presumed hybridization within Strombidae.
Exceptional are observations made by Reed (1995a, b),
who observed reproductive behavior of several species
currently assigned to Lobatus , viz. L gigas; L gallus;
L. costatus (Gmelin, 1791) andL. raninus (Gmelin, 1791).
She observed mating between several species of Lobatus
both in the field and in the laboratory. Apart from the
observation that males occasionally attempted copulation
with a male that already was engaged in copulation with
a female, mating with non nonspecific females was also
observed. Reed (1995a) reported the following combi-
nations: L. gallus <S x L. costatus $ ; L. raninus 3 x
L. costatus $; L. raninus 3 x L. gallus 9; and L. gallus 3 x
L. raninus 9- Reed (1995b) also observed one case of
multiple partners, i.e. copulation of L. costatus 9 with
L. gallus 3 and L. raninus 3- None of the females
spawned after these copulations. Although L. gigas was
present during these experiments, no mating between
L. gigas and L. gallus or any other species of Lobatus,
was reported.
It should be noted that "In a separate collection of
Strombus pugilis and S. alatus individuals mated at
random with no apparent distinction between species,
and females spawned fertile egg masses, regardless of
the male involved.” (Reed, 1995a: 329). However, we
don’t know anything about further results, i.e., whether
the eggs hatched or not. Within the genus Lobatus,
L. gallus and L. gigas appear to be the most closely
related species according to the eladogram presented
by Latiolais et al. (2006). These authors did not incor-
porate L. goliath (Schroter, 1805) in their analysis. In
an analysis based on anatomy (Simone, 2005), L. gigas
and L. goliath group as sister taxa, which in turn are
sister taxa to L. costatus and L. gallus.
Recently, Oxenford (2011) described a small popula-
tion of L. goliath off Barbados, reported from two local-
ities on the western coast of that island. In Barbadian
waters, several species of Strombidae were already
reported, among them L. gigas (Oxenford, 201 1). Oxenford
already noted that, although it is quite unlikely that
L. goliath would have a serious negative impact on the
marine ecosystem, it would be desirable to monitor
this / these population(s) to record any developments.
Taking into consideration the relatively large number of
putative cases of hybridization in Strombidae (Kronenberg,
2008), the close relationship between L. goliath and
L. gigas (Simone, 2005), and the behavior of Strombidae
as reported by Reed (1995a, b), we cannot exclude the
possibility that hybridization will occur. Of course,
these species should also occur syntopic, i.e. not only
living in the same geographical range but also sharing
the same habitat.
ACKNOWLEDGMENTS
I thank Dr. Harry G. Lee, Jacksonville, Florida, for drawing
my attention to the specimen discussed herein; Dr. Pete
Stimpson, Loudon, Tennessee for sending the specimen on
loan; and Dr. Gregory Herbert, University of South Florida,
for additional comments on possible results of hybridiza-
tion. Dr. Lee also reviewed a first draft of the manuscript
and provided useful comments. Two anonymous reviewers
provided useful comments that further helped to improve
this paper. Mr. Eeleo Kruidenier, Naturalis Biodiversity
Center made the photographs. I thank my partner, Ms.
Marianne Matthijssen for her abiding support.
LITERATURE CITED
Abbott, R.T. 1960. The genus Strombus in the Indo-Pacific.
Indo-Pacifie Mollusea 1(2): 33-146.
G.C. Kronenberg, 2013
Page 39
Concli-L [Archives], www.listserv.uga.edu (last accessed: 23
July 2012)
Iredale, T. 1921. Molluscan nomenclatural problems and solu-
tions. Proceedings of the Malacological Society of London
14: 198-208.
Frank, W. (webmaster) www.jaxshells.org/strombsslax.htrn
(last accessed: 23 July 2012)
Jung, P. and A. Heitz. 2001. The subgenus Lentigo
(Gastropoda: Strombidae) in tropical America, fossil and
living. The Veliger 44: 20-53.
Kronenberg, G.C. 2008. An intergeneric hybrid (Gastropoda:
Caenogastropoda: Strombidae) with remarks on the
subdivision of Indo-Pacific Tricornis. Basteria 72:
331-343.
Kronenberg, G.C. and H.G. Lee. 2007. Genera of American
strombid gastropods (Gastropoda: Strombidae) and
remarks on their phylogeny. The Veliger 49: 256-264.
Landau, B.M., G.C. Kronenberg, and G. Herbert. 2008. A
large new species of Lohatus (Gastropoda: Strombidae)
from the Neogene of the Dominican Republic, with notes
on die genus. The Veliger 50: 31-38.
Latiolais J. M., M S. Taylor, K. Roy and M E. Hellberg. 2006.
A molecular phylogenetic analysis of strombid gastropod
morphological diversity. Molecular Phylogenetics and
Evolution 41: 436-444.
Oxenford, II A. 2011. A giant leap? Goliath conch a Brazilian
endemic species is found in Barbados. Journal of the
Barbados Museum and Historical Society 57: 192-201.
Reed, S. E. 1995a. Reproductive anatomy and biology of the
genus Strombus in the Caribbean: I. Males. Journal of
Shellfish Research 14: 325-330.
Reed, S. E. 1995b. Reproductive anatomy and biology of the
genus Strombus in the Caribbean: II Females. Journal of
Shellfish Research 14: 331-336.
Simone, L.R.L. 2005. Comparative morphological study of rep-
resentatives of the three families of Stromboidea and the
Xenophoroidea (Mollusca, Caenogastropoda), with an
assessment of their phylogeny. Arquivos de Zoologia 37:
141-267.
[Swainson, W.] 1837. Catalogue of die Foreign Shells in the pos-
session of the Manchester Natural History Society, arranged
according to the system of Lamarck. 99 pp. [not seen]
New distribution record for the rare limpet
A croloxus coloradensis (Henderson, 1930)
(Gastropoda: Acroloxidae) from Montana
Tl le Rocky Mountain Capshell, Acroloxus coloradensis
(Henderson, 1930), the only North American member
of the basommatophoran family Acroloxidae, is broadly
distributed across southern Canada and south into the
Rocky Mountains in the USA (Turgeon et ah, 1998; Lee
and Ackerman, 2000). Despite its wide geographic
range, A. coloradensis has been documented from < 30
locations, mostly in British Columbia, Alberta, Ontario,
and Quebec (Lee and Ackerman, 2000; Anderson, 2005).
Relict populations of A. coloradensis in the USA have
been documented from only 6 sites in Colorado and
2 sites in Glacier National Park (Glacier NP), Montana
(Anderson, 2005; Ellis et ah, 2004). In Glacier NP,
A. coloradensis was first reported from Lost Lake
(Figure 1; Russell and Brunson, 1967). A second popu-
lation in the park was discovered in Trout Lake in 2001
(Ellis et ah, 2004). In both lakes, A. coloradensis was
found primarily under rocks and other cover objects.
Figure 1. Documented locations of the Rocky Mountain
Capshell ( Acroloxus coloradensis ) in Glacier National Park,
Montana. The location of the newly discovered population is
marked with a triangle. The Lost Lake and Trout Lake
populations are marked with circles.
We report the incidental collection of a single Acroloxis
coloradensis specimen (5mm total length) from a small,
unnamed beaver-dammed lake (1.04 ha, 1391 m eleva-
tion; N 48.996°, W -13.678°) in the Belly River Valley in
Glacier NP (Figures 1-2). The limpet was collected
during a site survey for amphibians on 30 July 2012.
The shallow margins of the lake are dominated by
organic substrates that support extensive emergent veg-
etation. Electrical conductivity of the lake was 370 pS,
which is high compared to other small lakes in Glacier NP
(BRH, unpublished data). A secondary goal of these
surveys was to collect < 30 large gastropods that get
captured while using dip-nets (4.75 mm mesh) to sample
for amphibian larvae. The A. coloradensis specimen,
characterized by its flat shell and apex that points to
the rear and left, was found among several preserved
Planorbella trivolvis and Phijsella sp. Because our survey
protocol for amphibian larvae does not include turning
cover objects and it is unlikely the limpet would have
been picked from the bottom of a net, we suspect the
A. coloradensis specimen was along with Planorbella
trivolvis and Phijsella sp.
Acroloxus coloradensis is classified as vulnerable by the
IUCN because of the small number of documented pop-
ulations and their isolation from one another (Bogan, 1996).
The rarity of this species makes the discovery of new
populations important for accurate risk assessments.
Figure 2. Rocky Mountain Capshell ( Acroloxus coloradensis)
collected from the Belly River Valley, Glacier National Park,
Montana. Total length of the specimen was approximately 5 mm.
B.R. Hossack and R.L. Newell, 2013
Page 4 1
Our record of A. coloradensis in the northeast corner
of Glacier NP in a habitat that differs from the other
2 locations in the park suggests it may be more widely
distributed in the area than records indicate. Addi-
tional work will be required to determine the relative
abundance of A. coloradensis at tliis newly documented
population and to determine if it is present in neigh-
boring water bodies.
ACKNOWLEDGMENTS
We thank P. Scarr and S. Dykman for their careful field
work and S. Schwartz for help with the photograph.
R.T. Dillon (College of Charleston) confirmed identifica-
tion of the specimen. Reviews by A.E. Bogan, B. Roth,
and J. Giersch improved the manuscript. This work was
funded by USGS Amphibian Research and Monitoring
Initiative (ARM I) and was conducted under research
permit GLAC-201 l-SCI-0025. This note is ARMI
product no. 430.
LITERATURE CITED
Anderson, T. 2005. Hocky Mountain capshell snail
( Acroloxus coloradensis): a technical conservation assess-
ment. USDA Forest Service, Rocky Mountain Region
Available: http://www.fs.fed.us/r2/prcjects/scp/assessments/
rockymountaincapshellsnail.pdf. (31 January 2013).
Bogan, A.E. 1996. Acroloxus coloradensis. In: IUCN 2012.
IUCN Red List of Threatened Species. Version 2012.1.
<www.iucnredlist.org> . (16 September 2012).
Ellis, B. K., L. Marnell, M. A. Anderson, J. A. Stanford,
C. Albrecht and T. Wilke. 2004. Status and ecology of a
glacial relict mollusk, the Rocky Mountain capshell limpet
(Acroloxus coloradensis ), in relation to the Limnology of
Lost Lake, Glacier National Park, Montana (USA). Open
File Report 186-05. Prepared for die National Park Service,
Glacier National Park, West Glacier, Montana by Flathead
Lake Biological Station, University of Montana. 63pp.
Henderson, | 1930. Ancylus coloradensis , new name for
A. hendersoni Walker, 1925, not 1908. The Nautilus 44: 31.
Lee, J. S., and J. D Ackerman. 2000. Freshwater molluscs at
risk in British Columbia: Three examples of "risk”. Pro-
ceedings of a Conference on the Biology and Management
of Species and Habitats at Risk, 2000, Kamloops, B.C.,
Volume One, B.C. Ministry of Environment, Lands
and Parks, Victoria, B.C. and University ol the Cariboo,
Kamloops, B.C. 490pp.
Russell, R. H. and R. B. Brunson. 1967. Acroloxus coloradensis
from Montana. The Nautilus 81: 33.
Turgeon, D.D., J.F. Quinn, Jr., A.E Bogan, E.V. Coan, F.G.
Hochberg, W.G. Lyons, PM. Mikkelsen, R.J. Neves,
C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema,
F.G. Thompson, VI. Vecchione, and J.D. Williams. 1998.
Common and scientific names of aquatic invertebrates
from the United States and Canada: mollusks, 2nd edition
American Fisheries Society, Special Publication 26,
Bethesda, ix 4- 526 pp.
Blake R. Hossack
U.S. Geological Survey
Northern Rocky Mountain Science Center
790 East Beckwith Avenue
Missoula, Montana, 59801 USA
[email protected]
Robert L. Newell
102 South 8th Street
Dayton, WA 99328 USA
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THE0NAUTILUS
Volume 127, Number 2
June 21, 2013
ISSN 0028-1344
CONTENTS
Yuri I. Kantor Antarctica, where turrids and whelks converge: A revision of
M.G. Harasewych Falsimohnia Powell, 1951 (Neogastropoda: Buccinoidea) and a
description of a new genus 43
Alisa Kosyan Aidemofusus ignotus new genus and new species, a new buccinid
Yuri I. Kantor (Gastropoda: Neogastropoda) from the North Pacific Ocean 57
Michael J. Bolton A new species of Striostrea (Bivalvia: Flemingostreidae) Irom the
Roger W. Portell upper Pliocene and lower Pleistocene strata of Florida, USA 65
G. Thomas Watters Rediscovery of Choanopoma? smithianum Pfeitfer, 1866
Jozef Grego (Annulariidae) from Haiti and designation of a neotype, with the
Jozef Steffek description of two new species of Weinlandipoma Bartsch, 1946 78
Orso Angulo-Campillo Marionia kinoi (Nudibranchia: Tritoniidae): A new species
Hans Bertsch from the tropical eastern Pacific 85
Research Note
Timothy A. Pearce
Megan E. Paustian
Omphalotropis ilapiryensis, a replacement name lor O. costidata
Emberton and Pearce, 1999 (Gastropoda: Littorinimorpha:
Assimineidae)
90
THE NAUTILUS 127(2):43-56, 2013
Page 43
Antarctica, where turrids and whelks converge: A revision
of Falsimohnia Powell, 1951 (Neogastropoda: Buccinoidea)
and a description of a new genus
Yuri I. Kantor
A.N. Severtzov Institute of Ecology
and Evolution
Russian Academy ol Sciences
Leninskyj Prospect, 33
Moscow 119071, RUSSIA
M.G. Harasewych 1
Department of Invertebrate Zoology, MRC 163
National Museum of Natural History
Smithsonian Institution
P.O. Box 37012
Washington, DC 20013-7012 USA
[email protected]
ABSTRACT
A study of the type material of Antarctic species of conoideans
described by Herman Strebel revealed that four species,
all originally described in the genus Bela Gray, 1847
(Mangeliidae), are referable to the superfamily Buccinoidea
based on radular morphology. Three of the species: B fluvicans,
B. minor , and B anderssoni are transferred to the genus
Falsimohnia , which is here reviewed. The new buccinoidean
genus Strebela is proposed for the species originally described
as Bela notophila. Of the five species of the conchologically
similar genus Pareuthria Strebel, 1905 that have been recorded
within the Antarctic Convergence, two, Pareuthria innocens
(Smith, 1907) and P hoshiaii Numanami, 1996, are transferred
to the genus Falsimohnia based on radular morphology. The
appropriate generic allocation of Pareuthria plicata Thiele, 1912
is not yet clear; however, major differences in radular morphol-
ogy exclude it from the genus Pareuthria. The tax a Pareuthria
valdiviae (Thiele, 1925) and Pareuthria turriformis Egorova,
1982 are presently known only from their shell morphology.
We suggest that they will be referred to other genera when
anatomical material becomes available, and that the genus
Pareuthria wall be limited to the Magellanic region.
Additional Keywords: Gastropoda, Conoidea, Buccinoidea,
Antarctic Convergence, biogeography, morphological convergence
INTRODUCTION
Despite numerous and ongoing research programs
that have been sampling areas of the Southern Ocean
for more than a century, many of the species of
Neogastropoda described from Antarctic waters are
known from a small number of specimens, and a consid-
erable proportion of the taxa have yet to be re-sampled
since their original description. Therefore, it is not sur-
prising that some of the smaller species, especially those
1 Author for correspondence
having narrow distributions, remain poorly studied. Among
the insufficiently studied groups within Antarctic waters
are the Conoidea. This superfamily is one of the most
diverse clades of Neogastropoda (Bouchet, et ah, 2002;
Bouchet, et ah, 2009), particularly in the tropics. The
classification of the superfamily Conoidea has recently
been revised based on radular morphology (Tucker and
Tenorio, 2009) and molecular phylogeny (Puillandre
et ah, 2011), and now includes 15 families (Bouchet
et ah, 2011). Thirteen of these families had previously
been included in a single family, the Turridae, which
emerged as paraphyletic in the phylogenetic analysis.
These 13 families are collectively referred to here as
“turrids”. Currently, about 460 eonoidean genera and
subgenera are recognized, 358 of which belong to “turrid”
families and encompass more than 4000 named Recent
species (Tucker, 2004), with many hundreds of species
still unnamed (Bouchet, et ah, 2009).
The diversity of turrids decreases significantly in high
latitudes. Only about 40 species are known to inhabit the
waters of the Arctic Ocean (Kantor and Sysoev, 2006).
The diversity of turrids in Antarctic and subAntarcic
waters is also very low. At present, 37 named species have
been recorded from the Southern Ocean (Engl, 2012),
although there are still new species to be discovered and
named. Remarkably, the number of presently recognized
Arctic and Antarctic turrids is nearly the same.
The majority of the known Antarctic turrids were
described in publications based on the large Antarctic
expeditions of the early 20th Century (Strebel, 1908,
Thiele, 1912, Hedley, 1916). Hermann Strebel (1908)
described numerous gastropods using material collected
by the Swedish Antarctic Expedition (1901-1903) under
the command of Otto Nordenskjold. Among these were
1 1 new species of turrids, of which ten were from within
the Antarctic Convergence (Strebel, 1908:87), the highest
number of species of Antarctic Conoidea to be described
in a single publication. All of the species described by
Page 44 THE NAUTILUS, Vol. 127, No. 2
Strebel were considered valid by most subsequent
authors (e.g., Powell, 1951, Engl, 2012). Five of Strebels
species are rather common in collections and are often
referred to in the literature. 01 these, four serve as type
species of subsequently described genera:
• Surcula magnified Strebel, 1908 ( =Aforia magnified,
family Cochlespiridae Powell, 1942)
• Bela antarctica Strebel, 1908 (= Conorbela antarctica,
type species of the genus Conorbela Powell, 1951,
family Pseudomelatomidae Morrison, 1965)
• Bela pelseneri Strebel, 1908 (= Propebela ( Lorabela )
pelseneri, type species of the subgenus Lorabela
Powell, 1951, family Mangeliidae P. Fischer, 1883)
• Bela purissima Strebel, 1908 (= Tijphlodaphne
purissima, type species of the genus Tijphlodaphne
Powell, 1951 family Borsoniidae A. Bellardi, 1875).
• Bela turrita Strebel, 1908 (— Belaturricula turrita
turrita, type species of the genus Belaturricida Powell,
1951, family Borsoniidae)
The remaining five Antarctic species described by
Strebel (1908) as turrids are less well known, and all but
Pleurotornella bathybia are reviewed here. Thanks to the
courtesy of Dr. Anders Waren we were able to study the
type material of Strebels species in the collections of
the Swedish Museum of Natural History, Stockholm,
including the radulae of three of the species.
The holotype of Pleurotornella bathybia had been lost
some time ago, and a neotvpe (ZSM Mol 2002 1313) has
only recently been designated (Engl, 2012: 185, pi. 76,
fig. 2a) from off South Georgia, near the locality where
Strebel s holotype was collected. As this taxon is presently
known only from its shell and protoconch morphology,
it is provisionally retained within Conoidea.
The radulae and shell characters of the four species
treated herein (all originally described in the turrid genus
Bela ) clearly demonstrate that these taxa belong in the
superfamily Buecinoidea. Badular morphology indicates
a close affinity to the genus Falsimohnia Powell, 1951. In
this paper we review the genus Falsimohnia and include
a redescription of four of Strebel s species.
MATERIALS AND METHODS
The type specimens of species described by Strebel
are housed in Swedish Museum of Natural History. We
also examined specimens from the Museum national
d’Histoire naturelle, and the Institute Royal des Sciences
Naturelles de Belgique for comparative purposes.
Radulae were cleaned with diluted bleach (1 part of
commercially available bleach to 3-4 parts of distilled
water), mounted on round glass cover slips and air-dried.
Cover slips were then mounted onto SEM stubs, coated
with gold and examined under a }EOL JSM 840A scan-
ning electron microscope. Abbreviations used in text:
AL: Aperture length; AW: Aperture width; IRSN: Insti-
tute Royal des Sciences Naturelles de Belgique, Brussels;
MNHN: Museum national d’Histoire naturelle, Paris;
NHMUK: The Natural History Museum, London; SL:
Shell length; SMNH: Swedish Museum of Natural His-
tory, Stockholm; ZMB: Zoological Museum of Berlin, Berlin;
ZSM, Zoologisehe Staatssammlung Miinchen, Munich.
SYSTEMATICS
Class Gastropoda Cuvier, 1797
Order Neogastropoda Wenz, 1938
Superfamily Buecinoidea Rafinesque, 1815
Family Buccinulidae Finlay, 1928
Subfamily Cominellinae Gray, 1857
Genus Falsimohnia Powell, 1951
Falsimohnia Powell, 1951: 137; Engl, 2012:145.
Type Species: Bueeinum albozonatum Watson, 1882:358,
by original designation.
Description: Shell small to medium-sized (to 12 mm),
broadly fusiform, with high, convex last whorl and short
to slightly attenuated canal. Protoconch paucispiral,
smooth. Spiral sculpture usually of well-defined cords,
covering entire shell surface. Axial sculpture of weakly
pronounced, low, narrow, broadly spaced, and slightly
sigmoidal axial ribs most prominent on spire whorls.
Operculum oval, with terminal or subcentral nucleus,
situated in the lower corner of operculum. Radula with
rachidian teeth nearly as long as wide, V-shaped, with
deep anterior notch and long, single, medial cusp ema-
nating from posterior edge. Lateral teeth with long basal
plate bearing two long, curved cusps near middle and
inner margin of tooth. Outer portion of basal plate
tapered, without cusps. Outer cusp broader, longer,
inner cusp shorter, generally more curved.
Remarks: Powell (1951: 137) based his monotypical
genus Falsimohnia entirely on radular characters as a
“derivative of Pareuthria in which the normal three cusps
of the central tooth have been reduced to a single
member”, noting also the similarity in the opereula of
the two genera. He designated Bueeinum albozonatum
Watson, 1882 from Kerguelen Island as the type species,
and went on to diagnose this genus by emphasizing
eonehological similarities with the genera Pareuthria
Strebel, 1905 and Glypteuthria Strebel, 1905, but explic-
itly differentiated Falsimohnia on the basis of its having a
radula similar to that of the Arctic genus Mohnia Friele,
1878, in which the rachidian tooth has a single cusp
and the lateral teeth are bicuspid. The description of
Falsimohnia did not provide any additional eonehological
or anatomical diagnosis.
Falsimohnia albozonata (Watson, 1882)
(Figs 1-7, 13-19)
Bueeinum albozonatum Watson, 1882: 358; Watson,
1886: 212, pi. 13, fig. 7.
Y. I. Kantor and M.G. Harasewych, 2013
Page 45
Figures 1-12. Species of Fabimohnia. 1-7. Falsimohnia albozonata (Watson, 1882). 1-3. Shell, and 4. operculum, MNHN
uncataloged, Kerguelen Islands, lies Nuageuses, R/V Marion Dufresne, cruise 04, st. CP17, 48°47,1' S, 68°49.3' E, 70 in, (SL =
9.2 mm). 5-7. IRSN, IG 26482, Kerguelen Islands, S of lie Suhin, 40 m, (SL = 8.2 mm). 8-10. Falsimohnia cf. albozonata 8-9.
Shell and 10. operculum, MNHN uncataloged, Crozet Islands, RA7 Marion Dufresne, cmise 03, st. CP17, 46°24' S, 51°59' E, 180 in,
(SL = 6.0 mm). 11-12. Laclwsis australis Martens and Thiele, 1904. ZMB 59955, Syntypes. 11. This syntype (ZMR 59955a) is
designated as the lectotype of Lachesis australis Martens and Thiele, 1904, herein. 12. This syntype (ZMB 59955b) becomes the
paralectotype of L. australis. The 5 mm scale bar applies to all shells, die 1 mm scale bar applies to opercula.
Page 46
THE NAUTILUS, Vol. 127, No. 2
Figures 13-16. Anatomy of Falsimohnia albozonata. 13. Right lateral and 14. left lateral views of the anterior foregut. 15. Right
lateral view or anterior foregut with right salivary gland removed to show the duct of the gland of Leiblein and valve of Leiblein.
16. Detail of distal end of penis, aoe, anterior oesophagus; dgl, duet of the gland of Leiblein; gL, gland of Leiblein; lsd, duct of
the left salivary gland; lsg, left salivary gland; nr. circumoesophageal nerve ring; poe, posterior oesophagus; pr, proboscis; prr,
proboscis retractors; rhy, rhynchodaeum; rsd, duct of the right salivary gland; rsg, right salivary gland; sp, seminal papilla; vL, valve
of Leiblein.
Lachesis? australis Martens and Thiele, 1904: 62, pi. 5,
fig. 18.
Pareuthria albozonata — Thiele, 1912: 244.
Falsimohnia albozonata — Powell, 1951: 138.
? Mangelia nigropunctata Martens, 1885: 91-92.
? Mangelia antarctica Martens and Pfeffer, 1886: pi. 1,
figs. 5 a, b.
Type Localities: Royal Sound, Kerguelen Islands,
49°28' S, 70° 13' E, 28 fms [51 m] (B. albozonatum );
Kerguelen Islands (L. australis)-. South Georgia (M.
nigropunctata, M. antarctica).
Material Examined:, IRSN IG 26482, Kerguelen
Islands, S ol lie Suhin, 40 m, 3 live, 1 dissected (Figures 1
E-G); MNHN, uncataloged, Kerguelen Islands, lies
Nuageuses, R/V Marion Dufresne, cruise 04, st. CP17,
48°47.T S, 68°49.3' E, 70 m (Figures 1 A-D); MNHN,
uncataloged, Crozet Islands, R/V Marion Dufresne,
cruise 30, st. DC60, 46°25' S, 50°22' E, 105-120 m
(11 dead); MNHN, uncataloged, Crozet Islands, R/V
Marion Dufresne, cruise 03, st. CP17, 46°24' S, 51°59' E,
180 in (1 live, radula examined, light form) (Figures 8-10).
Description: Shell medium-sized (to 10 mm), strong,
broadly fusiform, with evenly convex whorls and
short, broad siphonal canal. Protoconch rounded,
smooth, paucispiral. Axial sculpture limited to growth
lines, occasionally with narrow, broadly spaced, low,
slightly prosoeline ribs more pronounced on upper
teleoeonch whorls. Spiral sculpture of low rounded cords
(9-11 on penultimate whorl, 16-19 on last whorl) slightly
Y. I. Kantor and M.G. Harasewych, 2013
Page 47
Figures 17-20. Falsimohnia albozonata radulae. 17-18. Radula of specimen in figures 1-3, MNHN uncataloged, Kerguelen
Islands, lies Nuageuses, Marion Dufresney , cruise 04, st. CP17, 48°47,1' S, 68° 49. 3' E, 70 m. 19. Radula of specimen in figures 5-7,
IRSN, IG 26482, Kerguelen Islands, S of lie Suliin, 40 m. 20. Falsimohnia cf albozonata . Radula of specimen in figures 8-9, MNHN
uneataloged, Crozet Islands, R/V Marion Dufresne, cruise 03, st. CP17, 46°24' S, 51°59' E, 180 m. Scale bars = 10 pm.
narrower that the interstices. Aperture elongated (AIV
SL ~ 0.55-0.57), narrow (AL/AW ss 2.2), elliptical, with
narrow callus extending onto parietal and columellar
regions of inner lip. Operculum oval, with terminal
nucleus. Periostraeum thin, tightly adherent, transpar-
ent. Shell light hrown to rust-colored, with a characteris-
tic whitish spiral band along shell periphery, whitish
columella and siphonal canal.
Gross Anatomy: Last whorl of the animal of a single
specimen (IRSN IG 26482, SL 8.2 mm) was extracted
from the shell and dissected. Preserved tissues lack pig-
mentation. Foot short, truncated posteriorly, operculum
missing, likely lost during fixation (opercular disk dis-
tinct). Mantle cavity deep (LAV ~ 2), mantle edge did
not cover head. Siphon short, very broad. Head small,
with long, closely spaced tentacles, large, hrown eyes on
Page 48 THE NAUTILUS, Vol. 127, No. 2
very small lobes at tentacle bases. Disposition of mantle
cavity organs typical for Buccinoidea. Ctenidium narrow,
spanning % of mantle cavity, ospbradium equal in width
to ctenidium, slightly shorter. Hypobranchial gland with-
out distinct folds. Rectum very short, broad. Proboscis
(Figure 14, pr) long, narrow, about 3.2 mm in length
(0.7 AL). Proboscis retractor muscles (Figures 13, 14, prr)
very thin, paired, symmetrical, attached to rhynchodaeum
(Figures 13-15, rhy) at mid-length. Salivary glands
(Figures 13, rsg, 14, 15, lsg) medium-sized, oval, fused
ventral to rhynchodaeum, covering nerve ring (Figure 15,
nr) and most of valve of Leiblein (Figure 15, vL). Salivary
ducts (Figures 13, rsd, 14,15, lsd) very thick, running
along both sides of anterior oesophagus. Valve of Leiblein
well defined, conical. Gland of Leiblein (Figures 13, 14,
gL) small, simple, tubular, colorless, joins esophagus
through a slightly constricted duct posterior to nerve ring.
Odontophore spans 2/3 of proboscis length. Subradular
cartillages fused along entire length. Radula equal to
odontophore in length. Radula (Figures 17-19) narrow,
about 75-90 pm wide (1.6-1. 8% of AL). Lateral teeth
with long base tapered toward the outer edge, with two
strong, curved cusps, the inner cusp slightly shorter,
emanating from the inner half of the tooth. Rachidian
tooth with a V-shaped base nearly as long as wide, deeply
notched anteriorly and with one strong, long cusp ema-
nating from posterior midline of the tooth.
Remarks: We did not examine the holotype of
Buccinum albozonatum, but studied several specimens
collected close to the type locality (Kerguelen).
Conehologically they are very similar to the published
description (Watson, 1882) and subsequent illustra-
tions (Watson, 1886:pl. 13, fig. 7) of this species. We do
note, however, that all specimens we examined had a
slightly different operculum. Watson (1886) stated that
“Operculum is intermediate in form between that of
Buccinum and Cominella ; for, as in the latter, the nucleus
is at the lower end, but it is not apical, but is within the
edge towards the outer margin as in the former.” In the
specimens we studied, most opereula are eroded to vary-
ing degrees (Figure 4), yet their general shape indicates
that their nuclei are terminal, not subcentral. In other
respects the shells of the specimens examined are very
similar to the figure published by Watson. The opereula
of the samples we examined matched the illustration of
the operculum of F. albozonatum that was published by
Powell (1951: fig. N, 127).
Powell (1951:138) considered Falsimohnia albozonata
to have a broad geographic range, and synonymized
Mangelia antarctica Martens and Pfeffer, 1886 (from
South Georgia Island) and Lachesis? australis Martens
and Thiele, 1903 (from Kerguelen Island). The name
Mangelia antarctica was unnecessary replacement name
for Mangelia nigropunctata Martens, 1885 from South
Georgia (Zelaya, 2005).
We were not able to examine the type material of
Mangelia nigropunctata . However, Martens and Pfeffer’s
illustrations (1886:pl. 1, fig. 5) indicate that their taxon is
very similar to F. albozonata, and also has the whitish
spiral band along the shell periphery that is present in
F albozonata. As we have not found any material of
F. albozonata from South Georgia, we cannot confirm the
presence of tliis taxon off South Georgia, nor its status as
senior synonym of M. nigropunctata (= M. antarctica) .
Powell (1951: fig. K, 59) illustrated the radular denti-
tion of a specimen collected off South Georgia Island
that he attributed to F. albozonata . A reexamination of
voucher material (station 149) identified by Powell and
deposited in the Natural History Museum (NI4MUK
1961499) revealed the specimens to be referable to
F.fulvicans (see below for further discussion).
Another species that Powell (1951) considered to be a
synonym of F. albozonata is Lachesis australis, also from
Kerguelen. Two syntypes of L. australis were located
in the Zoological Museum of Berlin (ZMB 59955)
(Figures 11-12). One of the specimens (Figure 11)
matches typical Falsimohnia albozonata, while the
other (Figure 12) differs in having a more slender shell
with a more elongated aperture and uniform colora-
tion, lacking the whitish band along the shell periphery
and siphonal canal. We examined a similar specimen
from Crozet Islands (Figures 8-10) and found the radula
(Figure 20) to be identical to that of typical specimens of
F. albozonata (Figures 17-19). It is at present unclear
whether die conchological differences are due to intra-
specific variation, or whether the two morphotypes repre-
sent different species of Falsimohnia. In order to stabilize
the nomenclature, we here designate the specimen illus-
trated in Figure 1 1 (ZMB 59955a) as lectotype of Lachesis
australis Martens and Thiele, 1904. The remaining speci-
men (Figure 12, ZMB 59955b) becomes a paraleetotype.
As the lectotype has all the morphological features of
Falsimohnia albozonata, Lachesis australis Martens and
Thiele, 1904, is a junior subjective synonym of Buccinum
albozonatum Watson, 1882.
Falsimohnia fulvicans (Strebel, 1908)
(Figures 21-27, 35-36)
Bela fulvicans Strebel, 1908: 15, Taf. 2, fig. 25 a-d;
Melvill and Standen, 1912: 356; Powell, 1951: 56
(Burwood Bank), 59 (South Georgia); Zelaya, 2005:
128, fig. 52 (figure erroneously labelled as Lorabela
pelseneeri); Engl, 2012: 181, pi. 73, fig. 7a-c.
Type Locality: South Georgia, Cumberland Bay, out-
side Grytviken, 54°22' S, 36°27' W, 24-52 m (Swedish
South Polar Expedition, sta. 25). [The coordinates provided
on the printed label accompanying the specimen and in
Strebel’s original description are likely erroneous, as they
correspond to a location on dry land close to Grytviken.]
Type Material: Lectotype, SMNH Type 6303 (des-
ignated by Engl, 2012: p. 181) (Figures 21-22).
Paralectotypes: SMNH Type 1030 (st. 34, Cumberland
Bay, 54° IP S, 36°18' W, 252G310 m) (Figures 24-26);
Y. I. Kantor and M.G. Harasewyeh, 2013
Page 49
Figures 21-32. Species of Falsimohnia. 21-27. Falsimohnia ftilvicans (Strebel, 1908). 21-22. Leetotype, (SMNH Type-6303),
(SL 8.1 mm). 23. Paralectotype, (SMNH Type-6159). 24-25. Body of paralectotype (SMNII Type 1030) 26. penis tip of
paralectotype in figures 24-25. 27. Specimen from South Georgia identified by Powell as Falsimohnia albozonata, NHMUK
1961499, (SL 7.6 mm). Figures 28-32. Falsimohnia minor (Strebel, 1908). Syntypes (SMNH Type-1057). 28. (SL 5.6 mm). 29.
(SL 5.8 mm). 30-32. (SL 6.5 mm). Left scale bar applies to figures 21,22, 27, right scale bar applies to figures 28-32. The 1mm
scale liar applies to figure 26.
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THE NAUTILUS, Vol. 127, No. 2
Figures 33-38. Radulae. 33-34. Falsimohnia minor (Strebel, 1908), radula from syntype in figure 29. 35-36. F fulvicans
(Strebel, 1908), radula from paralectotype in figures 24-25. (SMNH Type 1030). 37-38. Strebelona notophila (Strebel, 1908),
radula from holotype. Figures 44-45 (SMNH Type-1064). Seale Bars = 10 pm.
Y. I. Kantor and M.G. Harasewyeh, 2013
Page 51
SMNH Type 6159 (st. 95, off Astrolabe Island, 64°09' S,
58° 17' W, 95 m) (Figure 23).
Description (based on intact shell of lectotype):
Shell medium-sized (to 8.1 mm), strong, thick, narrowly
fusiform, with moderately convex whorls and attenuated,
narrow, siphonal canal. Protoconch eroded, rounded,
paucispiral. Whorls with narrow and slightly concave
subsutural ramp, evenly convex along periphery. Axial
sculpture ol narrow, weakly sinuate and weakly
prosocline ribs, extending from suture to suture on spire,
becoming obsolete on periphery of last whorl (~ 15 ribs
on penultimate and last whorls). Spiral sculpture of sharp,
raised spiral cords. Cords (7-9 on spire whorls, ~ 18 on
last whorl) broadly spaced adapically, with interven-
ing spaces 1-2 times broader than cords. Cords become
broader than intervening spaces along shell periph-
ery, indistinct on siphonal canal. Aperture large (AL/SL
« 0.53), elliptical (AL/AW « 2.0), with callus extending
along parietal region and columella. Operculum oval, with
terminal nucleus, shifted to left. Periostracum beige,
tightly adhering. Shell beige, lighter on siphonal canal.
Gross Anatomy: Animal of the paralectotype with a
broken shell (Figures 24-26, <$) was dissected. Anatomy
similar to Falsimohnia albozonata in most regards. Eyes
large. Penis long, about the length of mantle cavity, dor-
soventrally compressed, blunt distally with long but nar-
row seminal papilla shifted posteriorly, situated on dorsal
edge of penis (Figure 26). Proboscis of moderate length,
cylindrical, tapering distally. Radula (Figures 35-36) nar-
row (~60 pm wide), lateral teeth with broad base, elon-
gated and weakly recurved outer edge, two strong cusps,
inner cusp shorter, emanating from middle and inner
portions of base. Central tooth with nearly square basal
plate, its anterior edge V-shaped with indentation accom-
modating single medial cusp emerging from rounder pos-
terior edge of anteriorly adjacent tooth. Lateral edges of
basal plate of central tooth with thickened ridges parallel
to central cusp, possibly vestiges of lateral cusps. Stomach
with long posterior mixing area.
Remarks: The species is rather distinct, resembling the
shell outline of Probuccinum costatum (Thiele, 1912), but
differing from that taxon in having more numerous axial
ribs on the shell, and a different radula. In Probuccinum
the radula is typical buccinulid, with broad tricuspid
central teeth and tricuspid lateral teeth that lack an elon-
gated outer basal plate. Falsimohnia fulvicans appears to
be rare, as the only confirmed record since the origi-
nal description is that of Zelaya (2005). The radula of
F. fulvicans is very similar to that of F. albozonata as is
its penis morphology, both providing support for the
transfer of this species to the genus Falsimohnia.
In his discussion of F. albozonata , Powell (1951:138)
considered the range of this species to include South
Georgia Island. Two specimens from Discovery station
149 that had been identified by Powell as F. albozonata
(NHMUK 1961499) were examined (Figure 27). The
similarity of these specimens to the lectotype of
F fulvicans indicates that they were misidentified as
F albozonata by Powell. Discovery station 149 is very
close to Cumberland Bay, the type locality of F. fulvicans ,
and the depth is comparable to that from which a
paralectotype of F fulvicans was collected. In a survey of
the Mollusca of the Burwood Bank, Melvill and Standen
(1912:356) reported a single “imperfect, bleached speci-
men” that they tentatively identified as Bela fulvicans.
Distribution: South Georgia, and questionably
Burwood Bank.
Falsimohnia minor Strebeh 1908
(Figures 28-34)
Bela anderssoni var. minor ? Strebel, 1908: 15; Powell,
1951:59; Engl, 2012:181, pi. 73, fig. 6a.
Typhlodaphne innocentia Dell, 1990: Aldea and Troneoso,
2008: 82, fig. 64; Aldea and Troneoso, 2010: 144-145,
fig. 157.
Bela poly sarca var. minor Locard , 1897, Bela pyrrhogramma
var. minor Locard, 1897, and Bela anderssoni var.
minor? Strebel, 1908, were all proposed as taxa of sub-
specific rank (ICZN, 1999: Article 45.6.4). In our view,
both Bela polysarca var. minor Locard , 1897 and Bela
pyrrhogramma var. minor Locard, 1897 are nomina nuda ,
as neither name is accompanied by a description or defi-
nition of the taxon, nor by an indication (ICZN, 1999:
Article 12). As nomina nuda , these names are not avail-
able names and do not enter into homonomy (ICZN,
1999: Article 54).
Type Locality: South Georgia Island, outer Cumberland
Bay, 54° 11' S, 36°18' W, 252-310 m, Swedischen
Siidpolar-Expedition, sta. 34.
Type Material: 3 syntypes, SMNH Type-1057.
Description: Shell small (to 6.5 mm), strong, moder-
ately thick, with convex whorls and short broad canal,
with shallow but distinct notch. Protoconch bulbous,
eroded on all specimens. Profile of whorls between
suture and shoulder slightly concave. Axial sculpture of
narrow, slightly sinuous, nearly orthoeline ribs (14-15 on
penultimate and last whorls), extending from suture to
suture on spire and suture to siphonal canal on last
whorl. Spiral sculpture of low, sometimes indistinct,
closely spaced spiral cords that are broader than inter-
vening spaces. Aperture large (AL/SL = 0.54-0.61),
moderately broad (AL/AW« 1.9), elliptical, inner lip
with narrow callus on columellar and parietal portions.
Operculum oval, transluscent, with terminal nucleus,
shifted to left. Periostracum tightly adhering, beige to
yellowish. Shell orange to brownish beige, siphonal canal
and columella lighter, often whitish. Radula (Figures 33-
34) [narrow (~ 55 pm wide, 1 .7% of AL),] short. Lateral
teeth with long basal plate with elongated outer edge and
two strong, curved cusps, inner one slightly shorter.
Basal plate of central tooth nearly square, with shallow
Page 52
notcli in anterior margin to accommodate cusp or adja-
cent tooth, and one strong, long medial cusp emanating
from posterior edge. Small denticle on left side of cusp.
Remarks: Strebel (1908) attributed only varietal status
to this taxon. We regard Falsimophnia minor to be a
distinct species, differing from F. andersssoni in having
more pronounced, wider and more broadly spaced axial
ribs, and far less distinct spiral sculpture. The morphol-
ogy of the radula, which is nearly identical to that of
F. albozonata and F. fulvicans , confirms the placement
of this species in the bueeinulid genus Falsimohnia.
Conehologieally, F. minor resembles some speci-
mens of Chlanidotella modesta (Martens, 1885) from
South Georgia, but differs in its smaller shell size and
radular morphology. In C. modesta the lateral teeth have
4-5 cusps and the central tooth bears three distinct
cusps of similar size (Powell, 1951: fig. L80; unpub-
lished observations).
THE NAUTILUS, Vol. 127, No. 2
Distribution: South Georgia, Bellinghausen Sea, Peter I
Island, 90-310 m.
Falsimohnia anderssoni (Strebel, 1908)
(Figures 39-41)
Bela anderssoni Strebel, 1908: 14—15, Taf. 2, fig. 24 a-d;
Mellvill and Standen, 1912:355; Powell, 1951: 56;
Engl, 2012:180-181, pi. 73, fig. 6b.
Type Locality: SE of Seymour Island, 64°20'S,
56°38'W, 150 m, Swedischen Siidpolar- Expedition, sta. 5.
Holotype: SMNH Type-1063.
Description: Shell small (to 9.4 mm), fusiform, thick,
with evenly rounded, convex whorls and short broad
canal. Protoconch bulbous, eroded. Axial sculpture of
low, very narrow, slightly sinuous ribs (19 on penultimate
Figures 39-45. Species of Falsimohnia and Strebela new genus. 39—43. Falsimohnia anderssoni (Strebel, 1908). 39—41. Holotype
(SMNH Type-1063) (SL 9.4 mm). 42-43. USNM 870245, off Elephant Island, South Shetland Islands, 220-240 m. 44-45. Type
species of Strebela new genus: Bela notophila Strebel, 1908, holotype, (SMNH Type-1064), (SL 8.1 mm), (radula extracted).
Y. I. Kantor and M.G. Harasewych, 2013
Page 53
whorl) most pronounced on spire whorls, and raised
growth striae that are prevalent on final whorl. Spiral
sculpture of low, rounded, closely spaced spiral cords
(14 on penultimate whorl, ~30 on final whorl) that are
wider than intervening spaces. Aperture large (AI7SL =
0.56), broadly elliptical, (AL/AW ~ 1.9), inner lip with
narrow callus on parietal and columellar portions. Oper-
culum oval, transparent, with terminal nucleus shifted to
left, about half of aperture length. Periostracum beige,
peeling. Shell beige, lighter than periostracum. The rad-
ula of the holotype was not examined.
Remarks: The shell shape and sculpture of Falsimohnia
anderssoni are similar to those of F. albozonata and
“ Pareuthria ” innocens (Smith, 1907) with which it may
be confused [see discussion for comments regarding the
generic reallocation of Pareuthria innocens], Falsimohnia
anderssoni differs from typical F. albozonata in its uniform
beige shell color. It differs from “ Pareuthria ” innocens
[syntype, BMNH, illustrated by Dell (1991), fig. 301 as
“holotype”] in having a broader shell with more convex
whorls, a wider aperture, more prominent spiral cords
and low narrow axial ribs. We take a conservative
approach in recognizing F. anderssoni as a distinct species,
but more detailed studies as additional material becomes
available may reveal that it represents a variation of either
F. albozonata or “P innocens" . Some authors (eg.
Castellanos and Landoni, 1993) have considered
Falsimohnia anderssoni to be a junior synonym of r’Bela
michaelseni Strebel, 1905 from off Patagonia, although
B. michaelseni has a more slender shell with more pro-
nounced axial ribs. However, there is little overlap
between the gastropod fauna occuring within Antarctic
Convergence and that of the Magellanic Province, even at
the generic level (e.g., Harasewych et ah, 2000; I larasewyeh
and Kantor, 2004). Species originally believed to have a
broad range spanning both Magellanic and Antarctic
regions, generally on the basis of convergent shell morphol-
ogy, are often shown to represent taxa referable to differ-
ent genera and/or families when examined anatomically.
The geographic range of this species is extented to
include the South Shetland Islands based on a speci-
men (USNM 870245) in the Smithsonian collections
(Figures 42-43).
Distribution: SE of Seymour Island and Elephant
Island, South Shetland Islands, 150-240 m.
DISCUSSION OF THE GENUS FALSIMOHNIA
We had earlier speculated (Harasewych and Kantor,
2004:42) that, based on opercular and radular morphology,
Antarctodomus okutanii Numanami, 1996 may be refer-
able to Falsimohnia , a view that was adopted by Engl
(2012:145). Differences in radular morphology between
Antarctodomus okutanii and Falsimohnia albozonata,
the type species of Falsimohnia, most notably the lack of
a long, tapered outer edge of the lateral teeth of A. okutanii
as well as the broader placement of the cusps on the lateral
teeth in this species, indicate to us that more research is
needed for an accurate generic placement for this taxon.
Among the species described by Strebel (1908) as
Bela, B. notophila Strebel, 1908 has been attributed to
Lorabela (Powell, 1951; Zelaya, 2005) or conservatively
retained in Bela (Engl, 1912). As with the species
discussed above, the assignment of Bela notophila to the
Conoidea lias never been questioned. However, an
examination of the radula obtained from the holotype of
this species (Figures 37-38) undoubtedly places it within
Buecinoidea. This taxon differs from all other Antarctic
Buccinoidea in having a shell with axial sculpture of
sinuous broadly spaced prosocline ribs. It is superficially
similar to species of Lorabela, to which it was attributed
by Powell (1951) when he established the genus Lorabela.
The unique combination of radular and shell characters
in this species necessitates the description of a new genus
to include this species.
Genus Strebela new genus
Type Species: Bela notophila Strebel, 1908.
Description: Shell small, (to 8 mm), strong, thick,
fusiform, with convex whorls slightly angulated at shoul-
der, attenuated broad siphonal canal. Protoconch
paucispiral, smooth. Axial sculpture of wide, broadly
spaced, strongly prosocline ribs (8 on last whorl). Spiral
sculpture of few low sharp cords. Aperture moderately
broad, elliptical (~ Vi shell length). Operculum small,
oval, with terminal nucleus. Radula with long bicuspid
lateral teeth and unicuspid central tooth with nearly
square basal plate.
Remarks: This new genus differs from Falsimohnia in
having a shell with a distinct, angulated shoulder, pro-
nounced, widely spaced and strongly prosocline axial ribs,
stronger and fewer spiral cords, and a well demarcated
siphonal canal. The lateral teeth of the radula of Strebela
have a thin basal plate, and two thin cusps that are broadly
spaced, emanating from the inner and outer ends of the
basal plate. Lateral teeth of Falsimohnia are or compara-
ble length but much thicker and stouter. The two cusps
are broader, and emanate from near the center and inner
ends of the basal plate, which has a tapering outer edge.
Etymology: This new genus honors Hermann Strebel
in recognition of his pioneering studies of Antarctic and
subAntarctic Mollusca collected by the Swedish Antarc-
tic Expedition (1901-1903). It is a combination of the
name Strebel with the genus name Bela, to which he
originally assigned the type species.
Strebela notophila (Strebel, 1908)
(Figures 37-38, 44-45)
Bela notophila Strebel, 1908: 20-21, Taf. 2, fig. 28 a-d;
Engl, 2012:181, pi. 73, fig. 5.
Page 54 THE NAUTILUS, Vol. 127, No. 2
Lorabela notophila — Powell 1951: 171, fig. N, 144;
Zelaya, 2005: 128.
Type Locality: South Georgia, outer Cumberland Bay,
54° ll'S, 36°18'W, 252-310 m, Swedischen Sudpolar-
Expedition, sta. 34.
Holotype: SMNH Type-1064
Description: Shell medium-sized (to 8.1 mm), strong,
tliiek, fusiform, with convex whorls, angular shoulder, and
attenuated siphonal canal slightly recurved to left, demar-
cated by a shallow notch. Protoconch broken in holotype.
Whorl profile below suture slightly concave on penulti-
mate and last whorls. Axial sculpture of pronounced,
wide, broadly spaced, sinuous, strongly prosoeline ribs
(8 on last whorl) that extend from suture to suture on spire,
becoming obsolete on last whorl at transition to siphonal
canal. Spiral sculpture of low, narrow, widely spaced, well-
defined cords (4 on penultimate whorl, 7 on the last
whorl), absent on subsutural rim, distinct on the shell
periphery, nearly obsolete on the siphonal canal. Aperture
elongated (AL/SL ~ 0.47), narrow (AW/AL ~ 0.40), ellip-
tical, with narrow callus along parietal and columellar
regions. Siphonal canal well defined, anal sinus not pro-
nounced. Operculum small (~ 0.5 AL), thick, ovate, with
terminal nucleus shifted to left. Shell beige in color,
siphonal canal whitish. Radula (Figures 37-38) narrow
(~ 80 pm wide, 2.1% of AL). Lateral teeth with long, thin
basal plate, two narrow, broadly spaced curved cusps (one
at each end of tire basal plate), inner one slightly shorter.
Basal plate of central tooth nearly square, weakly notched
anteriorly, with one thin, long, medial cusp emanating
from posterior edge.
Distribution: South Georgia, 97-310 m.
Remarks: Powell (1951: fig. 114) illustrated the
protoconch of this species, showing it to be smooth
and paucispiral.
DISCUSSION
Within the Neogastropoda, a number of taxa unrelated to
Conoidea were originally described in genera belonging
to one of the “turrid” families on the basis of convergent
shell morphology, among them species later found to
belong to the genera Exilia (Ptyehatractidae), Antimitra
(Bueeinidae), and Daphnellopsis (Muricidae). Conversely,
some taxa originally described as buccinoideans have
been shown to be referable to Conoidea when their anat-
omy was examined (e.g., Kantor and Harasewyeh, 1999).
Conehologieally the species transferred here to
Falsimohnia are similar to several Antarctic species that
have been attributed to Pareuthria Strebel, 1905. They
share a similar elongate-oval shell with a large, smooth
protoconch and weak axial sculpture.
The genus Pareuthria Strebel, 1905, contains 20 spe-
cies [WoRMS (Bouchet and Rosenberg, 2012)], of which
only five [P. innocens (Smith, 1907), P. plicatula Thiele,
1912, P. valdiviae (Thiele, 1925), P. turriformis Egorova,
1982; P. hoshiaii Numanami, 1996], have been recorded
within the Antarctic Convergence. The majority of the
species of Paraeuthria , including the type species ( Fusus
plumbeus Philippi, 1844, by subsequent designation,
Tomlin, 1932) are limited to the Magellanic Province.
Radulae of the type species Pareuthria plumbeus
(G. Pastorino, personal communication), and of two other
Magellanic species, P. fuscana (Bruguiere, 1789) and
P. venustula Powell, 1951 (Powell, 1951: figs. 69, 70), have
a broad central tooth with three cusps that are similar in
size, and lateral teeth with two cusps (outer cusp longer)
that lack a lateral extension along the outer margin of
the basal plate.
Pareuthria innocens (Smith, 1907), a species with its
type locality in the Ross Sea (Hut Point, McMurdo
Sound, in 45-54 m) and with a circum-Antarctie distri-
bution at depths of 6 to 549 m, was originally described
in the conoidean genus Thesbia Jeffreys, 1867. It was
subsequently transferred to the genus Pareuthria on the
basis of its buccinoidean radula by Thiele (1912; 207),
who provided an illustration (Thiele, 1912: pi. 16, fig. 22)
that showed bicuspid lateral teeth and a tricusip central
tooth. Egorova (1982: fig. 50) showed this species to have
a central tooth with a single, median cusp. Hain (1990:
55) noted the differences in these figures, and provided
an SEM image of the radula from a specimen he attrib-
uted to P. cf. innocens (Hain, 1990: pi. 22, fig. 7) in which
the central tooth has a single, median cusp, but lateral
teeth have two thin cusps situated at the ends of a basal
plate that lacked a lateral extension. Numanami (1996:
fig. 121C) published a SEM of the radula that clearly
depicts a central tooth that is narrow, with a single,
median cusp that is flanked by small denticles. The lat-
eral teeth have two thick cusps as well as outer lateral
projections. These reports indicate that either there is
great morphological variability in the radula of this taxon,
or that specimens representing more than one species
contributed to these observations.
Pareuthria plicata Thiele, 1912, was described from
the Davis Sea (Gauss station, in 385 m) and differenti-
ated from P. innocens on the basis of shell sculpture.
Engl (2012: 149) noted that this species was based on
24 variable syntypes, and designated a lectotype (Engl,
2012: pi. 54, fig. 3a). Hain (1990: pi. 22, fig. 8) published
a SEM of the radula of a specimen identified as P. cf.
plicata that was very similar to his image of the radula of
P. cf. innocens. The squarish basal plate and long, medial
cusp of the central tooth were more prominent, while
the lateral teeth also had two, widely separated cusps
that lacked outer lateral projections.
Pareuthria hoshiaii Numanami, 1996, is presently
known only from its holotype, collected at Gunnerus
Bank near Syowa Station (Queen Maud Land) at 288 m.
Its radula (Numanami, 1996: fig. 125D) has a central
cusp with a single, broadly triangular cusp flanked
by one or more pronounced denticles on each side.
Lateral teeth have two thick cusps and an outer lat-
eral projection.
Y. I. Kantor and M.G. Harasewych, 2013
Page 55
Two additional species attributed to Pareuthria. Euthria
( Pareutliria ) valdiviae Thiele, 1925, from unspecified
depths (Valdivia Station 160) off Kerguelen Island, and
Pareuthria turrifomiis Egorova, 1982, from off the
Shackleton Ice Shelf, Davis Sea at depths of 460 m, were
described solely on the basis of shell morphology, and
their radulae remain unknown.
Of the five species attributed to the genus Pareuthria
that occur within the Antarctic Convergence for which
radular morphology is known, it is clear that the radulae
of Pareuthria innocens and P. hoshiaii differ from that of
P. plumbeus, the type species of the genus. They more
closely resemble the radulae of species of Falsimohnia ,
including F alhozonata , the type species, most notably in
having a central tooth with a squarish basal plate and
single prominent medial cusp (that may be flanked by
denticles) as well as lateral teeth with a long basal plate
with a tapering outer edge, and two thick cusps along
the inner 2/3 of the tooth. We therefore transfer the
taxa Pareuthria innocens (Smith, 1907) and P. hoshiaii
Numanami, 1996 to the genus Falsimohnia. The result-
ing binomina are Falsinwhnia innocens (Smith, 1907)
new combination and Falsimohnia hoshiaii (Numanami,
1996) new combination.
The new genus Strehela is characterized by radula
similar to that of Falsimohnia , but differs in having a
central tooth with a squarish basal plate that appears
thinner, with less well defined lateral edges, a more shal-
lowly concave anterior that lacks an indentation for the
cusp of the anteriorly adjacent tooth, and a thinner
median cusp emerging from the posterior tooth margin.
The lateral teeth have a narrower basal plate with two
much thinner and longer cusps that are broadly spaced.
The outer cusp emerges from the outer margin of the
lateral tooth, which lacks the lateral projection.
A radula with this morphology was reported for
Pareuthria plicatula [Hain, 1990: pi. 22, fig. 8 (as
Pareuthria cf. plicatula ); Numanami, 1996: fig. 123D],
as well as for a specimen identified as Pareuthria cf.
innocens (Hain, 1990: pi. 22, fig. 7). However, there are
conspicuous differences in shell morphology between
Strebela notophila (strongly shouldered shell with
8 prominent, sinuous and strongly prosocline axial
ribs and pronounced, widely spaced spiral cords) and
Pareuthria plicatula (shell with rounded shoulder, ~ 40 thin,
weakly sinuous, nearly orthocline axial ribs and lack of
pronounced spiral sculpture). The appropriate generic
allocation of “ Pareuthria ’’ plicatula is not yet clear, but
major differences in radular morphology exclude it from
the genus Pareuthria.
The remaining two species from within the Antarctic
convergence that have been attributed to Pareuthria are
known only from their shell morphology. Pareuthria
valdiviae (Thiele, 1925), from Kerguelen Island was col-
lected with Falsimohnia albozonata, which it resembles
in shell morphology, but has more pronounced axial
striae and lacks the whitish band across the periphery.
Pareuthria turriformis Egorova, 1982 from the Ross Sea,
was attributed to the genus with doubts, as it differs in
shell morphology from both the Magellanic species of
Pareuthria and from Falsimohnia .
A review of the buccinoidean genera of the Antarctic
and Magellanic regions (Harasewych and Kantor, 2004:
Appendix 1) revealed only three genera that are repre-
sented by species in both regions: Meteuthria Thiele,
1912, Pareuthria , and Falsitromina Dell, 1990. Results
of the present study dispute the occurrence of the genus
Pareuthria within the Antarctic Convergence, and sug-
gest that both Pareuthria’ valdiviae and P. ' turriformis
will be referred to other genera when anatomical mate-
rial becomes available. A more detailed study of
Meteuthria and Falsitromina may reveal similar biogeo-
graphic patterns.
ACKNOWLEDGMENTS
We thank Dr. Anders Waren and Dr. Philippe Bouchet
for access to collections in their care. We are particularly
grateful to Dr. John Tucker for his thorough and
thoughtful review of this paper.
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THE NAUTILUS 127(2):57-64, 2013
Page 57
Aidemofusus ignotus new genus and new species, a new buccinid
(Gastropoda: Neogastropoda) from the North Pacific Ocean
Alisa Kosyan
Yuri I. Kantor
A.N. Severtzov Institute of Ecology
and Evolution
Russian Academy of Sciences
Leninsk-yj Prospect, 33
Moscow 119071, RUSSIA
[email protected]
ABSTRACT
Aidemofusus ignotus, a new species and genus of North Pacific
buccinids is described. Shell and operculum morphology as
well as general foregut anatomy are similar to several Colinae
(Buccinidae) genera, while radula structure is very unusual for
the family, precluding inclusion of the new genus into any
recognized subfamily of Buccinidae.
Additional Keywords: Buccinidae, taxonomy, Russia, Okhotsk Sea
INTRODUCTION
Buccinoidea is a large and diverse superfamily of pred-
atory marine gastropods, widely distributed in polar,
temperate and tropical waters of the World Ocean. It
encompasses several families, few of which are well
defined conchologically and anatomically (Nassariidae,
Melongenidae, Colubrariidae, Columbellidae) and well
supported in molecular phylogenies (Oliverio and Modica,
2010). Two families, Fasciolariidae and Buccinidae were
not retrieved as monophyletic in a few molecular analyses
and may be paraphyletic. Buccinidae (in currently accepted
sense - Bouchet and Roeroi, 2005) are especially hetero-
geneous, and include groups that are conchologically and
anatomically very different, which is reflected in their
complicated taxonomic structure. The state of the art clas-
sification presented in Bouchet and Roeroi (2005) recog-
nizes 6 subfamilies and 1 1 tribes. Despite tire high number
of described and accepted suprageneric taxa, there
remained several genera of Buccinoidea with uncertain
taxonomic position, among them Buccipagoda Ponder,
2010, Costaria Golikov, 1977, and Troschelia Morch, 1876.
Systematic^ of the Buccinidae is still based mostly on
eonehologieal characters with occasional use of radulae.
Relatively few buccinids have been included in molecular
phylogenetic studies (e.g., Hayashi, 2005; Kosyan et ah,
2009). The main reason is unavailability of properly pre-
served material, especially of boreal and Arctic species,
vast collections of which had been accumulated over the
years in European and American museums. Therefore mor-
phology and conchology remain the principal approaches
to buccinid alpha-taxonomy at the moment.
The practical needs for biodiversity studies require
descriptions of new taxa, even in groups for which a robust
phylogeny and classification are future goals. Many of
species remain rare and probably will not be recollected
in the near future.
While preparing a revision of the north-western Pacific
buccinids, we found a peculiar species that cannot be
confidently placed in any recognized suprageneric taxon.
The combination of eonehologieal, anatomical and radu-
lar characters preclude including it in any known genus.
Therefore a new species and genus are proposed, and
tentatively attributed to Buccinidae.
MATERIALS AND METHODS
The material for the study is deposited at P.P. Shirshov
Institute of Oceanology, Moscow, and Zoological Insti-
tute of Russian Academy of Sciences, St. Petersburg.
Standard methods were used, including manual dissec-
tion and scanning electron microscopy for examination
of the radulae. Abbreviations used in the text are: AL:
aperture length; D: shell diameter; H: height of the
shell; h: height of the last whorl; spm: specimen; IO: P.P.
Shirshov Institute of Oceanology, Russian Academy of
Sciences, Moscow, Russia; ZIN: Zoological Institute, Russian
Academy of Sciences; ZMMU: Zoological Museum of
Moscow State University, Russia.
SYSTEM ATICS
Order Neogastropoda Wenz, 1938
Family Buccinidae Rafinesque, 1815
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THE NAUTILUS, Vol. 127, No. 2
Genus Aidemofusus new genus
Type Species: Aidemofusus ignotus new species, by
original designation.
Description: Shell small, up to 25 mm in length,
broadly fusiform, with low spire and moderately long,
strongly recurved siphonal canal. Spiral sculpture of
low, closely spaced, wavy cords, with much narrower
interstices; about 12 cords on penultimate whorl and
30 on last whorl and canal. Axial sculpture of weak, low,
slightly recurved ribs, about 15 on last whorl. Aperture
high > V2 SL, elongate oval, strongly constricted on
passing to siphonal canal. Columellar margin weakly
concave, nearly straight. Columellar and parietal mar-
gins with narrow callus. Operculum with terminal nucleus,
strongly turned to left, about 0.8 AT. Central tooth of
radula small, with subsquare base and serrated posterior
margin. Lateral teeth square, with nearly straight posterior
edge with numerous (7-12) short sharp cusps. Salivary
ducts form widened pouches at base of inverted proboscis.
Remarks: Aidemofusus can be easily distinguished
by its unusual radula. It has a distant similarity to radulae
of Parancistrolepis Azuma, 1965 in the morphology of its
lateral teeth, but the central tooth is different as is the
operculum. Costaria borealis Golikov, 1977 and Troschelia
bemiciensis (King, 1846), possess wide lateral radular
teeth with multiple cusps, but the cusps are much longer
and the central tooth has a different shape in these genera.
Shell of Aidemofusus may be easily confused with shells of
small-sized buccinid genera possessing axial ribs such as
Retifusus Dali, 1916 and Retimohnia McLean, 1995.
Etymology: The genus name (masculine gender) derives
from aidemos (Greek: bashful, modest), recognizing the
unremarkable shell, lacking any diagnostic characters.
Aidemofusus ignotus new species
(Figures 1-24, Table 1)
Description: Shell small, thick, solid, broadlv fusi-
form, with low spire, of 5+ convex whorls (Figures 1-6).
Protoconch and upper telecoconch whorls abraded in
all specimens. Shell surface usually strongly eroded. Last
whorl high to very high, strongly convex. Spiral sculpture
of low, closely spaced wavy cords, with interstices four
times narrower than cords; about 12 cords on penulti-
mate whorl and 30 on last whorl and canal. Axial sculp-
ture of weak, low, slightly recurved ribs, about 15 on last
whorl. Aperture high, > V2 SL, elongate oval, strongly
constricted on passing to siphonal canal. Outer lip usu-
ally evenly convex, sometimes flattened in middle part.
Siphonal canal medium-long, broad, strongly recurved to
left. Columellar margin weakly concave, nearly straight.
Columellar and parietal margins with narrow callus.
Shell pale in color, covered with thin yellowish-beige
periostracum. Operculum (Figure 14) with terminal
nucleus, strongly turned to left (almost subspiral), com-
prising about 0.8 AL.
External Anatomy: Head wide, with moderately-long
contracted tentacles and large black eyes on lobes at base
of tentacles (Figures 8, 13). Foot contracted, with medium-
wide propodium (Figure 8). Mantle length equals width,
ctenidium spans 2/3 (paratype 1, Figure 9) or nearly entire
mantle length, more broad in posterior part (spm. 1,
Figure 15). Osphradium about % (paratype 1) or V2 of
ctenidium length (spm. 1). Penis of spm. 1 with large
conical seminal papilla, not surrounded by fold of skin
(Figure 17, sp). Capsule gland of paratypes 1 (Figure 9,
eg) and 3 large, occupying about 1/3 width and almost
complete length of mantle cavity. Bursa copulatrix
represented by narrow moderately thick-walled tube,
lined inside with multiple longitudinal folds (Figure 9,
be). Female orifice, a very small pore on tip of bursa
copulatrix tube (Figure 9, fo).
Digestive System: Description based on holotype,
paratype 1 and spm. L Proboscis short, completely (in
paratype 1, Figures 8, 12) or partially everted (spm. 1,
Figure 16). Proboscis retractors attached to base of pro-
boscis (Figures 12, 16, prr). Salivary glands paired, long,
equal to proboscis length (Figures 12, 16, sg). Salivary
ducts very thick, widen to form salivary poaches either
inside proboscis (holotype and paratype 1, Figures 10,
11, spo) or at mid length of anterior oesophagus, and
entering proboscis along oesophagus (Figure 16, spo).
In holotype and paratype 1 salivary poaches become
narrow close to buccal cavity (Figure 10, 11); in spm. 1
salivary poaches wider than oesophagus, opening into
buccal cavity at proboscis tip without narrowing. Buccal
mass as long as proboscis in holotype (Figure 10, bm),
about 2/3 proboscis length in paratype 1 (Figure 11, bm)
and spm. 1. Radula sac slightly shorter than buccal mass.
Radula of holotype 2.5 mm long and about 80 pm wide.
Table 1. Aidemofusus ignotus new species. Dimensions and gender of specimens examined in this study. AL: aperture length;
D: shell diameter; H: height of the shell; h: height of the last whorl.
A. Kosyan and Y. I. Kantor, 2013
Page 59
Figures 1-6. Shells of Aidemofusus ignotus new species. 1. Holotype. 2. Par a type 1. 3. Paratype 2. 4. Paratype 3. 5. Para type 4.
6. Spm. 1, from the western Bering Sea, 1960 m.
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THE NAUTILUS, Vol. 127, No. 2
Figures 7-12. Anatomy of Aidemofusus ignotus new species. 7-9, 11-12. Paratype 1. 10. Holotype. 7. Soft parts, ventral view.
8. Cephalopodium, dorsal view. 9. Mande. 10. Opened proboscis, holotype. 11. Opened proboscis, paratype 1. 12. Foregut.
Abbreviations: aoe, anterior oesophagus; be, bursa eopulatrix; bm, buccal mass; eg, capsular gland; ct, ctenidium; eye, eye; fo,
female orifice; ft, foot; gl, gland of Leiblein; hd, head; mo, mouth opening; nirr, medial radular retractor muscle; odr, odontophoral
retractor muscles; op, operculum; os, osphradium; pr, proboscis; pip, propodium; prpg, propodia] groove; prr, proboscis retractors;
pw, proboscis wall; r, radula; rd, rhynchodaeum; sd, salivary duct; sg, salivary gland; spo, salivary poaches.
consisting of 108 rows of teeth (Figures 19-20). Central
tooth small, with subsquarish base and trapezoidal, ser-
rated posterior margin. Lateral teeth are almost of the
same height and twice the width of the central tooth,
square, with about 8 equal short sharp cusps in the right
and 1 1 in the left longitudinal tooth row, some bearing
additional smaller denticles. Number of cusps and shape
of denticles differs from row to row. Radula ol paratype 1
is 3 mm long and about 75 pm wide, consisting of 98 rows
of teeth (Figure 21); central tooth is the same as above
described, lateral teeth bear about 13 and 8 extremely
short cusps respectively. Radula of spm. 1 is 3 mm long
and about 75 pm wide, consisting of 112 rows of teeth
with laterals bearing about 7 rather long cusps in each
longitudinal row (Figures 22-24). Gland of Leiblein very
small in paratype 1 (Figure 12, gl) and not found in spm.
A. Kosyan and Y. I. Kantor, 2013
Page 61
Figures 1.3-18. Anatomy ol Aidemofusas ignotus from the western Bering Sea, 1960 m. 13. Head. 14. Operculum 15. Mantle.
16. Foregut. 17. Distal part of penis 18. Stomach, general view. Scale bar 1 mm. Abbreviations: aoe, anterior oesophagus; cep.t,
cephalic tentacles; cm, eolumellar muscle; ct, ctenidium; tig, digestive gland; eye, eye; gon, gonad; int, intestine; kd, kidney;
nr, nerve ring; os, osphradium; poe, posterior oesophagus; pr, proboscis; prr, proboscis retractors; rd, rhynehodaeum; re, rectum;
s, siphon; sd, salivary duct; sg, salivary gland; sp, seminal papilla; spo, salivary poaches; st, stomach; vl, valve of Leiblein.
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THE NAUTILUS, Vol. 127, No. 2
Figures 19-24. Radulae of Aidemofusus ignotus new species. 19, 20. Holotype. 21. Paratype 1. 22-24. Specimen from the western
Bering Sea, 1960 m. Scale bar 50 pm.
1 (due to poor fixation). Valve of Leiblen large, swollen,
situated immediately anterior to nerve ring at proboscis
base in paratype I and at a distance from nerve ring
in spm. 1 (Figure 16, vl). Nerve ring medium large.
Posterior oesophagus slightly wider than anterior. Stom-
ach (Figure 18, st) spans about 1/4 of whorl, parallel to
its longitudinal axis and bordering kidney (Figure 18,
kd). Posterior mixing area absent. Stomach walls lined
with multiple low transverse folds; posterior oesophagus
lined with tall longitudinal folds. More detailed study
was not possible because of poor fixation.
Remarks: Shells of type specimens are strongly
eroded (paratype 5 lacks any spiral sculpture and not
figured). Specimens from the Sea of Okhotsk (holotype,
paratypes 1 and 2) differ slightly from specimens from
the Bering Sea (paratypes 3-5 and spm.l) in having a
higher spire. In specimens from the Okhotsk Sea, the
shell periphery is in the middle part of last whorl, while
in specimens from the Bering Sea the shell periphery is
in the upper third of the last whorl and the shells are
more globose. We consider these distinctions as resulting
from intraspecific geographic variability. A specimen
A. Kosyan and Y. I. Kantor, 2013
Page 63
from the western Bering Sea (Figure 6), collected in
1960 m is similar conchologieally to types, but we did
not include it in the type series because of the very large
bathymetric differences of habitat, and attribute it to the
new species conditionally.
Type locality: 50°5F N 156°01' E, Okhotsk Sea,
depth 94 m.
Type Material: Holotype ZMMU LC 39504, Okhotsk
Sea, off Shumshu Island, northern Kurile Islands, R/V
Vityaz, sta. 2821, 50°51' N, 156°01' E, depth 94 m, 27
May 1954. Paratypes 1,2 ZMMU LC 39505, sta. 2817,
51°00' N, 156° 09' E, depth 114 m, 26 May 1954.
Paratypes 3-5: ZIN 61623, Northern Bering Sea, R/V
Dalnevostochnik sta. 26, 64° 45.5' N, 169°35.5' W, 47 m,
17 Aug. 1932, coll. Ivanov, Makarov.
Other Material Examined: IO, Western Bering Sea,
R/V Vittaz, sta. 1030, 60°24.7' N, 173°46.9' E, depth
1960 m, 16 Oct. 1951 (1 live spm).
Distribution: Northern Bering Sea, Northern Kurile
Islands, 47-1960 m.
Etymology: Species name originates from ignotus (Eat.
unknown, strange), to reflect its unusual radula morphology.
DISCUSSION
The systematics of the large family Bueeinidae is very
complicated and not fully resolved. There are several
genera that can be allocated to either Bueeinidae or
Easciolariidae (e.g., Costaria , Troschelia ), and the attempts
to resolve their phylogeny have been unsuccessful, which
resulted in a paraph yletic Bueeinidae (Kosyan et ah,
2009). The shell of Aidemofusus ignotus can be easily
confused with other small-sized buccinids possessing axial
ribs (e.g., Retifusus and Retirnohnia). A distinctive charac-
ter of foregut of the new species are the extremely wide
salivary ducts that form thick-walled pouches. Salivary
pouches were found in several small-sized buceinid species
in the subfamily Colinae: Para retifusus kosugei Kosyan
2006, Retifusus roseus (Dali, 1877), Coins latericeus
(Moller, 1842), and Retirnohnia sp. (Kosyan, 2006, 2007;
Kosyan and Kantor 2009), but these all have thin walls.
In specimens of R. roseus with an inverted proboscis,
salivary pouches are usually posterior to the proboscis
base. During proboscis evertion, the salivary pouches
move inside and become more elongated. Transverse
sections of salivary ducts of R roseus show the presence
of a rather thick outer layer of circular muscle fibers
(Kosyan, 2007); this indicates the possibility of peristaltic
contractions. The function of the pouches is not clear.
They are probably used for accumulation and quick
delivery of large amounts of saliva during feeding.
The radula structure in the new genus is very unusual
for Bueeinidae. In Parancistrolepis , the lateral teeth are
somewhat similar, having numerous small cusps (although
longer than in the new genus), but the central tooth bears
3 or 4 distinct cusps, and the breadth of base of central
tooth exceeds length (Habe, 1972). In Costaria borealis
from the Kurile Islands and Troschelia bemiciensis (North
Atlantic), lateral teeth have multiple cusps, but the cusps
are much longer, and the teeth are much broader
(Golikov, 1977; Bouchet and Waren, 1985). The central
teeth of these taxa have only one or two cusps that are
distinct and relatively much longer cusps than those in
Aidemofusus. Cuspless central teeth are also found in the
family Columbellidae (e.g., Bandel, 1984).
The above discussion indicates that the taxonomic
position of Aidemofusus ignotus remains uncertain.
Conchologieally it can be attributed to the subfamily
Colinae, but its radula precludes its inclusion in any
recognized suprageneric taxon.
ACKNOWLEDGMENTS
The authors are very grateful to Dr. A.V. Gebruck,
Dr. E.M. Krylova, and T.A. Savilova from IO RAN
(Moscow, Russia) and Dr. B.I. Sirenko and R.A. Pikalova
from ZIN RAN (Saint-Petersburg, Russia) for access to
die collections in their care and for assistance while
working with molluscan collections and archives. Many
thanks to N.N. Surovenkova from IPEE RAN (Moscow,
Russia) who much helped with scanning electron
microscopy of die radulae, and to Dr. M.G. Harasewyeh
for valuable corrections of the manuscript. The study
was supported by RFBR grants No. 1 1-04-0 1284-a and
12-05-00082-a.
LITERATURE CITED
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(Kuroda) and A. troclioideus ovoideus Habe et Ito. Venus
24: 127-129.
Bandel, K. 1984. The radulae ol Caribbean and other
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Bouchet, P. and J.-P. Rocroi. 2005. Classification and noinen-
clator of gastropod families. Malaeologia 47: 1-397.
Bouchet, P. and A. Waren. 1985. Revision of the Northeast
Atlantic bathyal and abyssal Neogastropoda excluding
Turridae (Mollusca, Gastropoda). Bollettino Malacologico,
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Golikov, A.N. 1977. Investigation of prosobranchs of the family
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fauna 21(29): 102-104. [In Russian],
Habe, T. 1972. Notes on die genus Parancistrolepis Azuma
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Hayashi, S. 2005. The molecular phylogeny of the Bueeinidae
(Caenogastropoda: Neogastropoda) as inferred from the
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complete mitochondrial 16S rRNA gene sequences of
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forms found on the coast of Northumberland and of Durham.
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Kosuge, S. 1967. On the transfer of “ Phymorhijnchus ?”
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Kosyan, A.R. 2006. Two new species of the genus Pararetifusus
Kosuge, 1967 (Buccinidae: Colinae), with notes on die mor-
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pods of die subfamily Colinae (Neogastropoda: Buccinidae)
of die Far-East seas of Russia PhD dissertation. A N.
Severtzov Institute of Ecology and Evolution, Russian Ac.
Sci., Moscow, 223 pp. [In Russian].
Kosyan, A.R., M.V. Modica, and M. Oliverio. 2009. The anatomy
and relationships of Troschelia (Neogastropoda: Buccinidae):
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ship? The Nautilus 123: 95-105.
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83-94.
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THE NAUTILUS 127(2):65-77, 2013
Page 65
A new species of Striostrea (Bivalvia: Flemingostreidae) from the
upper Pliocene and lower Pleistocene strata of Florida, USA
Michael J. Bolton
Ohio EPA
4675 Homer Ohio Lane
Groveport, Ohio, 43125 USA
mike. bolton@epa. state, oh. us
Roger W. Port ell
Florida Museum of Natural History
P.O. Box 17800
University of Florida
Gainesville, FL 3261 1 USA
[email protected]
ABSTRACT
Faunal studies of the Tamiami and Caloosahatehee formations
(upper Pliocene to lower Pleistocene) in southern peninsular
Florida have revealed a new crassostreine oyster, Striostrea
paucichomata Bolton new species. Although similar in appear-
ance, this bivalve differs from Crassostrea virginica (Gmelin,
1791) by having weakly developed chomata (not visible in about
30% of the specimens examined), the right valve having a
riblet-bearing surfieial layer (visible only on exceptionally well-
preserved specimens), and an adductor muscle attachment
that is situated more dorsally. The geology of the type area of
S. paucichomata in northern Sarasota County, and a strati-
graphic nomenclatural history ol the Tamiami and Caloosa-
hatchee formations are reviewed. A key to Cenozoie crassostreine
oysters known from the southeastern United States is also
provided. The following new generic placements are proposed
for four previously named species: Mijrakeena sculpturata
(Conrad, 1840) new combination, Mijrakeena lawrencei
(Ward and Blackwelder, 1987) new combination, Mijrakeena
g reeni (Ward, 1992) new combination, Undulostrea locklini
(Gardner, 1945) new combination and Striostrea cahohasensis
(Pilsbryand Brown. 1917) new combination.
Additional Keywords: Mollusca, Crassostreinae, Striostrea
gigantissirna (Finch, 1824), Conradostrea , Fossil
INTRODUCTION
Investigations of two mounds of construction fill in
Manatee County, Florida (the fill probably originated
from northern Sarasota County quarries) in 1996 and
1998, which contained molluscan fauna typical of the
Pinecrest beds of the Tamiami Formation (upper
Pliocene to lower Pleistocene), revealed a crassostreine
oyster species different from Crassostrea virginica
(Gmelin, 1791) and herein described as Striostrea
paucichomata Bolton new species. A review of published
faunal lists for crassostreine oysters from the Pliocene
and early Pleistocene of Florida found only C. virginica
(see Mansfield, 1932, 1939; Olsson and Harbison, 1953;
DuBar, 1958, 1962; Stanley, 1986; Campbell, 1993).
Examination of the extensive fossil collections at the
Florida Museum of Natural History (FLMNH) found that
S. paucichomata is a common component of the Tamiami
Formation and also present in the Caloosahatehee Forma-
tion (lower Pleistocene) in southern peninsular Florida. All
of the type, figured and referred specimens are deposited
in the University of Florida, Florida Museum of Natural
History, Invertebrate Paleontology Collection and are
cataloged with the prefix UF and a lot number.
Specimens of Striostrea paucichomata found in the
Tamiami Formation were commonly found in association
with the oysters Hijotissa haitensis (G.B. Sowerhy I,
1850), Ostrea compressirostra Say, 1824, and Mijrakeena
sculpturata (Conrad, 1840) new combination. Oysters
less commonly associated with S. paucichomata were
Undulostrea locklini (Gardner. 1945) new combination,
C. virginica , Cuhitostrea coxi (Gardner, 1945), and
Dendostrea frons (Linnaeus, 1758). Mijrakeena sculpturata
was placed in the genus Conradostrea Ward and
Blackwelder, 1987 along with Mijrakeena lawrencei
(Ward and Blackwelder, 1987) new combination and
Mijrakeena g reeni (Ward, 1992) new combination.
The diagnostic shell characters of Conradostrea are the
same as those of Mijrakeena Harry, 1985 and therefore
Conradostrea should be considered a junior synonym
of Mijrakeena. The shell morphology of U. locklini is
consistent with the description of Undulostrea Harry,
1985 and therefore should be included in that genus.
Mijrakeena and Undulostrea have similar spatial and
temporal distributions as the genus Placunanomia
Broderip, 1832 (Anomiidae Rafinesque, 1815). These
three genera inhabited the eastern Pacific and western
Atlantic in the Pliocene and became extirpated from the
western Atlantic by the middle of the Pleistocene with
one or two species still extant in the eastern Pacific
[Mijrakeena angelica (Rochebrune, 1895); Undulostrea
megodon (Hanley, 1846); Placunanomia cumingii Broderip,
1832; Placunanomia panamensis Olsson, 1942].
Stenzel (1971: N1128) in his revision of the oysters
stated that fossil crassostreines (as nonineubatory genera
within the subfamily Ostreinae) “are recognized by their
left valve umbonal cavity and similarity to living
Crassostrea ' . Harry (1985: 149) in his revision of the living
oysters recognized the subfamily Crassostreinae pro-
posed by Torigoe (1981) and characterized the shells as
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THE NAUTILUS, Vol. 127, No. 2
“medium to large size, usually elongated dorsoventrally,
occasionally subcircular. The left valve is usually deeply
concave, and the right one is usually nearly flat. Shell
plications are usually limited to the left valve, often indif-
ferently developed or absent. The early part of the right
valve exterior has continuous growth of the outer shell
layer, and later it often forms fragile, appressed,
overlapping lamellae, but the outer surface is frequently
eroded during life, obliterating the sculpture. The
chomata are ostreine, or absent. The muscle scars tend
to be more darkly colored than the surrounding shell,
in one or both valves.”
Stenzel (1971) and Harry (1985) listed the presence
of a riblet-bearing surficial layer on the right valve and
the presence of chomata as characters that separate
Striostrea from Crassostrea. Stenzel (1971) described
the riblet-bearing surficial layer as: “This layer is thin
and delicate and flakes off readily. In fossil species, only
a few exceptionally well-preserved specimens retain it
on the outer face of the right valve (see Fig. 107,1c).
Commonly the layer is dark-colored because it is either
made entirely of conchiolin or is a prismatic calcite layer
rich in conchiolin. Riblets are restricted to this surficial
layer, and the immediately underlying, more calcareous
and lighter-colored layer shows a faint trace of them at
best. Because of its delicate consistency the riblet-bearing
layer is better preserved in very young and still fragile
oyster shells and dehisces in older individuals. Old indi-
viduals may show riblets only on the marginal conchiolin
fringes.” (p. N979) and “Right valve covered by many
thin, readily dehiscent, conchiolin-rieh imbricating
layers that have prismatic shell structure and carry on
their tops many narrow (1.3 mm. or less wide) dichoto-
mous flat-topped radial riblets separated by narrower
interspaces, riblets converging and diverging irregularly
from place to place, producing shaggy appearing surface,
becoming less abundant and less prominent in later
growth stages.” (p. N1136).
Stenzel (1971: N979) provided Striostrea alabamiensis
(I. Lea, 1833) as an example of an extinct species of
Striostrea based on the description and illustrations in
Harris (1919). Striostrea alabamiensis is actually a syno-
nym of Striostrea gigantissima (Finch, 1824) as proposed
by Harris (1919) and Lawrence (1995: 193). Harris (1919)
thought that Finch’s description was not sufficient to be
valid. Howe (1937) argued that Finch’s description was as
informative as those of some of his contemporaries and
should be considered valid. Other extinct erassostreine
species with a riblet-bearing surficial layer based on
the literature include Ostrea dorsata Deshayes, 1824
(Deshayes, 1824; J.D.C. Sowerby, 1850; Wood, 1861-
1871), Ostrea spatulata Lamarck, 1806 (Deshayes, 1824),
Ostrea tenera f. Sowerby, 1821 (J.D.C. Sowerby, 1850;
Wood, 1861-1871), Ostrea velata Wood, 1861 (Wood,
1861-1871) and Crassostrea cahobasensis (Pilsbry and
Brown, 1917) (Woodring, 1982).
Lawrence (1995) argued that all of the erassostreine
genera should be included in Crassostrea. Part of his
argument was based on the presence of chomata on
C. gigantissima and C. cahobasensis and a riblet-bearing
surficial layer on C. gigantissima. However, since both of
these species have chomata and a riblet-bearing surficial
layer and since these are currently considered diagnostic-
characters for the genus Striostrea , then at least for
Striostrea his argument is not valid.
Carter et al. (201 1 ) proposed placing the erassostreine
oysters in the family Flemingostreidae Stenzel, 1971.
According to Carter et al. (2011), this family contains
the extinct paraphyletic subfamilies Flemingostreinae
Stenzel, 1971 and Liostreinae Vialov, 1983 and the extant
subfamily Crassostreinae Scarlato and Starobogatov,
1979. Tbe living erassostreine oysters were originally
placed in their own family (Crassostreidae) by Scarlato
and Starobogatov (1979: 46) on the basis: “Non-incubating
oysters, because of the presence in them of such mor-
phological structures as a promyal cavity (promyal pas-
sage) and peculiarities of reproduction ” and “The family
is characterized by the development of a promyal cavity,
the pericardium shifted before tbe adductor muscle
anteriorly, and to the right, and the union of the anterior
part of the suprabranchial cavity with its excurrent
part. In the left (attached) valve there is usually a deep
subumbonal cavity. The eggs develop internally, the
sexes are separate (possibly protandric hermaphrodites,
but with a regular and complete sex change phase).” In
order to include the Flemingostreinae and Liostreinae
in the same family-group rank with the Crassostreinae,
the obligate choice for family name was Flemingostreidae
(Nikolaus Malchus, personal communication). See syno-
nymy list under the systematic^ section.
OCCURRENCES, STRATIGRAPHY, AND AGE
Over forty complete valves (some paired) of Striostrea
paucichomata Bolton new species have been examined
in the FLMNH Invertebrate Paleontology Collections.
All were derived from the two most densely-packed and
species-rich molluscan units of southern Florida; namely
the Pinecrest beds of tbe Tamiami Formation and the
Caloosahatchee Formation. The majority was recorded
from the Pinecrest beds in northwestern Sarasota County
with the remainder collected from Broward, Charlotte,
De Soto, and Hendry counties (see Figure 1).
Much confusion and controversy surround surface
and near-surface deposits (especially the Tamiami For-
mation) of southern Florida. As summarized by Jones
(1997: 107) this is because “Pliocene and Pleistocene
deposits in the region consist of siliciclastic and carbon-
ate lithologies whose lateral and temporal relationships
are obscured by 1) thinness and discontinuous distribu-
tion of units, 2) limited exposures, 3) rapid facies
changes, and 4) repeated advance and retreat of the sea
over this low-elevation region in response to the many
sea-level oscillations of the Plio-Pleistocene”. Addition-
ally, many stratigraphic units have been erected not based
on lithology (as now required by the North American
Stratigraphic Code) but fossil content (Scott, 1992).
M. J. Bolton and R. W. Ported, 2013
Page 67
defunct APAC Sarasota Mines (formerly Macasphalt Shell Pits,
Newburn Road Pit, Warren Brothers Pits) and active SMR
Aggregates Pits (formerly Wendell Kent Pit, Richardson Road
Shell Pits, Quality Aggregates Shell Pits). All counties - Sarasota
(S), De Soto (D), Charlotte (C), Hendry (H) and Broward (B)
where occurrences are known are shaded.
Herein, we use widely-accepted stratigraphic terminol-
ogy for both units (Tamiami and Caloosahatchee forma-
tions) in which S. paucichomata is known to occur (e.g.,
Zullo and Harris, 1992). For a more thorough review
of southern Florida stratigraphy refer to Lyons (1991).
The name “Tamiami limestone” was first applied by
Mansfield (1939) for deposits exposed during road con-
struction in Collier and Monroe counties. Parker and
Cooke (1944) broadened the concept of the Tamiami
limestone and designated it a formation. They also
included the sands near Pinecrest, as described in
Mansfield (1931), in their concept and concluded that
the Buckingham limestone of Mansfield (1939) was a
facies of the Tamiami Formation. Parker (1951) placed
the Buckingham limestone in the Tamiami Formation,
and Olsson (1964) informally proposed the “Pinecrest
beds” for fossil deposits younger than the Tamiami For-
mation and older than the Caloosahatchee marl. Hunter
(1968) divided the Tamiami Formation into five, major,
members based on lithostratigraphy; Bayshore Clay,
Murdock Station member, Pinecrest sand, Ochopee
limestone, and Buckingham limestone. She considered
the three youngest members, the Pinecrest sand,
Ochopee limestone, and Buckingham limestone, to be
lateral equivalents; her oldest member being the
Bayshore clay. However, Missimer (1992, p. 63) reported
that the Tamiami Formation “consists of at least nine
mappable members or facies” including the Pinecrest
Sand, Ochopee Limestone and Buckingham Limestone.
Due to the poorly defined, lithologically (carbonates,
siliciclastics, and mixed siliciclastics-carbonates), and
temporally complex nature of the Tamiami Formation,
Zullo and Harris (1992) employed sequence stratigraphy
to help unravel both its temporal and spatial relation-
ships. For the purposes of this study we follow their
nomenclature of this marine deposit, especially in the
type area of S. paucichomata (Figures 1 and 2).
Today, the Pinecrest beds are best exposed at
Sell roeder- Manatee Ranch Aggregates, Inc. (SMR)
excavations (formerly Richardson Road Shell Pits and
Quality Aggregates Shell Pits) in Sarasota County (Figure 1);
now that the more westward Ashland Petroleum and
Asphalt Corporation (APAC) Sarasota pits (formerly
Newburn Road Pit, Warren Brothers Pits, and
Macasphalt Shell Pits) are water-filled. At APAC, Petuch
(1982) divided the exposed beds into twelve units. Based
on the aforementioned sequence stratigraphic analysis
of Zullo and Harris (1992) at APAC and SMR, they
concluded that Petuch Units 0-1 belong to the Caloosa-
hatchee Formation and Units 2-11 were Tamiami
Formation. Units 2-9 were divided into the Upper
Tamiami Formation and Units 10-11 were Lower
Tamiami Formation. Further subdivision placed Units 2-3
into the upper Pinecrest beds and Units 4-9 into the
lower Pinecrest beds (see Figure 2). Herein, we follow
the stratigraphic organization proposed Zullo and Harris
(1992) although Petuch and Drolshagen (201 1 ) now con-
sider Units 2^4 to belong to the Fruitville
Member (Tamiami Formation), Units 5-9 to belong to
the Pinecrest Member (Tamiami Formation), Unit 10 to
belong to the Buckingham Member (Tamiami Forma-
tion), anti Unit I I to be the Sarasota Member (Murdock
Station Formation).
Jones et al. (1991) estimated the age of Petuch s (1982)
Units 2 4 as being 2.25 (+/— 0.25) Ma and Units 5-10
as being 3.0 (+/— 0.5) Ma based on s‘Sr/s,1Sr isotope
bivalve dating, paleomagnetism, and invertebrate and
vertebrate biochronology. Allmon (1993) concluded
that Units 5-10 are upper Pliocene (3. 0-3.5 Ma) and
Units 2-4 are much younger (2. 0-2. 5 Ma). Gibbard
et al. (2009) places the boundary between the Pliocene
and the Pleistocene at 2.588 Ma. Therefore, Units 2-4
with a minimum age of 2.0 Ma and maximum age of
2.5 Ma is lower Pleistocene. Units 5-10 with a minimum
age of 2.5 Ma and a maximum age of 3.5 Ma is mostly,
if not wholly, upper Pliocene. The underlying Unit If is
therefore at least upper Pliocene.
The name “Caloosahatchee beds of marls” was applied
by Dali (1887) for shell horizons exposed along the
Caloosahatchee River. Matson and Clapp (1909) later
referred to the unit as “Caloosahatchee marl” and DuBar
(1974: 216) elevated it to formational status because “of
the diversity of lithologies and the vagueness of the term
marls". Today, the southern peninsular Florida marine,
brackish, and freshwater units which are younger than
the Tamiami Formation and older than the Bermont
Formation are placed within this unit (DuBar, 1974).
Page 68
THE NAUTILUS, Vol. 127, No. 2
Figure 2. Stratigraphic nomenclature for the type area of Striostrea paucichomata Bolton new species in Sarasota County
modified from Zullo and Harris (1992) and incorporating stratigraphic units of Petuch (1982). The new species has thus far been
recorded irom Units 3, 5 through 8 and 10.
The Caloosahatchee Formation disconformably over-
lies the Tamiami Formation and has been estimated to
be about 1.8 Ma using He/U coral dating (Muhs et al.,
1992). This places the unit in the late lower Pleistocene.
SYSTEM ATICS
Class Bivalvia Linnaeus, 1758
Order Ostreida Ferussac, 1822 in 1821-1822
Superfamily Ostreoidea Rafinesque, 1815
Family Flemingostreidae Stenzel, 1971
Flemingostreini Stenzel, 1971
Crassostreidae Searlato and Starobogatov, 1979
Crassostreini Chiplankar and Badve, 1979
Crassostreinae Torigoe, 1981
Crassostreinae Freneix, 1982
Liostreinae Vialov, 1983
Subfamily Crassostreinae Searlato and Starobogatov, 1979
Tribe Striostreini Harry, 1985
Genus Striostrea Vialov, 1936
Type Species: Ostrea procellosa Lamy, 1929, which is
a junior synonym of Ostrea margaritacea Lamarck, 1819.
Recent, along the coast of South Africa and the western
Indian Ocean as far north as the Arabian Peninsula
(Huber, 2010).
Striostrea paucichomata Bolton new species
(Figures 3-12)
Diagnosis: Right valve with riblet-bearing surfieial
layer (only visible on exceptionally well-preserved spec-
imens), weakly developed ehomata usually present on
both valves (not visible in about 30% of specimens), left
valve external surface usually with characteristic irregu-
lar and undulating growth intervals, maximum height
about 8 cm, posterior adductor muscle imprint situated
about mid-point between ventral edge of hinge and
ventral margin of shell.
Description: Shell usually elongate dorsoventrally
(Figures 3-6, 1 1-12). Holotype maximum dimensions:
left valve 6.56 cm high, 3.08 cm long, 1.72 cm wide;
right valve 5.58 cm high, 2.33 cm long, 0.2 cm wide.
Maximum height about 8 cm. Left valve usually without
extensive attachment area; usually moderately to
deeply concave; umbonal cavity weakly to strongly
developed depending on degree of valve concavity;
external surface usually with characteristic irregular
and undulating growth intervals (Figure 3). Right valve
M. J. Bolton and R. W. Portell, 2013
Page 69
Figures 3-6. Paired valves of Striostrea paucichomata Bolton new species. Holotype (UF 34779). 3. Exterior of left valve.
4. Interior of left valve. 5. Exterior of right valve. 6. Interior of right valve.
Page 70
THE NAUTILUS, Vol. 127, No, 2
Figures 7-12. Striostrea paucichomata Bolton new species. 7. Exterior detail of right valve near hinge showing the riblet-bearing
surficial layer, Holotype (UF 34779). 8. Interior detail of right valve near hinge showing anachomata (see arrows). Holotype (UF
34779) 9. Interior detail of left valve near hinge showing catachomata (see arrows), same specimen as Figure 10. Paratype (UF
200351). 10. Exterior of left valve showing variation in appearance. Paratype (UF 200351)- 11. Exterior of left valve showing variation
in appearance, same specimen as Figure 12. Paratype (UF 29811). 12. Interior of left valve. Paratype (UF 29811).
M. J. Bolton and R. W. Portell, 2013
Page 7 1
usually flat; with regular, closely spaced growth lines
(Figure 5); exceptionally well-preserved specimens
with riblet-bearing surficial layer consistent with the
description in Stenzel (1971: N979, N1136) (Figure 7),
may only be present near hinge, riblets 0. 1-0.2 mm
wide. Both valves usually with weakly developed
ostreine chomata (Figures 8-9), not visible in about
30% of specimens, only present near hinge, older spec-
imens usually with relict chomata or chomata absent,
anachomata 0. 1-0.3 mm wide with 0. 1-1.5 mm gap
between them, chomata may be difficult to see without
magnification. Shell not thick as S. gigantissima and
S. cahobasensis commonly are. Posterior adductor
muscle imprint situated about mid-point between ven-
tral edge of hinge and ventral margin of shell (compared
to ventral to the mid-point in Crassostrea virginica ),
usually semilunar in outline (Figures 4, 6, 12).
Holotype (Figures 3-8): UF 34779, left and right
valves (pair), USA, Florida, Sarasota Co., Macasphalt
Shell Pit (SO001), T36S, RISE, Plio-Pleistocene, spoil,
1 Nov. 1986, R.J. Britt, Jr.
Paratypes: UF 200351, two left valves, USA, Florida,
Sarasota Co., Quality Aggregates Phase 07 (SO022),
T36S, R19E, Pliocene, upper Tamiami Formation, upper
Pinecrest beds, Petueh Unit 3?, 7 June 1994, R. Portell
et al. (Figures 9-10); UF 216676, one left valve, USA,
Florida, Sarasota Co., Macasphalt Shell Pit B (SO017),
T36S, RISE, Pliocene, upper Tamiami Formation, lower
Pinecrest beds, Petueh Unit 5, 16 March 1988, R. Portell
and D. Jones; UF 38987, two left valves, USA, Florida,
Sarasota Co., Macasphalt Shell Pit B (SO017), T36S,
R18E, Pliocene, upper Tamiami Formation, lower
Pinecrest beds. Section 2, Petueh Unit 6, 16 March
1988 R. Portell and D. Kendrick; UF 53225, three left
valves, USA, Florida, Sarasota Co., Richardson Road
Shell Pit 01B (SO013), T36S, R19E, Pliocene, upper
Tamiami Formation, lower Pinecrest beds, Petueh Unit 7,
19 April 1991, R. Portell and D. Jones; UF 178522, two
right valves, USA, Florida, Sarasota Co., Quality Aggre-
gates Phase 8 REU-2 (SO049), T36S, R19E, Pliocene,
upper Tamiami Formation, lower Pinecrest beds, REU
Unit 2A, 3-1 June 2006, USF REU; UF 53629, one
right valve, USA, Florida, Sarasota Co., Richardson
Road Shell Pit 01B (SO013), T36S, R19E, Pliocene,
upper Tamiami Formation, lower Pinecrest beds,
Petueh Unit 8, 19 April 1991, R. Portell and D. Jones;
UF 95889, one left valve, USA, Florida, Sarasota Co.,
Richardson Road Shell Pit 01C (SO021), Pliocene, lower
Tamiami Formation, Petueh Unit 10, R. Portell and
D. Jones; UF 29811, left and right valves (pair), USA,
Florida, Sarasota Co., Macasphalt Shell Pit (SO001),
T36S, RISE, Plio-Pleistocene, spoil, 1969-1978, E. and
E. Bradley (Figures 11-12).
Additional Specimens from Other Locations or
Formations: UF 93046, one right valve, USA, Florida,
Charlotte Co., Acline Shell Pit '(CH010), T41S, R23E,
Pliocene, Tamiami Formation, Pinecrest beds, Florida
Geological Survey/C. R. Loeklin; UF 208478, two left
valves, USA, Florida, Hendry Co., Interceptor Canal 01
(HN027), T48S, R34E, Pliocene, Tamiami Formation,
Pinerest beds, 1968, II. K. Brooks and D. Townsend; UF
208483, two right valves, USA, Florida, Broward Co.,
south of Seminole Indian Reservation headquarters,
just north of lock on drainage canal (5422), Pliocene,
Tamiami Formation, Pinecrest beds, 1969, H.K. Brooks
et al.; UF 200355, three left valves and two right valves,
USA, Florida, De Soto Co., De Soto Shell Pit 05
(DE010), T39S, R25E, lower Pleistocene, Caloosa-
hatchee Formation, Portell Bed 7, 7 March, 1991, R.
Portell and K. Schindler; UF 200354, two right valves,
USA, Florida, Hendry Co., Caloosahatchee River 09,
T45S, R28E, early Pleistocene, Caloosahatchee Forma-
tion, DuBar Horizon 5, 1953, J. DuBar; UF 2654,
left and right valves (pair) and one right valve, USA,
Florida, Hendry Co., Caloosahatchee River 01
(HN002), T43S, R28/29E, lower Pleistocene, Caloosa-
hatchee Formation, spoil, J.C. Macbeth.
Occurrence: Striostrea paucichomata is known from
the upper Pliocene to lower Pleistocene Tamiami For-
mation and lower Pleistocene Caloosahatchee Forma-
tion in Sarasota (type area), Charlotte, De Soto, Hendry
and Broward counties, Florida (Figure 1). Specimens
have been found at the type location in the upper
Tamiami Formation, Pinecrest beds in Petueh Units 3,
5, 6, 7, 8 and lower Tamiami Formation Unit 10.
Etymology: The species name is derived from the
Latin panel meaning few and “chomata” which are the
tubercles and pits on the periphery of inner surface of
shells, usually near the hinge. This name is in reference
to the usually low number of chomata that are often
difficult to see or absent.
Discussion: Identification of fossil oysters has been
confused and neglected due to the high amount of
morphological variation associated with environmental
factors. This condition has led to the publication of
many synonyms based on either different eeophenotypes
or the differences between young and old specimens
(Stenzel, 1963). The proliferation of species names has
also been the result of some authors describing new
species based on a very limited number of specimens.
However, given adequate material and using the charac-
ters described in Stenzel (1971) and Harry (1985), accu-
rate generic identifications of most Cenozoic oysters
should be possible.
Use of the genus Striostrea for erassostreine oysters
with a riblet-bearing surficial layer and chomata has
not been widely used in North America. This may par-
tially be due to the riblet-bearing layer being fragile
and only present on exceptionally well-preserved spec-
imens and the chomata are not visible on all specimens
either because they were only present on young spec-
imens or are present as relict or active chomata only on
a percentage of older individuals. For S. paucichomata
these characters are also often difficult to see without
Page 72
THE NAUTILUS, Vol. 127, No. 2
Figures 13-16. Striostrea gigantissima (Finch, 1824). 13. Exterior of left valve, same specimen as Figure 14 (UF 228893).
14. Exterior of right valve (UF 228893). 15. Exterior of right valve, same specimen as Figures 16 and 17 (UF 228894).
16. Interior of right valve (UF 228894).
M. J. Bolton and R. W. Portell, 2013
Page 73
Figures 17-21. Striostrea gigantissima (Finch, 1824) (17, 18) 17. Interior detail of right valve near hinge showing anachomata
(see arrows). (UF 228894). 18. Exterior of right valve showing the riblet-bearing surlicial layer. (UF 228895). Striostrea cahohasensis
(Pilsbry and Brown, 1917) (19-21 ). 19. Exterior of left valve, same specimen as Figures 20 and 21 (UF 191980). 20. Interior of
left valve (UF 191980). 21. Interior detail of left valve near hinge showing catachomata (see arrows). (UF 191980).
Page 74
THE NAUTILUS, Vol. 127, No. 2
magnification. For these reasons, to accurately identify
fossil Striostrea it is important to have sufficient
numbers of exceptionally well-preserved specimens
including ones of different age classes.
Fossil Striostrea known from the western Atlantic
include S. gigantissima (Finch) (lower Eocene through
upper Oligoeene; USA: North Carolina-Texas), S.
cahobasensis (Pilsbry and Brown) new combination
(upper Oligoeene through middle Miocene; Caribbean:
Venezuela, Panama, Puerto Rico, Haiti, Mexico; USA:
FL), and S. paucichomata Bolton new species (upper
Pliocene through lower Pleistocene; USA: FL). Photos
of young specimens of S. gigantissima (Figures 13-18)
from the upper Eocene of Georgia and S. cahobasensis
from the lower Miocene (Figures 19-21) and upper Oli-
gocene (Figures 22, 23) of Florida are provided for
comparison. A key for the Cenozoic crassostreine oysters
known from southeastern United States is provided
below. There are also specimens of a large crassostreine
oyster in the FLMNH Invertebrate Paleontology Collec-
tion from the Pliocene of Curasao (an island off the coast
of Venezuela) that may be a Striostrea. Young specimens
have chomata (UF 114702, UF 1 16000, UF 116005), but
no surface riblets were present on the right valves.
However, the specimens were not sufficiently preserved
to exhibit a riblet-bearing surfieial layer. Furthermore,
they very much resemble the eastern Pacific species
Crassostrea titan (Conrad, 1853) in shape and size. Addi-
tional study is required to determine if they are con-
specific. The complete temporal and paleogeographic
distribution of Striostrea will not be known until
museum collections are reexamined using the characters
and caveats discussed in this paper. Four living species
of Striostrea are known from the eastern Atlantic Ocean
along the coast of tropical West Africa, the coast of
South Africa and western Indian Ocean along the coast
of East Africa as far north as the Arabian Peninsula,
northern Pacific Ocean from japan to Taiwan, and trop-
ical eastern Pacific Ocean (Huber, 2010).
Key to Cenozoic Crassostreine Oysters Known
From the Southeastern United States:
1 Shell without chomata; right valve without a riblet-
bearing surfieial layer (visible only on exceptionally
well-preserved specimens); shell may have costae
(primarily on the left valve) and may have plicae
along the ventral margin; posterior adductor
muscle imprint usually situated ventral to the
mid-point between ventral edge of hinge and ven-
tral margin of shell; upper Oligoeene - present.
Crassostrea virginica (Gmelin, 1791)
1 ' Shell with ostreine chomata (may be absent in older
shells and various ecomorphs) (Figures 8, 9, 17, 21,
23); right valve with a riblet-bearing surfieial layer
(visible only on exceptionally well-preserved
Figures 22-23. Striostrea cahobasensis (Pilsbry and Brown, 1917). 22. Exterior of left valve, same specimen as Figure 23 (UF
27389). 23. Interior detail of left valve near hinge showing cataehomata (see arrows). (UF 27389).
M. J. Bolton and R. W. Ported, 2013
Page 75
specimens, especially younger ones) (Figures 7,
18); shell usually without costae or plicae 2
2(F) Both valves of similar convexity (especially old
shells) or left valve slightly to moderately more
convex and capacious; maximum shell height about
56 cm; shell may be extremely thick (see Harris,
1919: Pis. 1-6; Howe, 1937: Pi. 44 Figs. 1-6;
Toulmin, 1977: Pi. 14 Figs. 5, 6, Pi. 15 Figs. 1, 2,
Pi. 56 Fig. 7); lower Eocene - upper Oligo-
cene Striostrea gigantissima (Finch, 1824)
2' Left valve usually more convex and capacious than
right valve; maximum height less than 20 cm 3
3(2') Chomata moderately developed or absent
(Figures 21, 23); shell maximum height about 19.5
cm; shell may be thick (see Pilsbry and Brown,
1917: Pi. 6 Figs. 1, 8; Woodring, 1982: Pi. 90 Fig.
21, Pi. 93 Figs. 6, 7, 9-1 1, Pi. 94 Figs. 1, 3, 5, Pi. 102
Figs. 1,5, PI. 103 Fig. 8, PI. 106 Figs. 2, 6, 7); upper
Oligoeene - middle Miocene Striostrea
cahobasensis (Pilsbry and Brown, 1917)
3' Chomata weakly developed or absent (Figures 8,
9); shell maximum height about 8 cm; shell not
thick (Figures 3-6, 11-12); upper Pliocene - lower
Pleistocene Striostrea paucichomata
Bolton new species
ACKNOWLEDGMENTS
We thank Alex Kittle and Sean Roberts (Florida
M useum of Natural History) and Jose H. Leal (The
Bailey- Matthews Shell Museum) for collection assis-
tance; Sean Roberts for photographing the figured
specimens; Nikolaus Malchus (Universitat Autonoma
de Barcelona) for discussions on oyster morphology and
taxonomy; and Jason and David Bolton, Douglas Jones,
David Kendrick and Kevin Schindler for early field
assistance. RWP thanks the Florida Geological Survey
for transfer of invertebrate paleontology collections
formerly under their care which now reside at the
Florida Museum of Natural History and Schroeder
Manatee Ranch Aggregates Incorporated’s President
(Eugene Henshaw, Jr.), Mine Supervisor (Dave Myers),
Plant Superintendent (Steve Stombaugh), and Office
Manager (Nancy Plank) for permission and assistance
with quarry collecting. Financial support for RWP’s
fieldwork was provided by the McGinty Endowment
and by Barbara Toomey and James Toomey. This is
University of Florida Contribution to Paleobiology 647.
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THE NAUTILUS 127(2):78-84, 2013
Page 78
Rediscovery of Choanopoma? smithianum Pfeiffer, 1866
(Annulariidae) from Haiti and designation of a neotype,
with the description of two new species
of Weinlandipoma Bartsch, 1946
G. Thomas Watters
Department of Evolution, Ecology,
and Organismal Biology
Ohio State University
Columbus, OH 43212 USA
Watters [email protected]
Jozef Grego
Horna Micina 219
SK— 974 01
Banska Bystrica, Slovak Republic
Jozef Steffek1
Department of Applied Ecology
Faculty of Ecology and Environmental Science
Technical Univerzity
Zvolen, Slovak Republic
ABSTRACT
Choanopoma ? smithianum Pfeiffer, 1866, was described from
“monte Platon" on the western end of the Tiburon Peninsula
of Haiti. The species has never been figured and the type spec-
imens are apparently lost. Surveys of the region in 1984 and
2006 rediscovered this “lost" species, which is here allocated
to Weinlandipoma Bartsch, 1946. A neotype is designated based
on this material, the species is redescribed, and compared with
similar taxa. Two additional species are described as new,
Weinlandipoma auduboni and Weinlandipoma macaijaense.
Additional Keywords: Gastropoda, Annulariidae, Hispaniola,
Weinlandipoma , new species
INTRODUCTION
Ludwig Pfeiffer often described species without immedi-
ate illustration although some taxa might subsequently be
figured by him or others. In 1866 Pfeiffer described
Choanopoma ? smithianum from “in monte Platon insulae
Haiti.” He questioned whether his species belonged
in Choanopoma because none of Iris five specimens
retained an operculum, a diagnostic characteristic of the
genus. The species was never figured. Although the species
was mentioned again by Pfeiffer (1876) and listed by Tryon
(1867), Kobelt (1880), and Crosse (1891), no additional
information was given. Bartsch, in Iris 1946 review of tire
Hispaniolan annulariids, uncharacteristically overlooked
this name. Watters (2006), based on tire facts that the type
could not be located at The Natural History Museum (UK)
or tire Berlin Museum fur Naturkunde, where much
of Pfeiffer’s material that had not been destroyed during
World War II resided, turd drat it had never been illus-
trated, suggested diat dre taxon was a nomen (labium. But
1 Posthumously
investigation of the type locality (Grego and Steffek, 2007)
revealed a species that undoubtedly represents Pfeiffer’s lost
Choanopoma? smithianum as well as previously unrecog-
nized taxa.
Pfeiffer’s type locality of “in monte Platon insulae
Haiti” was stated to be “30 engl. Meilen nordostliclr von
Aux-Cayes.” This measurement is based on road miles,
not straight distance. This site is identifiable as the area
of the well-known Citadelle de Platons, on “monte
Platon ” located on the Plain of Fornron south of Pic
Macaya (dre second highest mountain in Haiti at 2,347 m)
and Pic Formon (2,219 m). It is reasonable to assume drat
the original collector had visited dre area to view this
impressive fortress. Citadelle de Platons was used during
the Haitian Revolution (1791-1804) to guard against
Napoleons expeditionary force led by Charles Leelere
to recapture tire French colony and reinstate slavery. There
is also an adjacent village of Les Platons. Collections
at Citadelle de Platons and 27 otirer neighboring sites near
Monre Cavalier, Cay Demiere Jeudi, and dre Pic Ouasac
ridge revealed only four annulariid species: Weinlandipoma
gonavense (Weinland, 1880), and three additional species
of Weinlandipoma. Pfeiffers brief description could only
apply to one of the four species encountered based
on circumstantial evidence presented here. We believe
drat the four Weinlandipoma taxa near and at dre type
locality are Weinlandipoma smithianum, Weinlandipoma
gonavense , Weinlandipoma auduboni, new species, and
Weinlandipoma macaijaense, new species.
Clench (1935) collected in the general area but reported
only on dre Urocoptidae. Thompson (1986) collected exten-
sively in the (dren proposed) Parc National Pic Macaya. This
material is at the Florida Museum of Natural History (UF)
and is included in dris study. The collections by Grego and
Steffek at drirteen sites and the collections by Thompson
at eleven sites resulted in 1,558 specimens of four
Weinlandipoma species. Some of these sites were figured
and discussed in more detail in Grego and Steffek (2007).
G. Thomas Watters et al., 2013
Page 79
MATERIALS AND METHODS
Unless noted otherwise, all sites are in the Department
du Sud, Haiti. Table 1 lists the material examined and
the collections sites. Descriptions and measurements
were based on shells oriented with the spire up and the
aperture facing the viewer. Minimum and maximum
dimensions were based on all adult specimens available.
Abbreviations used in text are: BMSM, The Bailey-
Matthews Shell Museum, Sanibel, Florida, USA; GTW,
Collection of G.T. Watters, USA; Grego coll., Jozef Grego
Collection, Slovak Republic; NHMUK, The Natural History
Museum, London, UK; OSUM, Ohio State University
Museum of Biological Diversity, Columbus, USA; UF,
Florida Museum of Natural History, Gainesville, USA.
SYSTEMATICS
Family Annulariidae Henderson and Bartsch, 1920
Subfamily Annulariinae Henderson and Bartsch, 1920
Genus Weinlandipoma Bartsch, 1940
Type Species: Choanopoma blandii Weinland, 1880,
by original designation.
Weinlandipoma smithianum (Pfeiffer, 1866)
(Figures 1-4, 18)
Choanopoma ? Smithianum Pfeiffer, 1866: 88-89; Pfeiffer,
1876: 157; Watters, 2006: 484, 557 [as a nomendubium\.
Choanopoma smithiana Pfeiffer, 1866. Tryon, 1867: 99.
Table 1. Localities and specimens. A, Weinlandipoma auduboni ; G, W. g onavense- M, W. macaijaense ; S, W. smithianum
Locality A
Grego coll., Citadelle de Platons, cleared area around fortress, 727 m elevation, 1
18.2702° N, -73.9709° W.
Grego coll., near Morne Cavalier, sinkholes in wet limestone forest, 1,1 16 m elevation, 3
18.3227° N, -74.0204° W
Grego coll., near Morn Cavalier, Parc National Pic Macaya, foot of Mt. Ouasac, 4
wet limestone outcrops in cleared forest, 1,341 m elevation, 18.3391° N, —74.0304° W
Grego coll., Pic Formon, Parc National Pic Macaya, limestone boulders in cleared 0
wet limestone forest, 1,200 m elevation [coordinates unknown]
Grego coll., Cay Derniere Jeudi, Parc National Pic Macaya, wet limestone outcrops 0
at edge of forest, 1,306 m elevation, 18.3397° N, -74.0210° W
Grego coll., Cay Derniere Jeudi, Parc National Pic Macaya, wet limestone outcrops 0
at edge of forest, 1,331 m elevation, 18.3397° N, —74.0210° W
Grego coll., near Morne Cavalier, 1 km W of Cay Michel, Parc National Pic Macaya, 2
wet limestone outcrops, 1,191 m elevation, 18.3267° N, —74.0297° W
Grego coll., Pic Formon, off path between Morne Cavalier and Cay Michel, Parc National 0
Pic Macaya, wet limestone outcrops, 1,202 m elevation, 18.3254° N, —74.0262° W.
Grego coll.. Pic Ouasac, deforested pine slope, Parc National Pic Macaya, 2,122 m elevation, 0
18.3576° N, -74.0279° W
Grego coll., near Morne Cavalier, 2 km W of Cay Michel, Parc National Pic Macaya, 0
wet limestone outcrops, 1,258 m elevation, 18.3267° N, —74.0297° W
Grego coll.. Pic Ouasac, Parc National Pic Macaya, deforested pine slope, 1,818 m elevation, 14
18.3519° N, -74.0282° W
Grego coll.. Pic Ouasac, Parc National Pic Macaya, deforested pine slope, 1,818 m elevation, 0
18.3519° N, -74.0282° W
Grego coll.. Pic Formon, Parc National Pic Macaya, around cave 700 m W of school, 0
cleared wet limestone forest, 1,020 m elevation, 18.3247° N, —74.0130° W
UF 77446, 77448, 77453, Morne Formond, Parc National Pic Macaya, 1,650 m elevation, 0
ca. 18.3525° N, -74.0229° W
UF 32226, 6 km E of Cavaillon, 200 m elevation, 18.3035° N, -73.6013° W 0
UF 32278, Camp Perrin, 200 m elevation, 18.3274° N, -73.8602° W 0
UF 32269, 1 km S of Camp Perrin, 200 m elevation, 18.3171° N, -73.8601° W 0
UF 32732, Trou Wocli Sa Wo, ca. 1 km SSW of Camp Perrin, 18.3188° N, -73.8553° W 0
UF 32323, 7 km NW of Port Saint, 18.1433° N, -73.9647° W 0
UF 32385, 10 km NW of Port Salut, 18.1663° N, -73.9797° W 0
UF 33265, 6 km SSE of Fond des Blancs, 210 m elevation, 18.2341° N, -73.0984° W 0
UF 33375, 3 km E of La Vallee, Nippes Department, ca. 18.4333° N, -73.4346° W 0
UF 32361, 1 km SE of Roche- a-Bateau, 18.1785° N, -73.9913° W 0
UF 48134 - Plaine Sa Wo, [not located, presumably near Trou Woch Sa Wo, 0
ca. 1 km SSW of Camp Perrin, 18.3188° N, -73.8553° W.]
Totals 24
G M
4 1
102 10
60 5
351 0
366 0
114 0
20 6
17 9
73 0
20 0
33 0
77 0
21 0
26 0
23 0
9 0
38 0
8 0
7 0
5 0
30 0
6 0
3 0
0 0
1413 31
S
3
0
0
0
0
0
0
32
0
0
1
0
0
0
0
0
53
0
0
0
0
0
0
1
90
Page 80
THE NAUTILUS, Vol. 127, No. 2
Figures 1-19. Weinlandipoma and their distribution. 1-4. Weinlandipoma smithianum (Pfeiffer, 1866). All from Citadelle de
Platons. 1. Grego coll., 9.5 mm length; 2-3. Neotype of Choanopoma smithianum Pfeiffer, 1866. NHMUK 20120258, 9.6 mm
length; 4. Grego coll., 10.6 mm length. 5-8. Weinlandipoma auduboni new species. 5-6. Holotype UF 451538, near Morne Cavalier,
1 km W of Cay Michel, 7.6 mm length. 7. Paratype OSUM 37268, Citadelle de Platons, 6.8 mm length; 8. Paratype BMSM 17935,
near Morne Cavalier, 1 km W of Cay Michel, 7.2mm length. 9-12. Weinlandipoma macatjaense new species. 9-10. Holotype UF
451540, near Morne Cavalier, 1 km W of Cay Michel, 9.1 mm length; 11. Paratype OSUM 37269, Citadelle de Platons. 9.0 mm
length; 12. Paratype BMSM 17936, near Morne Cavalier, 1 km W of Cay Michel, 7.4 mm length. 13-17. Weinlandipoma g onavense
Bartsch, 1946. 13. Grego coll., Mt. Formon, 8.7 mm length 14. Grego coll., Citadelle de Platons. 10.6 mm length; 15. Grego coll.
Mt. Formon. 9.9 mm length. 16. Grego coll. Mt. Formon, 8.9 mm length. 17. Grego coll., Mt. Ouasac, 17 mm length. 18. Radula
of Weinlandipoma smithianum (Pfeiffer, 1866), UF 48134, Plaine Sa Wo. 19. Distribution of Weinlandipoma. Areas are approxima-
tions. A - W. meridianum Bartsch, 1946. B - W. milleri Bartsch, 1946. C - W. blandii (Weinland, 1880). D - W. orcuttii Bartsch, 1946.
E - W. strictecostatum (Maltzan, 1888). F - W. excisum Bartsch, 1946. G - W. gonavense gonavense (Weinland, 1880). H - W. gonavense
ssp. of Bartsch, 1946. I — W. gonavense conception Bartsch, 1946. J - W. gonavense robustum Barstch, 1946. K - W. smithianum
(Pfeiffer, 1866), W. macatjaense new species, W. auduboni new species. Base map courtesy of NASA.
G. Thomas Watters et ah, 2013
Page 81
Choanopoma smithianum Pleiffer, 1866. Kobelt, 1880:
277; Crosse, 1891: 165.
Redescription: Shell small (neotype 9.6 mm length,
decollate x 5.8 mm maximum width including peri-
stome), elongate conic. Protoconch unknown, decollate
in adults. Teleoconch of 4-4.25 whorls. Axial sculpture
of weakly scalloped or undulating cords, regularly spaced,
65-75 on f inal whorl, more widely spaced on early whorls.
On final and penultimate whorls these are grouped
in bundles of 4-5 cords where first cord is barely discern-
able and then each subsequent one increases in size until
last cord is longest, then series begins again; this results
in an smoothly undulating sutural line. Sculpture is more
randomly spaced on earlier whorls. Primary spiral sculp-
ture present only as very weakly scalloped margins of axial
cords on earliest whorls and in umbilicus resulbng
in an angular aspect to base. Secondary spiral sculpture
of numerous, microscopic threads between axial cords,
often worn away. Suture strongly indented. Aperture
double (rarely single), circular (3.5 mm in maximum
width in neotype), barely adnate. Inner lip smooth, erect.
Outer lip broadly expanded, slightly narrower facing
umbilicus, consisting of numerous fused lamella, slightly
auriculate. Shell dirty' white to pale lavender with 7-8 spiral
rows of brown spots and dashes; spots do not continue
as bands over lip. Operculum multispiral with a slightly
oblique, erect calcareous lamella. Radula taenioglossate
with digitate first marginal (Figure 18).
Neotype: NHMUK 20120258 (Figures 2, 3).
Type Locality: The neotype is from the type locality
“in monte Platon insulae Haiti," here interpreted as
Citadelle de Platons, specifically the cleared area around
the fortress, 727 m elevation. Department du Sud, Haiti,
18.2702° N, -73.9709° W.
Distribution: Specimens are only known from the
type locality. Pic Formon, and near Camp Perrin (UF
32254, 48134). It probably has a narrow range in the
southern foothills of the Pic Macaya range of the Massif
de la Hotte.
Variation in specimens: Ninety specimens seen. The
degree of color pattern intensity varies from dark to
barely discernible.
Comparison with Other Species: Weinlandipoma
smithianum, W. macayaense, and W auduhoni are most
similar to each other and to W. orcuttii Bartseh, 1946,
described from a hill north of Coteaux. That species is
distributed along the southwestern edge of the Tiburon
Peninsula from Coteaux to as far east as Aquin. It appears
to be a coastal, lowland species whereas W. smithianum,
W. macayaense , and W. auduhoni are montane.
Weinlandipoma orcuttii differs from W. smithianum,
W. macayaense , and W. auduhoni in sculptural differences.
In W orcuttii the axial ribs produce swollen sutural
cusps of a uniform length resulting in a regular, even
row of denticles lining the suture. In W. smithianum , W.
macayaense , and W. auduhoni these ribs are not of
uniform length and are gathered in bundles forming
an undulating or ragged row of denticles at the suture.
Weinlandipoma smithianum, W. macayaense, and W.
auduhoni also bear a striking resemblance to Articulipoma
fluxum (Bartseh, 1946), a species Bartseh described as
“exceedingly puzzling” (p. 48). It extends across most of
the Tiburon Peninsula from Port-au-Prince to Les Cayes
but has not been found in the Citadelle de Platons area. It
differs from Weinlandipoma in having a paueispiral, flat,
chondroid operculum rather than a multispiral operculum
with an erect lamella and in lacking microscopic spiral
tiireads between the axial sculpture. However, the overall
shell sculpture is very similar.
Weinlandipoma smithianum differs from W. auduhoni
and W. macayaense in being larger with much weaker
scalloped axial sculpture, a peculiar pattern of sutural
sculpture, and a barely adnate aperture.
Weinlandipoma auduhoni new species
(Figures 5-8)
Description: Shell small (holotype 7.6 mm length,
decollate x 4.4 mm maximum width including peri-
stome), short conic. Protoconch lost in all examples.
Teleoconch of 3.75 whorls. Axial sculpture of undulating
and strongly scalloped, narrow, erect lamella, 60-70 on
final whorl, more widely spaced on early whorls. These
are often grouped in bundles of 2-3 lamella into unfused
cusps at suture, separated by a wide space containing
1-3 very fine threads that do not form cusps, but this
pattern is not consistent on entire shell; this results in a
ragged sutural line. Sculpture is more widely spaced on
earlier whorls. Primary spiral sculpture present only as
strongly scalloped margins of axial lamella; three scallops
on spire whorls, seven on final whorl, three of these are
prominent in umbilicus. Secondary spiral sculpture of
numerous, microscopic direads between axial lamella,
often worn away. Suture strongly indented. Aperture
double, circular (2.7 mm in maximum width in holotype),
always solute. Inner lip smooth, slightly erect. Outer lip
expanded, narrower facing umbilicus, consisting of fine,
fused lamella, slightly auriculate. Shell dirty white with
6-7 spiral rows of widely separated, small, brown spots
and dashes axially aligned; spots continue as bands over
both sides of lip and form spots on inner lip. Operculum
multispiral with a vertical, erect calcareous lamella.
Holotype: UF 451538.
Type locality: Near Morne Cavalier, 1 km W of Cay
Michel, Parc National Pic Macaya, wet limestone out-
crops, 1,191 m elevation. Department du Sud, Haiti,
18,3267° N, -74.0299° W.
Paratypes: UF 451541, near Morne Cavalier, foot of
Mount Ouasac 1,341 m elevation. Department du Sud,
Haiti, 18,3391° N, -74.0304° W (1 specimen); OSUM
37268, Citadelle de Platons, cleared area around fortress,
727 m elevation. Department du Sud, Haiti, 18.2702° N,
Page 82
THE NAUTILUS, Vol. 127, No. 2
—73.9709° W (1 specimen); BMSM 17935, near Morne
Cavalier, 1 km W of Cay Michel, Pare National Pic
Macaya, wet limestone outcrops, 1,191 m elevation.
Department du Sud, Haiti, 18.3267° N, —74.0297° W
(1 specimen).
Other Material Examined: Grego coll., Pic Ouasac,
Parc National Pic Macaya, deforested pine slope, 1,818 m
elevation, 18.3519° N, -74.0282° W (14 specimens);
Grego coll., near Morne Cavalier, sinkholes in wet
limestone forest, 1,116 m elevation, 18.3227° N,
—74.0204° W. (3 specimens); Grego coll., near Morn
Cavalier, foot of Mt. Ouasac, wet limestone outcrops in
cleared forest, 1,341 nr elevation, 18.3391° N, —74.0304°
W (3 specimens).
Distribution: Known from Mom Cavalier and Citadelle
de Platons between 727-1,818 m elevation; it probably
occurs throughout the middle elevations of die Pic
Macaya range.
Variation in Specimens: Twenty-four specimens seen.
The examples we have seen are very uniform in their charac-
teristics, differing only in tire number of axial lamella (60-70).
Comparison with Other Species. Weinlandipoma
auduboni differs from W. mcicayaense in being smaller
(6. 2-7. 2 mm decollate length vs. 7. 2-9. 6 mm for W.
mcicayaense), in having coarser sculpture, and in having
fewer axial lamella (60-70 vs. 70-90 for W mcicayaense) .
Etymology: Named for John James Audubon (17 85 —
1851), ornithologist, naturalist, and painter who was
born at nearby Les Cayes, and for the Societe Audubon
d' Haiti, without whose assistance these collections could
not have been made.
Weinlandipoma macatjaense new species
(Figures 9-12)
Description: Shell small (holotype 9. 1 mm length,
decollate x 5.4 mm maximum width including peri-
stome), elongate conic. Protoconch of 1.75 smooth,
prominent whorls, usually decollate in adults. Teleoconcli
of 3.75-4 whorls. Axial sculpture of scalloped lamella,
70-90 on final whorl, more widely spaced on early
whorls. On final and penultimate whorls these are
grouped in bundles of 3-5 cords with wide species
between bundles having 1-5 fine threads. Sculpture is
more randomly spaced on earlier whorls. Primary spiral
sculpture present only as scalloped margins of axial; 3-
4 scallops on spire whorls, 7-8 on final whorl, three
pronounced scalloped cords in umbilicus. Secondary
spiral sculpture of numerous, microscopic threads
between axial cords, often worn away. Suture strongly
indented. Aperture double, circular (2.8 mm in maxi-
mum width in neotype), barely solute. Inner lip smooth,
erect. Outer lip expanded, narrower facing umbilicus,
consisting of about five narrow, erect lamella, slightly
aurieulate. Shell dirty white with 6-7 spiral rows of
widely separated, small, brown spots axially aligned;
spots continue as bands over both sides of lip and form
spots on inner lip. Operculum multispiral with a verti-
cal, erect calcareous lamella.
Holotype: UF 451540.
Type locality: Near Morne Cavalier, Parc National
Pic Macaya, 1 km W of Cay Michel, wet limestone out-
crops, 1,191 m elevation. Department du Sud, Haiti,
18.3267° N, -74.0299° W.
Paratypes: UF 451537, Pic Formon, off path between
Morne Cavalier and Cay Michel, Parc National Pic
Macaya, wet limestone outcrops, 1,202 m elevation.
Department du Sud, Haiti, 18.3254° N, -74.0262° W
(1 specimen); OSUM 37269, Citadelle de Platons, cleared
area around fortress, 727 m elevation. Department du Sud,
Haiti, 18.2702° N, -73.9709° W (1 specimen); BMSM
17936, near Morne Cavalier, 1 km W of Cay Michel, wet
limestone outcrops, 1,191 m elevation. Department du
Sud, Haiti, 18.3267° N, —74.0299° W (1 specimen);
OSUM 37270, near Morn Cavalier, Parc National Pic
Macaya, foot of Mt. Ouasac, wet limestone outcrops in
cleared forest. Department du Sud, Haiti, 1,341 m eleva-
tion, 18.3391° N, -74.0304° W (1 specimen).
Other Material Examined: Grego coll., Pic Formon,
off path between Morne Cavalier and Cay Michel, Parc
National Pic Macaya, wet limestone outcrops, 1,202 m
elevation, 18.3254° N, —74.0262° W (8 specimens);
Grego coll., near Morne Cavalier, Parc National Pie
Macaya, 1 km W of Cay Michel, wet limestone outcrops,
1,191 m elevation, Department du Sud, Haiti, 18.3267°
N, —74.0299° W (4 specimens); Grego coll., near Morne
Cavalier, sinkholes in wet limestone forest, 1,116 m ele-
vation, 18.3227° N, —74.0204° W (10 specimens); Grego
coll., GTW 15157a, near Morn Cavalier, foot of Mt.
Ouasac, wet limestone outcrops in cleared forest, 1,341 m
elevation, 18.3391° N, —74.0304° W (4 specimens).
Distribution: Known only from the vicinity of Citadelle
de Platons and Morn Cavalier at 727-1,341 m elevation.
Variation in specimens: Thirty-one specimens seen.
The examples we have seen are very uniform in their
characteristics, differing only in the number of axial
lamellae (70-90).
Comparison with Other Species: See under W.
smithianum, above.
Etymology: L. mcicayaense, from the area of Parc
National Pic Macaya.
DISCUSSION
We have only Pfeiffer’s unfigured description of
Choanopoma? smithianum on which to base this species.
Only four species, which we wall refer to here as Species
A, B, C, and D, were found at the type locality and
surrounding areas (27 sites) despite careful examination.
G. Thomas Watters et ah, 2013
Page 83
All are members of Weinlandiponm . Pfeiffer’s descrip-
tion, taken as a whole, contains enough information to
identify which of these species represents C. smithianum
based on circumstantial evidence. We present all avail-
able evidence below.
(1) Pfeiffers description mentions his specimen being
11.5 mm long by 6 mm wide giving a lengtli/width
ratio = 1.9. Species A has a ratio of 1. 7-1.9. Species
B is more turbinate with a ratio of 1 .2. Species C has
a ratio of 1 .4—1.7. Species D is 1.6-1. 9. The length/
width ratio of Pfeiffer’s specimen most closely matches
Species A and D.
(2) Pfeiffer’s description mentions his specimen being
11.5 mm long in decollate length. Our specimens
of Species A vary from 9.0-11.0 mm in decollate
length. Species B varies from 8.8-11.3 mm in decol-
late length. Species C is the smallest with specimens
ranging from 6. 2-7. 2 mm in decollate length.
Species D is 7. 2-9. 6 mm in decollate length. The
decollate length of Pfeiffer’s specimen most closely
matches Species A and B.
(3) Pfeiffer (1866: 88) referred to “ .sutura costis
excurrentibus irregulariter crenata ” in his original
description. Species B has a set of cusps lining the
suture of a uniform height, but often alternating with
a smaller thread. Species A, C, and D have an undu-
lating row of cusps of various lengths. Unfortunately,
all four species could fit Pfeiffer’s description.
(4) Pfeiffer compared his species with Cyclostoma
serraticosta Weinland, 1862 and Choanoponia
puertoplaten.se Pfeiffer, 1858. The former species was
regarded as Orcuttipoma rollei serraticosta by Bartseh
(1946), and as Parachondria rollei serraticostus by
Watters (2006). The latter species was overlooked by
Bartseh (1946) and Watters (2006) suggested it might
be a Colonina based on the apparent spiral cords in the
original minute ( 10 mm) illustration. (However, in view
of Pfeiffer’s comparison it is more likely that the spiral
lines are bands of color rather than sculpture; this
would eliminate this species from Colonina but the
generic placement remains unknown as the type has
not been located.) Pfeiffer also included where he
would have placed his species in the 1865 second
supplement to his Monographia pneumonoponiorum
viventium-, he placed it at “7 1/2,” in between Cistula
tractum “Gundlaeh” (species 7) and Choanopoma
puertoplaten.se (species 8). Both of these species
resemble Species A, C, and D more so than Species B
in having a prominent color pattern; however the same
color pattern does occur in Species B but it is not as
well-defined. Again, Pfeiffer’s description of “seriebus
punctorum ruforum ornata” could apply to any of the
four species.
(5) Pfeiffer makes no note of the strongly scalloped axial
cords seen in Species C and D.
(6) The outer lip is narrowly adnate in Pfeiffer’s descrip-
tion of C. smithianum. Only Species A and B have
adnate outer lips.
(7) The peculiar folded lamella of the peristome found
in Species B, particularly in the Citadelle de Platons
specimens, not seen in any other species of the
genus, would certainly have been noted by Pfeiffer.
(8) Species D has not been found at the type locality.
Based on sculptural aspects, size, and degree of elonga-
tion we believe only Species A could fit the description
of Choanopoma ? smithianum Pfeiffer, 1866. Species B
is W g onavense (Weinland, 1880), species C is W
auduboni and D is W macayaense.
Weinlandiponm is restricted to the lower and middle
elevations of the Chaine de la Selle mountain range, which
forms the Tiburon Peninsula and continues into the
Barahona Peninsula as the Sierra de Baorueo (Figure 1 9).
It is also found on the neighboring islands of He tie la
Gonave, Petite Gonave, and lie a Vaehe. Species occur
from Polo in the eastern Barahona Peninsula and Port-
au-Prince across the Tiburon Peninsula to its western
end. This mountain range originated apart from the rest
of Hispaniola and continues to collide with it (with
recently disastrous results). The mountain range harbors
many unique species and genera of Annulariidae that
do not seem to have counterparts in the remainder of
Hispaniola. Watters (2006) hypothesized that these taxa
evolved on what is now the Tiburon Peninsula before it
merged with Hispaniola and that its taxa are not related
to those on tire rest of the island.
Tl le radula of Weinlandiponm is typical of most
annulariids, having a digitate first marginal (Figure 18).
It is this tooth that varies most in the family (previously
unpublished). Genetic studies of the Annulariidae are
now being completed.
Weinlandiponm smithianum , W macayaense , and W.
auduboni may co-occur with W. gonavense , although W.
gonavense appears to be more widespread. Weinlandiponm
smithianum has been found in the vicinity of Camp
Perrin at 200-210 m elevation (UF 32269, 48134),
Citadelle de Platons at 727 m elevation, and at Pic
Formon at 1,202 m elevation; it appears to occur at lower
elevations than W. auduboni and W. macayaense.
Weinlandiponm auduboni and W. macayaense are found
near Morn Cavalier and Citadelle de Platons between
727-1400 m elevation. These sites have wet limestone
outcrops and sinkholes in pine forests on the south slope
of the Pic Macaya range (although much of this area has
been deforested). Most of the sites are located within the
Parc National Pic Macaya, which includes the last virgin
cloud forest in Haiti and has as an extremely high level of
endemism of plants and animals (Thompson, 1986;
Grego and Steffek, 2007; Fernandez, 2007). Weinlandiponm
macayaense and W auduboni are sometimes, but not
always, found together. The condition of the specimens
does not permit a morphological examination of the geni-
talia but it is unlikely that the two species represent sexu-
ally dimorphic forms of one species; sexual dimorphism in
shells is unknown in Hispaniolan annulariids.
Weinlandiponm gonavense is characterized by a pecu-
liar apertural feature that is lacking in all other members
Page 84
THE NAUTILUS, Vol. 127, No. 2
of the genus (Figures 13-17). Several of the lamella
composing the outer lip are erect and folded back over
toward the aperture to various degrees, sometimes
forming a shelf. These folded lamella are usually among
the middle lamella (although in some populations the
lamella are outermost) and usually arise from the inner
(axial) portion of the lip. The degree of folding varies
considerably between and among populations, rarely
absent, but is prominent in the Citadelle de Platons
specimens. Specimens also vary in size, height, and color.
This variability caused Grego and Steffek (2007) to sug-
gest that additional species occurred at their study sites.
Thompson (1986) referred these taxa to Weinlandipoma
sp. Bartseh (1946) divided this taxon into four subspe-
cies, including one he did not formally name. These
subspecies ranged from lie de la Gonave to the south-
western coast of the Tiburon Peninsula. There appear to
be two distinct forms in our study areas, one tall and pale
that conforms to Bartseh’s subspecies conception , the
other short and dark and unnamed; some sites have both
forms, others only one. Both forms have folded aperture
lamella. However, a few specimens appear to be inter-
mediate between the two groups. At this time we are
referring to these specimens as W. pan accuse s.l. although
clearly more work is needed. A living individual of W.
gonavense was figured in Fernandez (2007, p. 195).
We believe the designation of a neotype of Choanopoma ?
smithianum Pfeiffer, 1866, for the purpose of clarify-
ing its taxonomic status is warranted at this time. The
species seems to have a much narrower range than
Weinlandipoma gonavense , with which it is occasionally
found and potentially confused. It may be a significantly
rare taxon deserving conservation protection that has
not been recognized because of the lack of illustration
or type specimens. Portions of its range have already
been deforested. A neotype is chosen from the type
locality and deposited at the Natural History Museum,
London, UK as NHMUK 20120258. The type locality of
“in monte Platon insulae Haiti ” is here amplified as
Citadelle de Platons, cleared area around fortress, 727
m elevation. Department du Sud, Haiti, 18.2702° N,
-73.9709° W.
ACKNOWLEDGMENTS
We greatly appreciate the support of Philippe Bayard and
Jean-Vilmond Hilllaire from the Soeiete du Audubon
Haiti and Eladio Fernandez from Sociedad Omitologica
Hispaniola for their excellent organization of the 2006
field trip to Parc National Pic Macaya. We thank Fred
Thompson, John Slapcinsky, and Gustav Paulay (UF) for
the use of their collections, and Kathie Way (NMHUK)
and Jose H. Leal (BMSM) for supplying catalogue
numbers. The manuscript was gready improved by two
anonymous reviewers.
LITERATURE CITED
Bartseh, P. 1946. The operculate land mcllusks of the family
Annulariidae of the island of Hispaniola and the Bahama
Archipelago. Bulletin of the U.S. National Museum 192:
264 pp., 38 pis.
Clench, W.J. 1935. Some new Urocoptidae from Hispaniola.
Proceedings of the Boston Society of Natural History 41:
1-12; pis. 1, 2.
Crosse, H. 1891. Faune malacologique terrestre et fluviatile de
file de Saint-Domingue. Journal de Conehyliologie 39:
73-210.
Fernandez, E. 2007. Hispaniola, a photographic journey
through island biodiversity. Belknap Press, Cambridge,
374 pp.
Grego, |. and J. Steffek. 2007. Biodiversity of Macaya National
Park and Biosphere Reserve in conditions of global change.
Pp. 8.5-112, in Proceedings from the Conference Stara
Lesna Tatry Biosphere Reserve, Slovakia, 2-6 June 2007.
Henderson, J.B. and P. Bartseh. 1920. A classification of
the American operculate land mollusks of the family
Annulariidae. Proceedings of die U.S. National Museum 58:
49-82.
Kohelt, W. 1880 Die geographische Verhreitung der Mollusken.
Jahrbiicher der Deutschen Malakozoologischen Gesellschaft
nebst Nachrichtsblatt 7(3): 241-286.
Pfeiffer, L. 1865. Monographia pneumonopomorum viventium,
Supplementum Secundum. Theodor Fischer, Cassellis, 284 pp.
Pfeiffer, L. 1866. Beschreibung neuer Landschnecken.
Malakozoologisehe Blatter 13: 76-91.
Pfeiffer, L. 1876. Monographia pneumonopomorum viventium,
accedente fossilium enumeratione. Supplementum tertium,
monograpliiae auriculaceomm. Parte seeunda auctum.
T. Fischer, Cassel, x + 479 pp.
Thompson, F.G. 1986. Land mollusks of the proposed National
Parks of Haiti. Prepared for SAID/ 1 laiti, Contract Number
521-01 69-C-00-3083-00 , 19 pp.
Tryon, G.W. 1867. Notices and reviews of new works.
American Journal of Conchology 3(1): 82-103.
Watters, G.T. 2006. The Caribbean land snail family
Annulariidae. Backhuys Publishers, Leiden. 557 pp.
THE NAUTILUS 127(2):85-89, 2013
Page 85
Mario7iia kinoi (Nudibranchia: Tritoniidae): A new species
from die tropical eastern Pacific
Orso Angulo-Campillo
Centro de Investigaciones Biologicas del Noroeste (CIBNOR)
Programa de Eeologia Pesquera, Instituto Politecnico Nacional
C.P 23096, La Paz, Baja California Sur, MEXICO
[email protected]
Hans Bertsch 1
Instituto de Investigaciones Oceanologicas
Universidad Autonoma de Baja California
Ensenada, MEXICO
and
Departamento de Ingeneria en Pesquerias
Universidad Autonoma de Baja California Sur
La Paz, MEXICO
[email protected]
ABSTRACT
Marionia kinoi , a new shallow water subtidal species of tritoniid
nudibranch, is described from the tropical eastern Pacific.
Along with a suite of internal characteristics (the number
of rodlet rows on the jaw and a distinctively shaped rachidian
tooth), its brilliant orange mid-dorsal color pattern, with whitish
margins, immediately distinguish this new species from its
Indo- and eastern Pacific congeners.
Additional Keywords: Gulf of California, Dendronotacea,
Eusebio Francisco Kino
INTRODUCTION
The most recent comprehensive review of Tritoniidae
taxonomy was published nearly 50 years ago (Odlmer,
1963). Since then, new species descriptions have called
into question his system of classification (e.g., Willan,
1988, and Smith and Gosliner, 2003, 2005, anti 2007).
When Bertsch et al. (2009) described the first species
of Tritoniidae known to feed on zoanthid (not aleyonarid)
Anthozoa, they presented a preliminary morphological
phylogenetic analysis of the family. Their strict consensus
phylogeny resulted in an unresolved polytomy of nine spe-
cies, distributed among four genera ( Marionia , Tritonia,
Tritoniella, and Tochuina), emphasizing the need for fur-
ther research to understand the evolutionary relationships
within this group.
The present paper contributes to our understanding
of the biodiversity of the tritoniids, essential for future
comparative studies of this elade. There are less than
30 currently accepted species of named Marionia world-
1 Mailing address: 192 Imperial Beach Blvd. #A, Imperial
Beach, CA 91932, USA
wide, with most occurring in the Indo-Paeific (Smith and
Gosliner, 2007; Gosliner, Behrens and Valdes, 2008; Garcia
and Bertsch, 2009). Adding to the many unresolved taxo-
nomic problems in this group, there are at least another
15 unnamed Indo-Paeific species (illustrated in Gosliner,
Behrens and Valdes, 2008) and 2 unnamed eastern Pacific
species, Marionia sp. 1 and M sp. 2 (illustrated in Behrens
and Hermosillo, 2005). Herein we name this Marionia
sp. 1 from tropical west America.
MATERIALS AND METHODS
The specimens were relaxed in iced water until no
response to external stimuli, then fixed in 4% Formalin.
Dissection was performed by a dorsal incision through the
length of the notum, from the base of the rhinopliores
to the base of the foot, allowing the removal of the entire
visceral mass in one piece. The jaws and the radula were
freed from the buccal mass, and put into 10% KOH solu-
tion. They were the rinsed in deionized water, air dried
and coated for electron microscopy. Scanning electron
micrographs (SEMs) were produced on a Hitaelu model
S-300D. Drawings were produced with the aid of a draw-
ing tube attached to a stereo microscope. All drawings
were digitalized by scanning and then composed and
edited for publication using Corel Photoshop rM. The
holotype and para type specimens are deposited in the
collections of California Academy of Sciences (CASIZ),
San Francisco.
SYSTEMATICS
Suborder Dendronotacea Odlmer, 1934
Family Tritoniidae Lamarck, 1809
Genus Marionia Vayssiere, 1877
Marionia kinoi new species
(Figures 1-10)
Page 86 THE NAUTILUS, Vol. 127, No. 2
Manama sp. I Behrens and Hermosillo, 2005*; Hermosillo
and Behrens, 2005; Hermosillo, 2006; Hermosillo
et ah, 2006*; Bertsch, 2010a.
Manama sp. Camacho-Garcia, et ah, 2005*.
* Indicates color photographs of living organ-
isms included
Description: External Anatomy (Figures 1-2): Lengths
of the living specimens were approximately 80mm. The
color of the animal is light orange to brownish red, with
small light colored tubercles that resemble a reticule that
covers the surface of the dorsum. The sheaths and shafts
of the rhinopl lores are orange, with tire apical portions a
light brown color. The body is elongate, subquadrilateral,
with the largest section 1/3 of the distance to die posterior
end of the foot. The oral veil extends beyond the front
of the body. There are eight velar papillae on each side on
the veil. The velar papillae are arranged in multifid groups,
with blunt rounded apices. The body is finely granulated,
except for the smooth sole of tire foot. The rhinophores
overhang from the margin of the notum. The rhinophore
shafts are typically tritoniid, with a central clavus sur-
rounded by a series of pinnate projections. The branchial
plumes are relatively short. There tire 11 plumes per side,
with the largest in the middle third of the body. The
branchial plumes are divided into four or five branches.
The gonopore is located on die right side, at die first third
of the body, below and between die 2Iul and 3rd branchial
plume. The anus is situated below die 3rd branchial plume,
at the same height as the gonopore (Figure 2).
Digestive System: The jaws are yellowish brown in
color, with a darker, reddish brown masticatory margin.
The masticatory margin appears smooth to the naked eye,
but scanning electron microscopy reveals the presence
of three to four rows of jaw rodlets. The jaw rodlets
at the anterior portion present three rows, which continue
halfway, were the fourth row begins to project (Figure 3),
until the posterior portion where the four rows are sepa-
rated. The jaw rodlets are slightly conical, resembling a
shark jaw (Figure 4). The radula of the paratype is large,
with a formula of 45 (35.1.1.1.35) at its broadest point
(Figures 5-7). The rachidian tootii is tricuspid, with a
base roughly rectangular, and resembles the shape of an
Figure 1. Mariania kinoi new species, photo of living speci-
men at type locality, 16 meters depth.
Figure 2. Lateral view of preserved specimen of Marionia
kinoi new species. Abbreviations: an, anus; gp, gonopore.
“M” (Figures 6-7). The central cusp is thick, asymmetri-
cal, incised in the shape of a “V,” with a profound canal
that begins at die base and fades to the right side of the
cusp. The top of the wedge forms two rectangular
projecting ridges that present scars along their interior
margins. The rows of teeth are arranged very closely
together. The first lateral teetii insert at the base of the
outer cusp of the rachidian tooth, and are differentiated
from the remaining lateral teeth, being more heavily
constructed, broader and curved. The remaining lateral
teeth are straight to slightly curved, and relatively broad
and stout. (Figures 5-6). The esophagus is half the size
of the digestive gland, forming a broad straight tube slighdy
distended at die insertion of die stomach on the ventral
side (Figure 8). The stomach is small, less than 0.5 cm
in diameter. There are approximately 30-38 subquadrate
(slightly more rectangular than square) stomach plates.
They are light cream colored, widi rounded comers and a
smooth surface. The intestine exits die stomach from the
anterior left portion, curving up around die anterior por-
tion of the digestive gland towards the right, becoming
wider for a portion before abruptly narrowing again to
terminate in die anus. The large digestive gland is shaped
like a chili pepper. The posterior portion has an anterior
hollow that fits around die posterior stomach. The smaller
anterior lobe covers the anterior portion of the stomach,
and is connected to the posterior digestive gland.
Reproductive System: A narrow hermaphroditic duct
connects to the narrow end of the ampulla (Figure 9).
The ampulla is muscular and convoluted, wide and
encased by the compact female gland mass. The smaller
albumen gland is easily discernible from the membrane
and mucus glands. The proximal vas deferens emerges
from the female gland mass as a thin, curved tube. The
distal portion slightly thickens before entering the base
of the conical unarmed penis, lying near the gonopore.
The muscular bursa copulatrix is an inflated oval 1 .5 cm
long. The vaginal duct is short, slightly widening at the
vaginal atrium. The oviduct exits the female gland mass
and opens to the vaginal atrium.
Nervous System: The ganglia of the central nervous
system sits on the dorsal esophagus, just behind the buccal
mass (Figure 10). The central nervous system is symetrical.
O. Angulo-Campillo and H. Bertsch, 2013
Page 87
Figures 3-7. SEMs oi jaw and radula of Marionia kinoi new species, paratype CASIZ 190511 . 3. Lateral view of the masticatory
border, showing the arrangement of jaw rodlets. 4. Detail of jaw rodlets. 5. Eleven rows of die radula, entire half-rows. 6. Closer view
of rachidian and lateral teeth (five rows). 7. Close-up of single rachidian tooth
Page 88
THE NAUTILUS, Vol. 127, No. 2
8 9
Figures 8-10. Drawings of internal anatomy of Marionia kinoi new species 8. Visceral mass. 9. Reproductive system. 10. Central
nervous system. Abbreviations: al, albumen gland; am, ampulla; an, anus; be, bursa eopulatrix; dga, anterior digestive gland; dgp,
posterior digestive gland; esc!, distal esophagus; esp, proximal esophagus; fgm, female gland mass; gp, gonopore; hd, hermaphro-
ditic duct; in, intestine; st, stomach; od, oviduct; pn, penis; va, vaginal atrium; vd, distal portion of vas deferens; vdp, proximal
portion of vas deferens.
The paired cerebral and pleural ganglia are distinct, joined
by a short connective. The pedal ganglia are below and on
either side of the cerebropleural ganglia, joined by short
thick connectives.
Type Material: Holotype: 3.6 cm in length, CASIZ
190509; Paratypes: 3.8 cm length (dissected, CASIZ
190511 ) and 4 cm in length (CASIZ 190510), all from type
locality, 9 March, 201 1, Orso Angulo and Jorge Zarate coll.
Type Locality: Two miles south of Punta Arenas, on tire
Gulf of California coastline east of La Paz, near La Riviera,
Baja California Sur, Mexico (23°29'24" N; 109°27'08" W),
16 m depth.
Etymology: The specific name kinoi is given in honor
of Padre Eusebio Francisco Kino, S.J. (1645-1711), the
first Pacific conchologist (Baily, 1935), and intrepid
explorer and cartographer of the Californias. His knowl-
edge of the distribution of abalones was crucial for his
determination that Lower California was a peninsula, not
an island (Bertseh, 2010b). He founded the first Jesuit
Catholic mission in the Baja California peninsula at La
Paz, which however only functioned from 2 April to
14 July 1683 (Bertseh, 201 1); no ruins are known to exist.
Distribution: Marionia kinoi new species is known from
three eastern Pacific faunal provinces: the Sea of Cortez,
Mexican and Panamic (. sensu Briggs, 1974). It has been
reported from the extreme southern portion of tire Gulf
of California (herein), from Bahia de Banderas, Jalisco/
Nayarit (Hermosillo, 2006), Faro de Bucerias, Michoacan,
and Ixtapa, Guerrero (Hermosillo and Behrens, 2005),
Costa Rica (Behrens and Hermosillo, 2005), and the Islas
Galapagos (Camaeho-Garcxa et al., 2005).
DISCUSSION
Marionia kinoi new species has been included in the
genus Marionia because of the number of rows in the
jaw (3 to 4) and the presence of stomach plates, in accor-
dance with the division proposed by Odhner (1963).
Currently, the genus consists of 23 named Indo-Pacific
species (compared and illustrated in Smith and Gosliner,
2007, and Gosliner, Behrens and Valdes, 2008), and five
named species in the Atlantic (listed in Garcia and
Bertseh, 2009).
This new species can be distinguished readily from other
species of Marionia based on coloration. For instance,
Marionia cyanobranchiata (Riippell and Leuckart, 1828),
and M. platyctenea (Willan, 1988) have dark green brown
or black gills. Marionia di.stincta Bergh, 1905, M. levis
Eliot, 1904, M. elongoreticulata Smith and Gosliner, 2007,
turd M. elongoviridis Smith and Gosliner, 2007, have trans-
verse light or dark lines on tire dorsum. Among Atlantic
species, the West African deeper-water species M. vanira
Marcus and Marcus, 1966 has a green body, and tire
Caribbean M. tedi (Marcus, 1983) has a translucent white
body and wlrite processes.
Gills situated on a more prominently-thickened basal
trunk separate Marionia kinoi new species from M. rubra
(Riippell and Leuckart, 1828), M. dakini (O’Donoghue,
1924), M. arborescens Bergh, 1890, and M. pustulosa
Odhner, 1936.
O. Angulo-Campillo and H. Bertsch, 2013
Page 89
The known feeding specificity of tritoniids on a single
species or small group within a family of Octoeorallia
(Smith and Gosliner, 2003) reliably distinguishes M. kinoi
new species from M. hawaiiensis (Pease, I860), which
preys on Anthelia , and M. bathycarolinensis Scott
and Gosliner, 2005, a specific predator on Paracis.
Neither of these prey genera of Octoeorallia are known
to occur in the Gulf of California (Hendriekx, Brusca,
and Findley, 2005).
In addition to these features, there are two external
characters drat differentiate the plate-bearing tritoniids
from our new species: the number of branchial plumes
and the number of velar papillae. Marionia kinoi new
species has 11 pairs of branchial plumes and eight pairs
of compound velar papillae. None of the other 20 Pacific
and two Atlantic species (contrasted in Table 1, Smith and
Gosliner, 2007) has this suite of features. Moreover, the
presence of 3 to 4 rows of jaw rodlets and 35 teeth rows
distinguish M kinoi new species from these congeners.
This new species differs from the only other known
Marionia species occurring in the American Pacific; that
still undescribed species is smaller in size, and the body,
lateral processes, oral veil and rhinophores are white.
ACKNOWLEDGMENTS
We thank Jorge Zarate for scuba diving assistance in
collecting the specimens of this new species.
LITERATURE CITED
Baily, Jr., J.L. 1935. The first Pacific conchologist. The Nautilus
48: 73-75.
Behrens, D.W. and A. Hermosillo. 2005. Eastern Pacific Nudi-
branchs. A Guide to the Opisthobranchs from Alaska to
Central America. Sea Challengers, Monterey, 137 pp.
Bertsch, H. 2010a. Las conchas azules (the blue shells): Father
Kino, abalones, and the island of California. The Nautilus
124: 188-191
Bertsch, H. 2010b. Biogeography of northeast Pacific opistho-
branchs: Comparative faunal province studies between
Point Conception, California, USA, and Punta Aguja, Piura,
Peru. In: Rangel Ruiz, L.J., J. Gamboa Aguilar, S.L. Arriaga
Weiss, and W.M. Contreras Sanchez (compliadores).
Perspectivas en Malacologia Mexicana. Universidad Juarez
Autonoma de Tabasco, Villahermosa: 219-259.
Bertsch, H. 2011. The missionary explorer-scientist. Father
Eusebio Kino, S.J.: Faith, maps, sea shells and Mission
San Xavier del Bac. Catholic Southwest, A Journal of His-
tory and Culture 22: 68-85.
Bertsch, H., A. Valdes, and T.M. Gosliner. 2009. A new species
of tritoniid nudibranch, the first found feeding on a
zoanthid anthozoan, with a preliminary phylogeny of the
Tritoniidae. Proceedings of the California Academy of
Sciences 60: 431-446.
Briggs, J.C. 1974. Marine Zoogeography. McGraw-Hill, New
York, 475 pp.
Camacho-Garcia, Y., T.M. Gosliner, and A. Valdes. 2005. Guia
de Campo de las Babosas Marinas del Pacifico Este
Tropical. California Academy of Sciences, San Francisco,
129 pp.
Garcia, F.G. and H. Bertsch. 2009. Diversity and distribution
of the Gastropoda Opisthobranchia from the Atlantic
Ocean: A global biogeographic approach. Scientia Marina
73: 153-160.
Gosliner, T. M., D.W. Behrens, and A. Valdes. 2008. Indo-Pacific
nudibranchs and sea slugs: A guide to the world's most
diverse fauna. Sea Challengers Natural History Books,
Etc., Gig Harbor, Washington, and California Academy
of Sciences, San Francisco, California. 426 pp.
Hendriekx, M E., R.C. Brusca, and L.T. Findley. 2005. Listado
y distribution tie la macrofauna del Golfo tie California,
Mexico. Parte 1. Invertebrados. A distributional checklist
of die macrofauna of the Gulf of California. Part 1 Inverte-
brates. [Bilingual] Arizona-Sonora Desert Museum, Tucson,
Arizona, xi + 429 pp.
Hermosillo, A. 2006. Ecologia de los opistobranquios (Mollusca)
de Bahia de Banderas, Jalisco-Nayarit, Mexico. Ph D. Tesis,
CUCBA, Universidad tie Guadalajara, 151 pp.
Hermosillo, A. and D.W. Behrens. 2005. The opisthobranch
fauna (Gastropoda, Opisdiobranchia) of the Mexican states
of Colima, Michoacan anti Guerrero: Filling in the faunal
gap. Vita Malacologiea 3: 11-22.
Hermosillo, A., D.W. Behrens, and E. Rios-Jara. 2006.
Opistobranquios de Mexico. Guia de babosas marinas del
Pacifico, Golfo de California y las islas ocean icas. Comision
National para el Conocimiento y Uso de la Biotliversidad
(CONABIO), Mexico, 143 pp.
Odhner, N. 1963. On the taxonomy of die family Tritoniidae.
The Veliger 6: 48-52.
Smith, V. G. and T.M. Gosliner. 2003. A new species of Tritonia
from Okinawa (Mollusca: Nudibranchia), and its asso-
ciation with a gorgonian octocoral. Proceedings of the
California Academy of Sciences 54: 255-278.
Smidi, V.G. and T.M. Gosliner. 2005. A new species of Marionia
(Gastropoda: Nudibranchia) from the Caroline Islands.
Proceedings of the California Academy of Sciences 56:
66-75.
Smith, V.G. and T.M. Gosliner. 2007. Two new species of
Marionia (Mollusca: Nudibranchia) from the Indo-Pacific
Region. The Veliger 48: 260-275.
Willan, R.C. 1988. The taxonomy of two host-specific, cryptic
dendronotoid nudibranch species (Mollusca: Gastropoda)
from Australia, including a new species description. Zoo-
logical Journal of the Linnean Society 94: 39-63.
Omphalotropis ilapiryensis, a replacement
name for O. costulata Emberton and
Pearce, 1999 (Gastropoda: Littorinimorpha:
Assimineidae)
Omphalotropis costulata Emberton and Pearce, 1999,
from Madagascar, is a caenogastropod (operculate) land
snail of the family Assimineidae. The genus is known from
the central Pacific westward through the Indian Ocean,
with Madagascar located at the extreme western edge of
its distribution. Sixteen species of Omphalotropis are
known from Madagascar (Pearce, 2003; Emberton 2004,
2009). Omphalotropis costulata Emberton and Pearce,
1999 has been reported from only two localities in a narrow
geographical range where it is endemic to a single moun-
tain range in southeastern Madagascar. The true ranges
and conservation statuses of Madagascar’s small, opercu-
late land snails are very poorly known (Emberton, 2009)
but the ongoing rapid destruction of rainforests in
Madagascar poses serious conservation concerns to this
and other species in Madagascar (Pearce, 2003).
We learned recently that the species name is a junior
homonym of Omphalotropis costulata (Mousson, 1870),
which occurs in Fiji, on Vanua Mbalavu Island. As per
Art. 52.2 of the ICZN (1999), the junior homonym
Omphalotropis costulata Emberton and Pearce, 1999
is an invalid name. No other synonym exists for this
Malagasy species, so a replacement name is required
(ICZN Art. 60.3). The species is currently listed in the
IUCN Red List (Seddon, 2012) as Omphalotropis sp.
nov. 2, a reversion to its original unassigned description
in the literature (Emberton et ah, 1996; Emberton, 1997).
Here we assign it a replacement name.
Omphalotropis ilapiryensis new name
Omphalotropis 02 (as OM02) — Emberton et ah, 1996:
210; Emberton, 1997: 1147, Table 5.
Omphalotropis sp. 2 — Emberton, 1997: 1143 (and as
“OM02” on p. 1147); Seddon, 2012
Omphalotropis costulata Emberton and Pearce, 1999:
Emberton and Pearce, 1999: 370-371, figs. 25, 48;
Pearce, 2003: 541, Table 8.10; Emberton, 2004: 84,
88, figs. 36-37, Table 3; not Omphalotropis costulata
(Mousson, 1870: 190-191, pi. vii, fig. 10) [orig. Realia
( Omphalotropis) costulata].
Type Material: Omphalotropis costulata: Holotype
USNM 860791 (Emberton, 2004: 88); paratypes UF
(Florida Museum of Natural History) 316989, 316990,
318440, 318441, 318442; all type material is from
Mount Ilapiry.
Additional Material: Additional specimen UF
282795, cataloged at UF as from Ramabeafo Mountain
(Emberton, 2004: 88, figs. 36-37).
Etymology: For its type locality, Mount Ilapiry
(Vohimena Chain, Tulear Province, southeastern
Madagascar; Emberton and Pearce, 1999).
Remarks: Ramabeafo Mountain (24.742° S, 46.854° E)
is located 20.5 km NW from Mount Ilapiry (24.859° S,
47.011° E). Later, Emberton (2009: 20-21, figs. 119-121)
used "Omphalotropis sp. 2” for a species found south
of Antsahanoro in northeastern Madagascar, but it is
certainly not the same species as O. ilapiryensis (which
was formerly referred to as “ Omphalotropis sp. 2”).
ACKNOWLEDGMENTS
We acknowledge Carl Christensen, Mary Seddon, and
Craig Hilton-Taylor (IUCN) for bringing the hom-
onymy of Omphalotropis costulata to our attention.
We are grateful to John Slapcinsky for assistance,
including obtaining museum specimen records. We
thank Mary Seddon and John Slapcinsky for reviewing
the manuscript.
LITERATURE CITED
Emberton, K.C. 1997. Diversities and distributions of 80 land-
snail species in southeastern-most Madagascan rainforests,
with a report that lowlands are richer than highlands in
endemic and rare species. Biodiversity and Conservation
6(8): 1137-1154.
Emberton, K.C. 2004. Madagascan Georissa, Cijclotus ,
Omphalotropis and so-called Chondrocyclus (Gastropoda:
Caenogastropoda: Hydrocenidae, Cyelophoridae, Assimineidae).
Archiv fur Molluskenkunde 133(1-2): 69-107.
Emberton, K.C. 2009. Operculate land snails from three
rainforest transects in northeastern Madagascar, with
descriptions of nine new species and one new subspecies
(Gastropoda: Caenogastropoda). Archiv fur Molluskenkunde
138(1): 1-41.
Emberton, K.C. and T.A. Pearce. 1999. Land caenogastropods
of Mounts Mahermana, Ilapiry, and Vasiha, southeastern
Madagascar, with conservation statuses of 17 species of
Boucardicus. The Veliger 42(4): 338-372.
Emberton, K.C., T.A. Pearce, and R. Randalana. 1996.
Quantitatively sampling land-snail species richness in
Madagascan rainforests. Malacologia 38: 203-212.
ICZN [International Commission on Zoological Nomenclature].
1999. International Code of Zoological Nomenclature.
T. A. Pearce and M. E. Paustian, 2013
Page 9 1
Fourth Edition. The International Trust for Zoological
Nomenclature, London, xxix+ 306 pp.
Mousson, A. 1870. Faune malacologique terrestre et fluviatile
des iles Viti, d’apres les envois de M. le Dr. Edouard
Graeffe. Journal de Conchyliologie 18 (1870): 109-135,
part 2: 179-236, pis. VII-VIlT. [in French]
Pearce, T.A. 2003. Gastropoda, terrestrial snails. In: Goodman,
S.M. and J.P. Benstead (eds.). Natural History ol
Madagascar. University of Chicago Press, Chicago:
529-574.
Seddon, M B. 2012. Omphalotropis sp. nov. 2. IUCN 2012. IUCN
Red List of Threatened Species. Version 2012. 1 . http://www
.iuenredlist.org/details/40081/0. (Access 23 Oct. 2012).
Timothy A. Pearce
Megan E. Paustian
Section of Mollusks
Carnegie Museum of Natural History
Pittsburgh, PA, 15213 USA
[email protected]
Sponsored in part by the State of
Florida, Department of State,
Division of Cultural Affairs and the
Florida Council on Arts and Culture
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CONTENTS
Hiromi Watanabe from a hydrothermal vent in the Iheya Ridge in the
Takenori Sasaki mid-Okinawa Trough, Japan 93
Emily A. Glover A new shallow water species of Nucinella from the Philippines
John D. Taylor (Bivalvia: Protobranchia: Nucinellidae), member of a tropical
seagrass chemosynthetic community 101
Jeffrey C. Nekola Vertigo marciae (Gastropoda: Vertiginidae), a new land snail
Gary Rosenberg from Jamaica 107
Juan-Franeisco Araya Comisepta guzmani new species: first species of genus confirmed
Daniel L. Geiger from Pacific cold seep environments off central Chile (Gastropoda:
Vetigastropoda: Fissurellidae) 115
Russel L. Minton A new species of Lithasia (Gastropoda: Pleuroceridae) from
the Buffalo River, Tennessee, USA 119
Martin Avery Snyder Description of Fusilaria garcicii new genus, new species (Gastropoda:
Fasciolariidae: Fasciolariinae) from the western Caribbean Sea 125
THE NAUTILUS 127(3):93-100, 2013
Page 93
A new species of Solemya (Bivalvia: Protobranchia:
Solemyidae) from a hydrothermal vent in the Iheya Ridge
in the mid- Okinawa Trough, Japan
Kei Sato Hiromi Watanabe
Department of Earth and Planetary Science Japan Agency for Marine-Earth Science and Technology
The University of Tokyo 2-15, Natsushima
7-3-1, Hongo, Bunkyo-ku Yokosuka 237-0061, |APAN
Tokyo 1 13-0033, JAPAN
and
The University Museum
The University of Tokyo
7-3-1 Hongo, Bunkyo-ku
Tokyo 1 13-0033, JAPAN
Takenori Sasaki
The University Museum
The University of Tokyo
7-3-1 Hongo, Bunkyo-ku
Tokyo 1 13-0033, JAPAN
ABSTRACT
A new species, Solemya {Solemya) flam, is described from a
hydrothermal vent field located in the Iheya Ridge in the mid
Okinawa Trough, Japan. The small-sized new species is char-
acterized by (1) a branched internal ligament attached to the
ehondrophore, (2) a bright yellowish brown periostraeum, and
(3) relatively short shell length relative to height (length/
height = ca. 2.2) for solemyid bivalves. This species is associated
with a vesicomyid clam, Calyptogena okutanii Kojima and Olita,
1997, which is dominant in chemosynthetic environments.
Additional Keywords: taxonomy, Acharax, Calyptogena site
INTRODUCTION
Solemyids are an ancient group of bivalves whose fossil
records date back to the Ordovician (Pojeta, 1988; Bailey,
2011). More than 30 Recent species are known (Huber,
2010) , and there is also a rich fossil record for the group
(e.g. Kiel, 2010; Amano and Ando, 2011; Taviani et ah,
2011) . They are morphologically characterized by an
elongate shell with a thick periostraeum, a posteriorly
situated and toothless hinge, enlarged etenidia, and
reduced digestive system. The periostraeum is much
larger than the calcified part of the shell and folded
inwards in living-animals. All known species are asso-
ciated with reduced environments including vents and
seeps, and they are one of several groups of chemo-
synthetic bivalves (Taylor and Glover, 2010), harboring
sulfide-oxidizing ehemoautotrophie bacteria in the etenidia
(Stewart and Cavanaugh, 2006).
Numerous chemosynthetic-based biological commu-
nities have been documented around Japan since 1980s
(see review by Sasaki et al., 2005; Watanabe et al., 2010).
Okinawa Trough is one of major localities with com-
munities sustained by active hydrothermal vents. During
a dive in the Iheya Ridge in mid-Okinawa Trough, an
unknown species of the Solemyidae was collected. This
was the first record of the family from the Okinawa
Trough, and, as a result of morphological comparisons,
the specimens were identified as a new species. In this
paper we describe this new solemyid species colonizing
the Calyptogena site.
MATERIALS AND METHODS
Three specimens of the new species were collected
from 27°32.994' N, 126°58.230' E at a depth of 1402 m
(Figures 1-3) with a scoop sampler from ROV Hyper-
Dolphin operated by the Japan Agency for Marine-
Earth Science and Technology (JAMSTEC). The specimens
were collected together with sediments and Calyptogena
okutanii during Dive #1246, February 10, 2011. The
locality has been surveyed since the 1990s, and its fauna
and bottom characters were documented by Ohta and
Kim (2001). The specimens were preserved during the
Page 94 THE NAUTILUS, Vol. 127, No. 3
Figures 1-3. The type locality of Solemya (Solemya) flava new species. 1. Map of Japan showing location of Okinawa. 2. Detailed
map of Okinawa Trough; bottom contour intervals are 1000 m. 3. Detailed bathymetrical map of the Iheya Ridge. Star indicates
the type locality.
cruise at — 30°C in light-shielded conditions, and fixed
in 99% ethanol in the laboratory. All specimens used for
description were dissected under a binocular microscope.
The type specimens were registered in the Department of
the Historical Geology and Paleontology, The University
of Tokyo (UMUT).
SYSTEMATICS
Family Solemyidae Gray, 1840
Genus Solemya Lamarck, 1818
Solemya (Solemya) flava new species
(Figures 4-15)
Diagnosis: Branched internal ligament attached to the
chondrophore, bright yellowish brown periostracum, and
relatively short shell length relative to shell height for
solemyid bivalves.
Description: Shell small, up to 14.2 mm in length,
elongate oval, laterally compressed, equivalve, inequi-
lateral, thin and flexible. Umbones not prominent, situ-
ated at posterior one-fourth of shell length. Lunule and
escutcheon absent. Antero-dorsal margin straight, almost
parallel to straight ventral margin. Both anterior and
posterior margins gently rounded. Postero-dorsal margin
slightly convex, positioned slightly below beak, with well-
expressed radial, oblique flexure corresponding to attach-
ment of chondrophore to inner shell surface. Shell length
small relative to shell height (length/height = ca. 2.2) for
solemyid bivalves. Periostracum thick, brownish yellow,
extending beyond shell margins and forming frills. Sur-
face with widely spaced weak radial ribs or furrows, espe-
cially on anterior and posterior regions. Internal surface
dull white. Ligament internal, thin, subtriangular, moder-
ately broad, attached to chondrophore, without expansion
in front of chondrophore. Chondrophore weak, narrow,
situated behind posterior adductor scar. Internal radial
rib anterior to posterior adductor scar inconspicuous,
narrow, extended from umbo and located anterior to pos-
terior adductor scar. Anterior adductor scar large, angular-
oval, feebly impressed. Posterior scar smaller, oval, not
adjoining antero-ventral margin of chondrophore or
extending behind chondropore. Pallia] line obscure.
Type Material: Holotype, UMUT RM30147, 11.5 x
5.8 mm with periostracum, 10.7 x 4.6 mm excluding
periostracum (Figures 4-7); paratype #1, UMUT RM30148,
15.4 x 9.3 mm with periostracum, 14.2 x 6.9 mm
excluding periostracum, (Figures 8-11); paratype #2,
UMUT RM30149, 13.1 x 6.9 mm with periostracum,
12.5 x 5.7 mm excluding periostracum (Figures 12-15).
Type Locality: Iheya Ridge in the Okinawa Trough,
27°32.994' N, 126°58.230' E, 1402 m deep.
Distribution: Known only from the type locality.
K. Sato et al., 2013
Page 95
Figures 4-15. Solemya ( Solemya) flava . 4-7. Holotype UMUT RM30147. 8-11. Paratype #1 UMUT RM30148. 12-15. Paratype
#2 UMUT RM 30 149.
Page 96
THE NAUTILUS, Vol. 127, No. 3
Figures 16-24. Solemyid bivalves for comparison (see also Sato et al., 2013). 16-18. Solemya ( Solemya ) pusilla (Gould, 1861)
UMUT RM 31050, Okinoshima-Island, Chiba Prefecture, Japan. 19-21. Solemija ( Solemya ) tagiri Okutani, Hashimoto and Miura,
2004, one of paratypes JAMSTEC 031588-031594, Kagoshima Bay, Kagoshima Prefecture, Japan. 22-24. Solemya (. Petrasma )
pervemicosa (Kuroda, 1948) |AMSTEC 053256, Joetsu Knoll, Niigata Prefecture, Japan.
K. Sato et al., 2013
Page 97
Figures 25-31. Solemyid hi\'alves for comparison (see also Sato et al., 2013). 25-28. Acharax johnsoni ( Dali, 1981) UMUT RM
28724. Off Otsuchi, Iwate Pref., Japan. 29-31. Acharax japonica (Dunker, 1882) UMUT RM 30934. Shimoda Marine Research
Center, Shizuoka Pref., Japan.
Etymology: The specific epithet is derived from the
Latin word flavus meaning yellow.
DISCUSSION
The new species is assigned to Solemya ( Solemya ) on the
basis of conchological characters. The genus Solemya is
distinguished from another valid genus of the family,
Acharax in having internal ligament and a slightly convex
postero-dorsal margin (Dali, 190S). Solemya is divided
into five subgenera: Solemya , Solemyarina, Petrasma,
Zesolemya, and Austrosolemya (Dali, 1908; Cox, 1969;
Taylor et al., 2008; Kamenev, 2009) which are diagnosed
as follows by features near the umbo such as the posterior
adductor muscle scar, internal ligament, chondrophore
Page 98
THE NAUTILUS, Vol. 127, No. 3
Figures 32—43. Internal umbonal areas of six solemyid species around Japan. Images in Figures 32-37 are shown as diagrams in
Figures 38-43. 32, 38. Acharax japonica. 33, 39. Acharax johnsoni. 34, 40. Solemya ( Petrasma ) peroemicosa. 35, 41. Solemya
(Solemya) pusilla. 36, 42. Solemya ( Solemya ) tagiri. 37, 43. Solemya (Solemya) flava new species. Abbreviations: c, chondrophore;
li, ligament; pa, posterior sdductor scar; pie, posterior outer-layer ligamental extension; r, internal radial rib.
and internal radial ribs: (1) Solemija s.s. has an opisthodetie
internal ligament and a short and thin chondrophore;
(2) Solemyarina has a heart-shaped internal ligament
expansion in front of the week and narrow chondrophore.
The posterior adductor scar is not impressed into the
inner shell surface; (3) In Petrasma, the anterior end of
the posterior adductor muscle scar is deeply impressed
and attached to the anteroventral margin of the
chondrophore; (4) Z esolemija is characterized by narrow
posterior expansions of the ligament, a bifurcated
chondrophore around the posterior adductor muscle
scar, and prominent anterior linear extensions of the
ligament; (5) Austrosolemya possesses an internal liga-
ment with lobate anterior extensions, a broad triangular
resilium, and strong chondrophoral ridges.
At the species level, the new species from off Okinawa
should be compared with known solemyids from Japan
and other adjacent regions of the Pacific geographically.
Around Japan five species have been recorded (Kuroda
et ah, 1971; Habe, 1977; Okutani, 2000), namely
Solemya ( Solemya ) pusilla (Gould, 1861) (Figures 16-18);
S. (S.) tagiri Okutani, Hashimoto and Miura, 2004
(Figures 19-21); S. ( Petrasma ) pervernicosa (Kuroda,
1948) (Figures 22-24); Acharax johnsoni (Dali, 1981)
(Figures 25-28), and A. japonica (Dunker, 1882)
(Figures 29-31). Solemya (S.) flava is distinct from these
species in five characters, viz. maximum shell length,
ligament position, configuration of posterior adductor
muscle scar, length and thickness of chondrophore,
and internal radial ribs (Table 1). Species from other
regions of the North Pacific ( Solemya ( Solemya ) volvulus ,
Solemya ( Petrasma ) panamensis ) are also distinguished
by these characters (see also Table 1).
In Solemya ( Solemya ), two species, S. (S. ) pusilla and
S. (S. ) tagiri , seem closely related to the new species in
shell morphology. Solemya (S.) pusilla is most similar in
maximum size to this species but differs in having a
much more elongate outline (L/H = 2.72). Solemya
( Solemya ) tagiri is also similar in shell size and having a
narrow and weak internal radial rib. However, the new
species is distinguished from these species in having
short shell length (L/H = 2.18) relative to shell height
and umbo situation. The umbo of S (S. ) tagiri is situated
in a more anterior position than that of this species.
Internal umbonal areas of six solemyid species are illus-
trated in Figures 32—43 for comparison.
From an ecological standpoint, the new species
might be related to S. (S. ) tagiri , since both species
are dependent on chemosynthesis-based, hydrother-
mal vent communities and their habitats are relatively
geographically close in Japan. However, the habitat of
S. (S.) tagiri is restricted to a narrow site at depth of
94-98 m in Kagoshima Bay which is nearly closed and
isolated from the open ocean. In contrast, the locality
of S. (S. ) flava is isolated in the middle part of Okinawa
Trough at a much greater depth (1402 m). Therefore,
each species is possibly endemic to Kagoshima Bay and
Okinawa Trough, respectively, and their habitats are
sharply separated.
ACKNOWLEDGMENTS
We thank the crew of the R/V Natsushima and the oper-
ation team of the BOV Hyper-Dolphin for assisting with
sample collection. This study was supported by Grant-in-
Aid for Scientific Research B (23370040) from Japan
Society for the Promotion of Science.
LITERATURE CITED
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Solemyidae) from the Miocene of Japan. The Nautilus
125: 207-212.
Bailey, J.B. 2011. Paleobiology, paleoecology, and systematic^
of Solemyidae (Mollusca: Bivalvia: Protobranchia) from
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Coan, E.V., P.V. Scott, and F. R. Bernard. 2000. Bivalve Sea-
shells of Western North America. Santa Barbara Museum
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Cox, L. B. 1969. Superfamily Solemyacea. In: R.C. Moore (ed.).
Treatise on Invertebrate Paleontology, Part N, Mollusca 6
(Bivalvia), Vol. 1. Geological Society of America and Uni-
versity of Kansas, Boulder, pp. 241-243.
Dali, W. H. 1908. A revision of the Solenomyacidae. The
Nautilus 22: 1-2.
Habe, T. 1977. Systematic^ of Mollusca in Japan. Bivalvia and
Scaphopoda. Hokuryukan, Tokyo, 372 pp. [in Japanese]
Huber, M. 2010. Compendium of Bivalves, Conchbooks,
Haekenheim, 901 pp.
Kamenev, G.M. 2009. North Pacific species of die genus
Solemya Lamarck, 1818 (Bivalvia: Solemyidae), widi notes
on Acharaxjohnsoni (Dali, 1981). Malacologia 51: 233-261.
Kiel, S. 2010. The fossil record of the vent and seep mollusks. In:
Kiel, S. (ed.). The Vent and Seep Biota. Aspects from
Microbes to Ecosystems. Springer, Dordrecht, pp. 255-277.
Kuroda, T., T. Habe, and K. Oyama. 1971 The Sea Shells
of Sagami Bay. Maruzen, Tokyo, 489 pp. + 121 pis. +
489 pp. + 51 pp.
Ohta, S. and D. Kim. 2001. Submersible Observations of the
Hydrodiermal Vent Communities on the Iheva Ridge,
Mid Okinawa Trough, Japan. Journal of Oceanography
57: 663—677.
Okutani, T. 2000. Family Solemyidae. In: Okutani, T. (ed.),
Marine Molluscs in Japan. Tokai University Press, Tokyo,
pp. 832-833.
Okutani, T., J. Hashimoto, and T. Miura. 2004. A new species
of solemyid bivalves from near submarine fumaroles in
Kagoshima Bay, Japan. Venus 62: 91-96.
Pojeta, J., Jr. 1988. The origin and Paleozoic diversification of
solemyoid pelecypods. New Mexico Bureau of Mines &
Mineral Resources Memoir 44: 201-222.
Sasaki, T., T. Okutani, and K. Fujikura. 2005. Molluscs from
hydrodiermal vents and cold seeps in Japan: A review of
taxa recorded in twenty recent years (1984-2004). Venus
64: 87-133.
Sato, K., R. Nakashima, R. Majima, H. Watanabe, and T. Sasaki.
2013. Shell microstructures of five Recent solemyids from
Japan (Mollusca: Bivalvia). Paleontological Research 17:
69-90.
Stewart, F.J. and C.M. Cavanaugh. 2006. Bacterial endosym-
bioses in Solerm/a (Mollusca: Bivalvia) — Model systems
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Taviani, M., L. Angeletti, and A. Ceregato. 2011. Chemo-
synthetic bivalves of the family Solemyidae (Bivalvia,
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Taylor, J.D., E.A. Glover, and S.T. Williams. 2008. Ancient
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THE NAUTILUS 127(3): 101-1 06, 2013
Page 101
A new shallow water species of Nucinella from the Philippines
(Bivalvia: Protobranchia: Nucinellidae), member of a tropical
seagrass chemosynthetic community
Emily A. Glover
John D. Taylor
Department of Life Sciences
The Natural History Museum
London SW7 5BD, UNITED KINGDOM
[email protected]
[email protected]
ABSTRACT
A new species of the protobranch bivalve Nucinella is described
from Panglao, central Philippines. It lives in very shallow
seagrass habitats and co-occurs with other chemosymbiotic
lueinid bivalves Pillucina and Cardiolucina. The species has
been included in recent molecular phylogenies. For compari-
son details of hinge teeth are provided for the type species
of Nucinella, Pleurodon ovalis , from the Pliocene of England.
Additional Keywords, protobranch, chemosymbiosis, new species
INTRODUCTION
Nucinellidae is a family of minute, mainly deep-water
protobranch bivalves recently recognised as chemo-
symbiotic (Oliver and Taylor, 2012; Bieler et al., in
press). Unusually, they are also all nronomyarian, with
only the anterior adductor muscle present. There are 21
known species in two genera comprising 15 Nucinella
and 6 Huxleyia (La Perna, 2005; Oliver and Taylor,
2012). Identification of Nucinella is problematic on
account of their small, smooth, external shells and
species discrimination is largely based on hinge den-
tition and ligament form. Most are <5nrm in height,
with ovoid, smooth shells, often with a blue-green
periostraeum; tbe largest living species (25 mm) is
Nucinella boucheti La Perna, 2005 described from
1600 m in the Philippines (La Perna, 2005). The liga-
ment is external, either opisthodetic or amphidetic
in Nucinella and internal opisthodetic in Huxleyia.
Tl le dentition is taxodont with up to 13 teeth located
beneath the umbones and a larger and generally elon-
gate, single anterior lateral tooth. There are persistent
reports that Huxleyia species are dimyarian (e.g. Coan
and Scott, 2012) but, after careful examination of sev-
eral species, Oliver and Taylor (2012) were unable to
confirm this and considered the genus monomyarian, as
did Hayami and Kase (1993). Anatomically, nucinellids
have large protobranch ctenidia, a large cleft foot that
is deeply digitate at the margins and small, finger-like
labial palps (Allen and Sanders, 1969; Kuznetsov and
Schilyeko, 1984; Oliver and Taylor, 2012).
Tbe depth distributions of living nucinellids are
known to range from the interidal zone to 3580 m
(Matsukuma et al., 1982; La Perna 2005; Oliver and
Taylor, 2012) although the majority of records are from
offshore to bathyal depths with N. owenensis Oliver and
Taylor, 2012 found at 3400 m off Oman. Only a few
species are known from very shallow water; Huxleyia
diabolica (Jousseaume, 1897) is recorded from 6-40 m
in seagrass and sand habitats of the northern Red Sea
(Zusehin and Oliver, 2003) and Huxleyia cavemicola
Hayami and Kase, 1993 is abundant in shallow sublit-
toral submarine caves of tbe Ryukyu Islands (Hayami
and Kase, 1993). Nucinella woodii (Dali, 1898) lives
in the Llorida Keys at subtidal depths, 65 to 188 m
(R. Bieler personal communication).
Reid (1990) speculated that nucinellids might be
chemosymbiotic and tbe presence of bacterial sym-
bionts in the ctenidial leaflets was later confirmed mor-
phologically for two Indian Ocean species (Oliver and
Taylor, 2012), by isotopic analysis of a New Zealand
fjord species (McLeod et al., 2010) and tbe symbiotic
Gammaproteobacteria identified moleeularly (Bieler et al.,
in press) for Nucinella giribeti new species, described
herein. Lossil monomyarian nucinellids, morphologi-
cally similar to living species, are recognized from the
early Jurassic where they are often associated with
dysaerobic environments (Harries and Little, 1999)
and others reported from Mesozoic hydrocarbon
seeps (Amano et al., 2007), with the likelihood that
they were also chemosymbiotic.
The relationship of Nucinellidae to other protobranchs
is still uncertain. Despite large morphological differences
in the shells Allen and Sanders (1969) suggested a close
Page 102
THE NAUTILUS, Vol. 127, No. 3
relationship with Solemyidae on the basis of anatomi-
cal similarity. Pojeta (1988) claimed a descent from
Palaeozoic “solemyoideans” through the dimyarian
Manzanella from the Permian. Since then nueinellids
have been classified in the superfamily Manzanelloidea
in the Solemyida, usually as the family Manzanellidae
(Bieler et al., 2010). For reasons discussed in Oliver
and Taylor (2012) we use the family name Nueinellidae
for Nucinella and Huxleyia rather than Manzanellidae,
which is based on the dimyarian Permian fossil
Manzanella (Chronic, 1952). Recent molecular results
provide little or no support for the monophyly of
Solemyidae +Nueinellidae (Bieler et al., in press; Sharma
et al., 2013).
We describe this new species from the Philippines,
firstly, because it is one of the few Nucinella known
from very shallow water and secondly, because it has
been included in molecular and morphological analyses
of bivalve phylogeny recorded only as Nucinella sp.
(Bieler et al., in press, Sharma et al., 2013). The other
nucinellid included in molecular analyses is Huxleyia
munita (Dali, 1898) from the northeastern Pacific.
MATERIALS AND METHODS
Nucinella shells were collected in the Philippines during
the Panglao 2004 Marine Biodiversity Project (see
Bouehet et al., [2009] for details). They were found
in a bulk sample collected by suction air-lift (station
S 1 1 ) and another single live-collected specimen from
station S39 was used for molecular analyses by Bieler
et al (in press).
Specimens were examined using Philips XL30 and
FEI Quanta 650 scanning electron microscopes (SEM)
following sputter coating in Au/Pd.
Institutional abbreviations used: MCZ, Museum of
Comparative Zoology, Harvard University, Cambridge,
USA; MNIIN, Museum National d’Histoire Naturelle,
Paris, France; NHMUK, Natural History Museum,
London, U K. Other abbreviations: H, height; L, lengtii,
PI, protoconch 1 All shell measurements in millimeters.
SYSTEMATICS
Subclass Protobranchia
Superfamily Manzanelloidea Chronic, 1952
Family Nueinellidae Vokes, 1956
Description: Shell obliquely oval, mostly higher than
long. Hinge with sub-umbonal taxodont teeth and
single, usually elongate lateral tooth on the anterior
dorsal margin. Ligament mostly opisthodetic, wholly
external or in a sunken resilifer. Shell sculpture of fine
com marginal lirae; periostraeum often glossy, persis-
tent, rather thick. Monomyarian, only anterior adduc-
tor muscle present. Anatomy protobranch with large
etenidia (Allen and Sanders, 1969; Kuznetsov and
Schileyko, 1984; Oliver and Taylor, 2012).
Remarks: Tl le main difference between Nucinella
and Huxleyia is the position of the ligament, external
or slightly sunken in the former or wholly internal in
the latter. For Nucinella, Oliver and Taylor (2012, fig. 7)
showed that there is considerable variation between
species in how deeply the ligament is sunken into the
hinge plate.
Genus Nucinella Wood, 1851
Type Species: Pleurodon ovalis Wood, 1840, non
Pleurodon Harlan, 1831. Syntype lots NHMUK Ramsholt
L4465, Sutton L4464, L4618. Features of N. ovalis, prob-
able syntypes, with details of the sub-umbonal teeth are
illustrated in Figures 1-5.
Diagnosis: As for the family, ligament external or in
a shallow resilifer.
Remarks: See Vokes (1956) for details of type
species designation. Nucinella ovalis is a Pliocene
fossil first described from the Coralline Crag, Ramsholt
and Sutton Members, Suffolk, England, (straddles
Pliocene stage boundary of Zanclean and Piacenzian,
3. 5-3.4 mya). Both localities have similar stratigraphy
and the palaeoenvironment is thought to be offshore,
shallow water to 50 m depth (Long and Zalasiewicz,
2011). Interestingly, Nucinella has been extinct since
the Pleistocene in the northeastern Atlantic and
Mediterranean (La Perna, 2004).
Nucinella giribeti new species
(Figures 6-17)
Description: Shell veiy small, largest valve L 2.8,
H 3.9 (largest live shell L 2.3, II 3.0), equivalve, outline
subovoid, external surface smooth with fine growth
increments only, periostraeum thick, ca. 5 pm, greenish,
shell colour white. Protoconch, PI 148 pm long (holo-
type), sharp boundary to post-larval shell. Ligament
opisthodetic short, set in shallow resilifer. Hinge, with
5-7 sub-umbonal teeth, slightly variable in size, younger
individuals may have fewer teeth (Figures 15-16). Holo-
type (Figures 11-14) right valve with 7 sub-umbonal
teeth, blade-like, slightly larger in posterior, anterior-
most tooth very small; lateral tooth prominent, short,
dorsal to adductor muscle scar; left valve with 6 sub-
umbonal teeth, 2 posterior larger, anteriormost very
small; lateral tooth as right valve. Anterior adductor
muscle scar large, ovoid, posterior scar absent. Inner
shell margin smooth. Image of live animal (Figure 6)
shows extended, multidigitate foot.
Type Material: Holotype, whole shell, MNHN 26701,
Figs 11-14, 17, L 2.0 H 2.7; Figured paratypes:
Three whole shells: NHMUK 20130108, L 2.2, H 2.9
(Figure 7) and MNHN 26702, L 1.9, H 2.6; L 2.3, H 3.0
(Figures 8-9); MNHN 26703, 1 right valve, L 1.5 H 2.1
(Figure 10); 2 right valves (Figures 15,16). Other
paratypes, 41 paired valves, 25 valves, MNHN 26704;
5 paired valves, 10 valves, NHMUK 20130108.
E.A. Glover and J.D. Taylor, 2013
Page 103
Figures 1-5. Nucinella ovalis (Wood, 1840), type species of Nucinella from Pliocene, Coralline Crag, Ramsholt, Suffolk, England.
Probable syntypes NHMUK L4465. 1. Exterior or right valve, L = 1.5 mm. 2-3. Interiors of right and left valves, L = 1.9 mm.
4-5. Details of hinges of left and right valves. Scale bar = 300 pm.
Type Locality: Philippines, Panglao Island, lagoon off
Poblacion 9°33.6' N, 123° 43.6' E, 2 m, fine sand and
seagrass, PANGLAO 2004, station SI 1
Other Material Examined: Sequenced specimen from
Philippines, Panglao Island, Tagbilaran-Panglao channel,
9°38.T N, 123° 5 1.4' E, 3-4 m, muddy sand beds of
Modiolus sp„ PANGLAO 2004, station S39. GenBank
numbers: 18S rRNA: KC429324; 28S rRNA: KC429414;
COI: KC429089.
Habitat and Distribution: Known only from or near
the type locality in shallow water sand with seagrass.
Etymology: Named for Gonzalo Giribet, evolution-
ary biologist responsible for the first sequencing of
Nueinellidae.
Remarks: The only other known species of Nucinella
from the Philippines is N. boucheti , which occurs at
depths of 1610-1580 m; this species, however, is much
larger and has 13 sub-umbonal teeth (La Perna, 2005).
In the tropical Indo-West Pacific three species of
Nucinella are recorded from Japan (Matsukuma et ah,
1982), a single species from off eastern Australia
(Vokes, 1956), a larger species off Zanzibar (Thiele
and Jaeckel, 1931), as well as the recently described
N. owenensis from off Oman; all are from deeper water
and differ in dentition from N. giribeti.
The type species of Nucinella, Pleurodon ovalis is a
Pliocene fossil from the Coralline Crag of England
that is similar in size and sub-umbonal dentition to
N. giribeti but has a much longer anterior lateral tooth.
(Figures 1-5).
DISCUSSION
Nucinella giribeti is the only species of the genus
recorded from intertidal depths and it is very sur-
prising that there are no other records from similar
habitats in Southeast Asia. In all probability they
have been overlooked because of their very small
size, but the discovery in Panglao indicates that they
can be quite abundant. The N. giribeti from the type
locality were associated with ehemosymbiotic lucinids
Pillucina new species and Cardiolucina siquijorensis,
both of which occurred in abundance in the seagrass
sediments (Taylor and Glover, 1997; Glover and Taylor,
in press). Both stations also contained several species of
cardiids including Fragum spp., Microfragum spp., and
Fulvia sp. (ter Poorten, 2009); other bivalves have yet to
be studied.
Most nueinellids, along with many other protobranch
mollusks, are found in deeper water from shelf to
hathyal depths, with one of the deepest so far recorded
being Nucinella owenensis recently described from
3400 m in an CU minimum zone, associated with
deposit feeding and other ehemosymbiotic bivalves
(Oliver and Taylor, 2012). The Japanese species
Nucinella viridis Matsukuma et ah, 1983 was found
at 3581 m at a likely hydrocarbon seep (Okutani and
Iwasaki, 2003; Sasaki et ah, 2005).
Life histories of Nueinellidae are poorly under-
stood, but in common with other protobranehs, they are
Page 104
THE NAUTILUS, Vol. 127, No. 3
Figures 6-17. Nucinella giribeti new species. 6. Live specimen, 0-3 m Panglao Island, Philippines. Image by Pierre Lozouet
MNHN. 7-9. Paratypes. 7. Left side (NI1MUK 20130108), H = 2.9 mm. 8. Right side, (MNHN 26702), H = 2.6 mm. 9. Left side
(MNIIN 26702), H = 3.0 mm. 10. Paratype, exterior of right valve (MNHN 26703), L = 1.5 mm, H = 2.1 mm. 11-12. Holotype
(MNHN 26701), interior of left and right valves, L = 2.0 mm, H = 2.7 mm. 13-14. Holotype, detail of hinge teeth of left
and right valves. Scale bar = 200 pm. 15. Paratype, right valve, detail of hinge to show tooth variation (MNHN 26703). Scale
bar = 200 pm. 16. Paratype, right valve detail of hinge teeth (MNHN 26703). Scale bar = 200 pm. 17. Protoconch of right
valve of holotype. Scale bar = 50 pm.
E.A. Glover and J.D. Taylor, 2013
Page 105
presumed to have only lecithotrophic planktonic devel-
opment. Protoconch sizes are variable: the type species
N. ova l is is at least 210 pm, N. giribeti is 148 pm and
N. seguenzae (Dali, 1898) (a Pleistocene fossil from
Spain) is 260 m (La Perna, 2004).
Nucinella giribeti and Huxleyia munita (Eastern
Pacific) are the first nucinellids to be included in molecular
phylogenies of bivalves (Sharma et ah, 2013; Bieler et ah,
in press), with results indicating a long independent his-
tory of these bivalves separate from tire Sclemyoidea.
The pre-Mesozoic history of the Nucinellidae is uncer-
tain but by early Jurassic times nucinellids, similar in
morphology to living forms, were present in dysaerobic
sediments (Harries and Little, 1999; Wignall et ah, 2005)
with later Cretaceous records from hydrocarbon seeps
(Amano et ah, 2007) indicating the high probability of
early acquisition of chemosymbiosis in the family.
ACKNOWLEDGMENTS
We are grateful to Dr. Philippe Bouehet and his team
(MNHN) for access to the Nucinella specimens. Pierre
Lozouet (MNHN) kindly allowed us permission to
publish Figure 6. We thank Alex Ball (N1IMUK) for
assistance with SEM, Riidiger Bieler (Field Museum,
Chicago) for details of die habitat of Nucinella woodii ,
and Liz Harper (University of Cambridge) for advice on
the Coralline Crag.
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THE NAUTILUS 127(3): 107-1 14, 2013
Page 107
Vertigo marciae (Gastropoda: Vertiginidae), a new land snail
from Jamaica
Jeffrey C. Nekola
Department of Biology
University of New Mexico
Albuquerque, NM 87131 USA
[email protected]
Gary Rosenberg
Department of Malacology
Academy of Natural Sciences of Drexel University
1900 Benjamin Franklin Parkway
Philadelphia, PA 19103 USA
[email protected]
ABSTRACT
Vertigo marciae , a new species of gastropod mollusk (Pupilloidea:
Vertiginidae), is described from Jamaica. This species is known
in the Recent fauna only from John Crow Peak in the Blue
Mountains, but also occurs as a Pleistocene fossil at Red Hills
Road Cave. Vertigo marciae has been confused with Vertigo
gouldii , but differs by its smaller shell size, lack of distinct
shell striation, lack of an angular lamella, and presence of a
flared aperture base. DNA sequence analyses document that
V marciae possesses unique mtDNA and nDNA sequences
and is most closely allied with Vertigo alabamensis. Vertigo
hebardi, and Vertigo oscariana. This group of species com-
prises a highly supported clade whose members are limited
either to the Caribbean or the southeastern USA
Additional Keywords: Vertigo, biogeography, southeastern USA,
Caribbean, DNA sequence analysis
INTRODUCTION
In their synopsis of the Jamaican land snail fauna,
Rosenberg and Muratov (2006) reported three species
from the genus Vertigo , all of which possess ranges
extending into the eastern half of North America
(Nekola and Coles, 2010): V. gouldii, V. milium , and
V. ovata. However, images of putative Jamaican
V gouldii’ on the Discover Life website ( h ttp ://pick4
.pick.uga.edu/mp/20q?search=Vertigo+gouldii&guide=l)
illustrate a shell quite unlike V gouldii from eastern
North America (Nekola and Coles, 2010) in its light-
yellow shell color, lack of sharp shell striation, and
absence of a basal lamella. Observation of the three
known Jamaican lots for this entity in the Academy of
Natural Sciences of Philadelphia (ANSP) collections
confirmed these differences. Although they looked most
like V. hebardi Vanatta, 1912 of the Llorida Keys, these
lots clearly differed from that species by their taller shell,
reduced striation, and absence of an angular lamella.
Thus, rather than representing a population of V7 gouldii
isolated by 2000 km from its nearest neighbors (Nekola
and Coles, 2010), these specimens appear to represent
an undescribed new species. Based on shell descrip-
tions, reports of Vertigo gouldii from the Pleistocene
Red Hills Road Cave deposits of Jamaica (Paul and
Donovan, 2005) also appeared to represent this new spe-
cies. This conclusion was subsequently confirmed via
observation of digital images.
We therefore investigated this putative new spe-
cies via analyses of both shell morphology and DNA
sequence data, and are now in the unusual situation of
describing anti reporting a new species simultaneously
from both living and fossil material, while also being
able to report on its phylogenetic relationships.
MATERIALS AND METHODS
Field Collection: Tl le three known Recent lots of the
putative new Jamaican species (ANSP 402244, 403039,
and 403040) were collected on John Crow Peak in the
Blue Mountains of eastern Jamaica on May 22, 1999,
during the Jamaican Biotic Survey. Collecting methods
were detailed in Rosenberg and Muratov (2006), and
included drying of soil litter samples over a Berlese
Lunnel. This encouraged snails to enter aestivation, allow-
ing them to mummify upon death. Lor this reason, it was
possible to successfully extract, amplify, and sequence
selected amplicons from both their mitochondrial and
nuclear genomes (Nekola et ah, 2009).
Shell Measurements: Measurements were deter-
mined in 0.1 mm increments for adult shells with
aperture facing up, using a dissecting microscope with
a calibrated ocular micrometer. Height and width were
measured as the dimensions of a bounding box with
long axis parallel to the shell axis and sides tangent
to the tip of the protoconch, the base of the lip, the
right-most margin of the aperture and the left-most
margin of the body whorl.
Imaging: Shells were imaged at 20 x magnification
using a digital camera attached to a stereomicroscope.
Page 108
Approximately 14 separate 1388x1040 pixel images
were made of each specimen with the image focal
lengths positioned at 75 pm increments from die front
to back of the shell. CombineZ5 freeware (http://www
.hadleyweb.pwp.blueyonder.co.uk/CZ5/combinez5.htm)
was used to assemble a final image from the focused
parts of each separate image. Assembled images were
imported into Adobe Photoshop, where brightness and
contrast were optimized anti the background made uni-
formly black. These images were then compiled into a
single plate.
DNA Sequence Analysis: Mitochondrial cytochrome
b ( Ci/tB ), I6S ribosomal RNA ( 16S ) and the internal tran-
scribed spacers 1 ( ITS-1 ) and 2 ( 1TS-2 ) of the nuclear
ribosomal RNA gene were investigated to test the dis-
tinctiveness of the putative new Jamacian species and to
resolve its nearest evolutionary neighbors. The mito-
chondria] cytochrome oxidase subunit 1 (COl) was not
analyzed as we were unable to amplify this gene in the
Jamaican species.
Sixteen specimens were chosen for analysis (Table 1).
This set includes not only two specimens of the
Jamaican species but also Vertigo alabamensis, V
gouldii, V hebardi, and V. oscariana. We attempted to
maximally spread these individuals across the known
geographic range of each species (see Nekola and
Coles, 2010). Because V. conecuhensis is likely a simple
shell form of V. alabamensis (Nekola and Coles, 2010),
THE NAUTILUS, Vol. 127, No. 3
only a single individual of this taxon was included.
Topotype or near-topotype material was selected for
V. conecuhensis (Pond Creek, Covington Co., Alabama,
about 50 km from the type locality in Evergreen,
Alabama) and V. hebardi (Long Key, Florida which is
the type locality). Previously analyzed 16S sequence
data from three Vertigo gouldii specimens analyzed
by Nekola et al. (2009) were retrieved from GeneBank
(Table 1). Vertigo pusilla , the type species of the genus,
was also included, as were Gastrocopta cristata and
Papilla muscorum for outgroup comparisons.
Genomic DNA was extracted from live, ethanol-
preserved, or mummified material using the OmegaBioTek
Mollusk DNA Extraction Kit. Because shell destruc-
tion was required, all shells were imaged prior to
extraction using the methods detailed above. PCR
amplification and sequencing of CijtB , 16S, ITS-1, and
ITS-2 were accomplished using standard methods
(Nekola et til., 2009). Resultant traces were examined,
primer ends removed, and aligned by eye. Because of
their lower evolutionary rates, the nDNA ITS-1 and
ITS-2 regions were concatenated to provide a roughly
similar number of nucleotide differences to either of
the mtDNA amplicons. Mega 5.0 (Tamura et ah, 201 1)
was used to calculate the average number of nucleotide
differences in the CytB, 16S, and concatenated ITS-1 +
ITS-2 regions between all putative species-level taxa.
Substitutions included both transitions and transversions
with pairwise gap deletion.
Table 1. Specimen information for material used in DNA sequence analysis.
Vertigo marciae new species
John Crow Peak, St. Andrew Parish Jamaica 18.1 132
John Crow Peak, St. Andrew Parish, Jamaica 18.1132
Vertigo alabamensis Clapp, 1915
Johnson Mill Bay, Bladen Co., North Carolina, USA 34.7125
Wolf Trap Bay, Leon Co., Florida, USA 30.3680
Vertigo conecuhensis Clapp, 1915
Pond Creek, Covington Co., Alabama 31.1036
Vertigo gouldii (A. Binney, 1843)
Brush Creek Canyon, Fayette Co. Iowa, USA 42.7796
Panther Creek, Searcy Co., Arkansas, USA 36.0858
1 1-Point River, Oregon Co., Missouri, USA 36.7931
Vertigo hebardi Vanatta, 1912
Long Key, Monroe County, Florida, USA 24.8146
Elliott Key, Miami-Dade County, Florida, USA 25.4553
Vertigo oscariana (Sterld, 1890)
Blanchard Springs, Stone Co., Arkansas, USA 35.9582
Wadboo Creek, Berkeley Co., South Carolina, USA 33.1971
Vertigo pusilla Muller, 1774
Podyji National Park, Moravia, Czech Republic 48.8586
Gastrocopta cristata (Pilsbry and Vanatta, 1900)
Albuquerque, Bernalillo Co., New Mexico, USA 35.0727
Pupilla muscorum (Linnaeus, 1758)
Masaryk University, Brno, Moravia, Czech Republic 49.2509
J.C. Nekola and G. Rosenberg, 2013
Page 109
Mega 5,0 was used to conduct nearest-neighbor join-
ing (NNJ), maximum parsimony (MP), and maximum
likelihood (ML) trees for the three focal regions. NNJ
was based on Maximum Composite Distance including
transitions and transversions with pairwise gap dele-
tion. MP used the close neighbor interchange search
option with the random addition of 10 replicate trees.
ML used all sites and was based on the Tamura-Nei
substitution model, a five-category Gamma Distribu-
tion for substitution rates, and the Nearest Neighbor
Interchange ML heuristic method. In all cases sup-
port values were estimated from 1000 bootstrap repli-
cates. Additionally, Bayesian trees were generated using
Mr Bayes 3.1 (H uelsenbeck and Ronquist, 2001) using
a GTR substitution model assuming gamma-shaped rate
variation over 1,000,000 generations with a sampling fre-
quency of once each 1000 generations.
Nomenclature: Taxonomic names and concepts follow
Nekola and Coles (2010). Apertural lamellae and fold
nomenclature follows that of Pilsbry (1948: 869, fig. 469),
i.e., parietal “teeth” are referred to as “folds” and all
other “teeth” are termed “lamellae”, whatever their form.
Also, we follow Pilsbry (1948) by referring to the raised
riblets on the surface of Vertigo shells as “striae.”
SYSTEMATICS
Class Gastropoda
Subclass Pulmonata
Order Stylommatophora
Family Vertiginidae
Genus Vertigo Muller, 1773
Vertigo marciae new species
(Figures 1-6; Tables 1, 2)
GenBank Accessions KF214477, KF214489, KF214502,
KF214503, KF214513, KF214514
Diagnosis: Minute shell reminiscent of Vertigo hebardi
but tiiller, less striate, and lacking angular lamella; shell
surface smooth and glossy with indistinct wrinkles; aper-
ture flared toward bottom; four lamellae/folds present,
including parietal, columellar, and two palatals.
Description: Shell 1.4-1. 6 mm tall x 0.8-1. 0 mm wide
(holotype 1.5 x 0.8 mm), columnar-ovoid to ovoid,
approximately 4-4.5 whorls, with moderately shallow
suture and domed apex. Translucent, pale yellow-brown
color. Body whorl approximately 60% of total height
(Figures 1-3, 5, 6). Protoconch and neanic whorls
smooth (Figure 4), with subsequent whorls having
irregular, infrequent, and weak wrinkles. Immediately
behind aperture sculpture takes the form of irregular
low striae (Figure 2). Aperture flared on bottom, making
it taller than wide and approximately 1/3 of shell
height. Lip unthickened and slightly reflexed, sinulus
moderate-weak, sometimes expressed as simple flatten-
ing of palatal wall. B as ally aperture abruptly inflates to
form rounded swelling but not crest (Figure 4). Umbili-
cus closed (Figure 3). Aperture with four lamellae/folds;
parietal lamella strong, slightly sinuous (Figures 1, 2, 5,
6); columellar lamella downward-sloping, peg-shaped;
two palatal folds with lower being approximately twice as
long as upper and extending approximately 0.2 whorls
into body whorl, lower slightly more immersed than
upper, both highest at mid length (Figures 1, 2, 5, 6).
Apertural end of lower palatal fold coincides with abrupt
inflation of basal aperture. Externally shell only slightly
impressed over palatal folds (Figure 4).
Type Material: Holotype (Figures 1 -4), ANSP 450580,
from type locality, May 22, 1999; paratypes (Figures 5-6),
ANSP 402244, 8 shells and Institute of Jamaica, 2 shells,
from type locality; ANSP 403039, Jamaica, St. Andrew'
Parish, John Crow Peak, Bine Mountains, elfin forest
with bamboo near summit, 18°05'45" N, 16° A0' OKA" W,
altitude 1755 nr. May 22, 1999; sta. JBS4a, 16 shells;
ANSP 403040, Jamaica, St. Andrew Parish, Vinegar Hill
Trail near head of Clyde River, 1 8°05' N, 76°39,18" W,
altitude 1520 m. May 22, 1999, sta. |BS 5, 3 shells.
Type Locality: Jamaica, St. Andrew Parish, John
Crow Peak, Blue Mountains; litter sample collected at
base of limestone cap, 18°5,50" N, 76°40,5" W, altitude
1550 m. May 22, 1999, sta. JBS4e.
Other Material Examined: Digital images of two speci-
mens from Red Hills Road Cave, Jamaica (Paul and
Donovan, 2005), Paul collection.
Etymology: Tbe specific name marciae refers to
Dr. M areia Mundle, then of the Jamaica Conservation
and Development Trust. Dr. Mundle arranged for the
vehicle used and park ranger guide that accompanied
the expedition to the type location, and was present
when the species was first collected.
RESULTS AND DISCUSSION
Variation: Vertigo marciae is rather constant in gen-
eral appearance in terms of its shape, color, sculpture,
and development of the apertural lamellae. While some
variation in size (especially height) was noted, this was
minor — only 16% difference between the largest and
smallest shells was observed.
Comparison with Other Vertigo Species: Vertigo
marciae differs from all other Vertigo species by its
small (<1.7 mm in height) yellow shell with indistinct
striae/wrinkles, flared aperture base, and lack of angular
and basal lamellae. It is closest in appearance to
V. hebardi of tbe Florida Keys (Figure 7), with which
it shares a small vellow shell, the lack of a basal lamella,
and preference for accumulations of tropical forest leaf
litter. However, it differs from this species in its taller
shell with less distinct striae and absence of an angular
lamella. It is also reminiscent of Vertigo marki Gulick,
Page 1 10
THE NAUTILUS, Vol. 127, No. 3
Figures 1-12. Stereomieroscope images of Vertigo marciae and related taxa. 1-4. Vertigo marciae , holotype, Jamaica, St. Andrew
Parish, John Crow Peak, Blue Mountains, 18°5'45" N, 76°40'8" W to 18°5'50" N, 76°40'5" W, ANSP 450580. 1. Apertural view.
2. Profile. 3. Umbilical view showing parietal and upper palatal lamellae. 4. Apical Mew showing protochoneh and the basal
apertural dilation. 5. Vertigo marciae, ANSP 402244, Paratype, second specimen from the type locality, exhibiting a shorter
shell. 6. Vetiigo marciae, ANSP 402244, Paratype, third specimen from the type locality, exhibiting a slightly more worn shell.
7. Vertigo hebardi. Long Key, Monroe County, Florida, 24°48'52" N, 80°49'14/' W, JCN 17479. 8. Vertigo goidelii (small southern
form), Tellieo Gorge, Monroe Co.. Tennessee, 35°19'49" N, 84°10'59" W, BFC 1332. 9. Vertigo alabamensis, Lanier Quarry, Pender
Co., North Carolina, 34°37'49" N, 77°40'27" W, JCN 10781. 10. Vertigo “ conecuhensis Pond Creek seep, Covington Co., Alabama,
31°6'12" N, 86°32'3" W, JCN 12364. 11. Vertigo oscariana, Wadboo Creek, Berkeley Co., South Carolina, 33°11'50" N, 79°56'46" W,
JCN 10908. 12. Vertigo gouldii (normal form), Deer Creek, Fillmore Co., Minnesota, 43°43'56" N, 92o20'39" W, JCN 14646.
J.C. Nekola and G. Rosenberg, 2013
Page 1 1 1
1904 from Bermuda, with which it shares a smooth,
yellowish shell that lack both basal and angular lamellae
(Pilsbry, 1919). However, V. marciae differs from this
species by its smaller shell height and lack of a callus
on the palatal wall of the aperture. Vertigo oscariana
from the southeastern USA (Figure 11) also has a
small, smooth yellow shell without a basal lamella,
but this species also lacks an upper palatal fold,
has a callus on the outer apertural margin, and a
shell that is widest in the middle, tapering both to
base and apex. Vertigo alabamemis of the southeastern
North American coastal plain (Figures 9, 10) also has a
yellowish shell lacking striae, but this species is larger
(generally > = 1.8 mm tall), possessing both angular and
basal lamellae, a strong crest on the apertural margin,
and a strong sinulus on the palatal wall of the aperture.
Rosenberg and Muratov (2006) identified Vertigo
marciae as Vertigo gouldii (Figures 8, 12) because of
its general shell shape and placement of the parietal
lamella and palatal folds, and also because V gouldii
was reported from Jamaica by Pilsbry and Cooke
(1919: 99). However, V gouldii has a larger shell of
brown color, has a duller shell luster from the pres-
ence of abundant strong but irregular striae, and always
possesses a basal lamella. Specimens documenting the
Pilsbry and Cooke record have not been traced, and are
apparently based on personal communication between
Victor Sterld and Pilsbry. Bequaert and Miller (1973: 95)
rejected Antillean records of V gouldii.
While Vertigo milium — which also occurs in Jamacian
tropical forest leaf litter — possesses a shell <1.7 mm tall;
this species is easily distinguished from V. marciae in
its dark red-brown color, presence of both angular and
basal lamellae, and its long, curved lower palatal fold
which deeply enters the shell.
Geographic Distribution and Ecology: Vertigo
marciae is currently known in the Recent fauna only
from the crest of John Crow Peak and its immediate
vicinity in the Blue Mountains of eastern Jamaica at ele-
vations of 1520-1755 m. John Crow Peak is capped by
an isolated limestone outlier known to have distinctive
plant communities, including endemic species (Grossman
et ak, 1993). (Note that John Crow Peak is not the same
location as tire John Crow Mountains, the easternmost
range of Jamaica.) Vertigo marciae is found mainly on or
adjacent to limestone boulders and outcrops in tropical
as well as scrub forest with bamboo, but has also been
found at one site (JBS 5) that lacks exposed limestone.
As a Pleistocene fossil, it is known only from Red Hills
Road Cave, which is about 21 kin west of Rosenberg
and Muratovs sites on John Crow Peak and 1000 m
lower in elevation (520 m, Paul and Donovan, 2005).
Vertigo marciae joins the two Radiodiscus species and
the Puuctum reported by Paul and Donovan (2005) on
tire basis of personal communication from Rosenberg
as species known in Jamaica only from John Crow Peak
and from the Red Hills Road Cave. Paul and Donovan
interpreted the faunal changes from the Red Hills Road
Cave fauna to the recent fauna around the cave as
suggesting drying of the climate since the Pleistocene.
As Vertigo marciae was found nowhere else in Jamaica
among the hundreds of sites sampled by Rosenberg
and Muratov, its current occurrence might represent a
relict distribution.
Although it must currently be considered a Jamaican
endemic, the general lack of local endemism in the
genus Vertigo (Nekola, 2009) suggests the possibility
that it may occur elsewhere in the adjacent Caribbean,
especially on carbonate substrates in mid- to high-elevation
montane forest.
Table 2.
Average number of base-pair differences between all pairwise combinations of Vertigo marciae and related species.
Page 112
THE NAUTILUS, Vol. 127, No, 3
Genetic Distinctness and Phylogenetic Relationships:
DNA sequence analysis clearly demonstrates that
V marciae is distinct at the species level. It pos-
sesses an average of 36 and 39 base pair dilferences
with V. hebardi and V. alabamensis , respectively,
in the 355 bp CytB amplicon (Table 2). This equates
to a 10-11% variation across the entire amplicon. In
addition, it possessed an average of 46.5 base pair
differences in CytB (13%) with V. oscariana and
63.8 differences (18%) with V. gouldii. In the more
slowly evolving 443 to 447 bp 16S region V. marciae
differed by 18 bases (4.0%) from V. hebardi and
V. alabamensis , 42 bases (9.4%) from V. oscariana ,
and 42.5 bases (9.6%) from V gouldii. In the
concatenated 1274-1284 bp ITS-1 + ITS-2 nDNA
amplicon, V. marciae possessed an average of 9 base-
pair (0.7%) differences with V. alabamensis , 12 (0.9%)
with V. hebardi , 23.5 (1.8%) with V. oscariana, and
37.3 (2.9%) with V gouldii. These levels of differ-
ence were considerably larger than those exhibited
between the conehologieally distinct V. alabamensis
and V. hebardi (Figures 7, 9, 10), which amounted
to an average difference of 8.3 bases (2.3%) in CytB,
4.0 (0.9%) in 16S, and 5.0 (0.4%) in concatenated
ITS-1 and ITS-2.
The topologies across phylogenetic reconstructions
were largely compatible, with the same highly supported
nodes being identified in all cases (Figures 13—15).
These demonstrate that V. marciae is clearly a member
of the genus Vertigo, being within the same highly
supported clade in Ci/tB and 16S that includes
V pusilla, the type species of the genus. While it was
not possible to root die concatenated ITS-1 + ITS-2 tree
because of profound differences with both Gastrocopta
cristata and Papilla muscorum sequences, V. marciae
was easily aligned with all Vertigo sequences.
Tt le phylogenetic reconstructions also demonstrate
that V. marciae is a member of a highly supported
clade that includes both V alabamensis and V. hebardi.
The existence of a single ancestor to all of these species
implies long distance dispersal from the southeastern
United States to Jamaica, perhaps with migrating birds
as a vector. This is not an unreasonable scenario, given
that much longer feats of long distance dispersal via
migrating birds have been documented in the eastern
Atlantic (Gittenberger et al., 2006).
95 95
88 92
99 99
100 97
\
V. conecuhensis - Alabama
V. alabamensis - North Carolina
99 99
100 99
95 94 gg 93 ^
100 94 79 9o
99 99
100 99
L V. alabamensis - Florida
V. hebardi - Elliott Key, Florida
V. hebardi - Long Key, Florida
V. marciae - Jamaica
V. marciae - Jamacia
95 95
97 87
— V. oscariana - South Carolina
V. oscariana - Arkansas
98 99
79 82
V. pusilla - Moravia, Czech Republic
V. gouldii - Missouri
V. gouldii - Arkansas
V. gouldii - Iowa
V. gouldii - Maine
Pupilla muscorum - Brno, Czech Republic
Gastrocopta cristata - New Mexico
0.05
Figure 13. The phylogenetic relationships of Vertigo marciae as reconstructed by maximum-likelihood analysis from the CytB
amplicon. Nodes with strong to moderate support across all four phylogenetic reconstruction methods have been labeled to the
left of that node by four support values: The upper left (normal font) is for nearest neighbor joining. The upper right (bold italic
font) is for maximum parsimony. The lower left (bold font) is for Bayesian. The lower right ( italic font) is for maximum likelihood.
J.C. Nekola and G. Rosenberg, 2013
Page 1 13
97 97
93 93 V. conecuhensis - Alabama
98 88 \
r V. alabamensis - Florida
100 S3
94 95
96 77
100 99 V. alabamensis - North Carolina
V. hebardi - Elliott Key, Florida
V. hebardi - Long Key, Florida
V. marciae - Jamaica
V. marciae - Jamaica
- V. pusilla - Moravia, Czech Republic
72 72
86 69
99 99
93 96
99 99
100 100
100 94
99 99
60 61
99 92 T S,0U^H " Arkansas
x r V gouldii - Missouri
V. gouldii - Maine
V. gouldii - Iowa
V. oscariana - South Carolina
V. oscariana - Arkansas
Pupilla muscorum - Brno, Czech Republic
Gastrocopta cristata - New Mexico
0.05
Figure 14. The phylogenetic relationships of Vertigo marciae as reconstructed by maximum-likelihood analysis from the 16S
amplicon. Nodes with strong to moderate support across all lour phylogenetic reconstruction methods have been labeled to the
left of that node by four support values: The upper left (normal font) is for nearest neighbor joining. The upper right (bold italic
font) is for maximum parsimony. The lower left (bold font) is for Bayesian The lower right (italic font) is for maximum likelihood.
100 100
100 100
85 85
94f W
98 99
100 98
/
85 86
100 78
V. alabamensis - Florida
V conecuhensis - Alabama
V. alabamensis - North Carolina
V. hebardi - Long Key, Florida
— V. hebardi -Elliott Key, Florida
V. marciae - Jamaica
100 100
— V. oscariana - Arkansas
100 wo
V. oscariana - South Carolina
V. gouldii - Maine
99 99
100 99
— V. gouldii - Iowa
V. gouldii - Missouri
V. pusilla - Moravia, Czech Republic
0.005
Figure 15. The phylogenetic relationships of Vertigo marciae as reconstructed by maximum-likelihood analysis from the con-
catenation of the ITS- 1 and ITS-2 amplicons. Nodes with strong to moderate support across all four phylogenetic reconstruction
methods have been labeled to the left of that node by four support values: The upper left (normal font) is for nearest neighbor
joining. The upper right (bold italic font) is for maximum parsimony. The lower left (bold font) is for Bayesian. The lower right
(italic font) is for maximum likelihood.
Page 114
THE NAUTILUS, Vol. 127, No. 3
While resolution of deeper nodes within the genus
Vertigo was not possible in the Ci/tB and 16S mtDNA
amplicons (Figures 13, 14), presumably due to base-
pair saturation, the concatenated ITS-1+ ITS-2 data
(Figure 15) demonstrated a very highly supported node
connecting the V. marciae / V. hebardi / V. alabamensis
elade to V. oscariana. This radiation includes some of
the most distinct members of the genus (Pilsbry, 1948).
Additional field work across the Caribbean — especially
in Cuba, Hispaniola, Puerto Rico, and Bermuda — will
be required to determine the actual number of species
contained within this group.
ACKNOWLEDGMENTS
John Slapcinsky of the Florida Museum of Natural His-
tory provided ethanol-preserved tissue samples of Vertigo
hebardi, while Michal Horsak of Masaryk University
in Brno, the Czech Republic, provided live samples
for Vertigo pusilla and Papilla muscorum. Ci/tB and 16S
sequences for V. pusilla , P. muscorum, and Gastrocopta
cristata were retrieved from full mitochondria] geno-
mic sequences provided hy Jason Marquardt and
Ulfar Bergthorsson of the University of New Mexico.
Christopher Paul, University of Bristol, provided digital
images of the Red Hills Road Cave fossils. Field work
which established the persistence of Vertigo hebardi in
the Florida Keys was supported hy The Bailey- Matthews
Shell Museum through an R.T. Abbott Visiting Curator-
ship to JCN. Additional funding for PCR and sequenc-
ing analysis was provided hy Michal Horsak. The field
work which resulted in the Recent samples of the new
species being collected was supported by NSF Grant
DEB-9870233 to Gary Rosenberg.
LITERATURE CITED
Bequaert. J.C. and W. B. Miller. 1973. The Mollusks of the
Arid Southwest. University of Arizona, Tucson, xvi+271 pp.
Gittenberger, E., D.S.J. Groenenberg, B. Kokshoorn and
R.C. Preece. 2006. Molecular trails from hitch-hiking snails.
Nature 439: 409.
Grossman, D.H., S. Iremonger, and D M. Muchoney.
1993. Jamaica: map of natural communities and
modified vegetation types. Jamaica: a rapid ecological
assessment. Phase 1: An island-wide characterization
of mapping of natural communities and modified
vegetation types. The Nature Conservancy, Washington,
DC., USA.”
Huelsenbeck, | P. and F. Ronquist. 2001. MB BAYES:
Bayesian inference of phylogeny. Bioinformatics 17:
754-755.
Nekola, J.C. and B.F. Coles. 2010. Pupillid land snails of
eastern North America. American M alacological Bulletin
28: 29-57.
Nekola, | C. 2009. Big ranges from small packages: North
American vertiginids more widespread than thought.
The Tentacle 17: 26-27.
Nekola, | C., B.F. Coles, and U. Bergthorsson. 2009. Evo-
lutionary pattern and process in the Vertigo gouldii
(Mollusca: Pulmonata, Pupillidae) group of minute North
American land snails. Molecular Phylogenetics and Evo-
lution 53: 1010-1024.
Paul, C.R.C. and S.K. Donovan. 2005. Quaternary and
Recent land snails (Mollusca: Gastropoda) from Red
Hills Road Cave, Jamaica Bulletin of the Mizunami
Fossil Museum 32: 109-144.
Pilsbry, H A. 1919. American Species of Vertigo. Manual
of Conchology. Second Series (Pulmonata) 25: 74-150,
pis. 6-13.
Pilsbry, H. A. 1948. Land Mollusca of North America (North of
Mexico). Academy of Natural Sciences of Philadelphia
Monographs 3, vol. 2(2): i-xlvii, 521-11 13.
Rosenberg, G. and IV. Muratov. 2006. Status report
on the terrestrial mollusca of Jamaica. Proceedings of
the Academy of Natural Sciences of Philadelphia 155:
117-161.
Tamura, K., D. Peterson, N. Peterson, G. Steelier, M. Nei,
and S. Kumar. 2011. MEGA5: molecular evolutionary
genetics analysis using maximum likelihood, evolutionary
distance, and maximum parsimony methods. Molecular
Biology and Evolution 28: 2731-2739.
THE NAUTILUS 127(3): 1 15-1 18, 2013
Page 115
Cornisepta guzmani new species: first species of genus confirmed
from Pacific cold seep environments off central Chile
(Gastropoda: Vetigastropoda: Fissurellidae)
Juan-Francisco Araya
Laboratories Cecilia Osorio
Universidad de Chile, Facultad de Ciencias
Las Palmeras 3425
Santiago, CHILE
[email protected]
Daniel L. Geiger
Santa Barbara Museum of Natural History
2559 Puesta del Sol
Santa Barbara, CA 93105 USA
[email protected]
ABSTRACT
The new bathyal gastropod species, Cornisepta guzmani new
species, collected at a depth of 846 metres from a methane
seepage site in the subduction zone off Concepcion (36°22' S;
73°43' W), central Chile, is described. The new species is
most similar to Cornisepta pacifica (Cowan, 1969), described
from offshore Alaska and Oregon, from which it differs in
shell morphology, sculpture, and geographic distribution.
Additional Keywords: Deep water, deep sea
INTRODUCTION
The genus Cornisepta was established to group a series
of small, deep-water fissurellids with high conical
shells, lacking a selenizone, with a prominent internal
septum, and that shed the protoconch as they age
(McLean and Geiger, 1998). The genus comprises thir-
teen extant worldwide species, of which seven have
been described from the Pacific Ocean: Cornisepta
antarctica (Egorova, 1972), C. festiva (Crozier, 1966),
C levinae McLean and Geiger, 1998, C. monsfuji Chino,
2009, C. pacifica (Cowan, 1969), C. soyoae (Habe, 1951)
and C. verenae McLean and Geiger, 1998. The genus
has records up to the early Oligocene, with two species
described for the early Rupelian of Germany; Cornisepta
anhaltina Muller, 2011 and C. g ranulocostata Muller,
2011 (Muller, 2011). Recent species of Cornisepta have
been recorded in deep-sea environments, in faunal asso-
ciations of seamounts (Beck et al., 2006), in reducing
systems at hydrothermal vents in mid-oceanic ridges
(Sasaki et al., 2010), and cold seeps at continental mar-
gins (Gracia et al., 2011). Little is known of their popula-
tion biology, ecology and conservation status.
This article describes Cornisepta guzmani based on
shell morphology and sculpture of two specimens col-
lected in methane-hydrate rubble and inside the valves
of a vesieomyid clam collected in a methane cold seep
off Concepcion Bay, central Chile. In addition, exter-
nal morphology and details of sculpture were com-
pared with that of congeneric species in the Pacific
Ocean and a species from the Atlantic Ocean.
MATERIALS AND METHODS
The description of the new species is based on two
specimens collected from rubble trawled from a methane
cold seep off tire coast of Concepcion Bay, central Chile
(36° 22' S; 73° 43' W). The holotype and paratype are
deposited under accession number 2013-001 in the col-
lections of the Santa Barbara Museum of Natural History,
Santa Barbara, California, USA (SBMNH).
For the morphological descriptions, the length (L),
height (H) and width (W) of the shells were measured
from photographs and scanning electron micrographs.
The position of the foramen (PF) was defined as the
distance of the centre of the foramen measured from
the anterior shell margin.
SYSTEM ATICS
Class Gastropoda Cuvier, 1 797
Suborder Vetigastropoda Salvini-Plawen, 1980
Family Fissurellidae Fleming, 1822
Subfamily Emarginulinae Children, 1834
Genus Cornisepta McLean and Geiger, 1998
Type Species: Fissurisepta antarctica Egorova, 1972
(by original designation). Recent, Weddell Sea, Antarctica
(McLean and Geiger, 1998: 18).
Remarks: Chino (2009) and Miiller (2011) errone-
ously attributed the genus name to McLean alone.
Page 116
THE NAUTILUS, Vol. 127, No. 3
Cornisepta guzmani new species
(Figures 1-7)
Diagnosis: Small fissurellid (up to 5.3 mm), conical
profile, foramen at summit of shell, anterior slope
convex, posterior slope concave, septum high, across,
thin, sculpture of widely spaced pustules randomly
hyperdispersed.
Description: Shell of moderate size for genus,
yellowish-white, conical, anterior slope convex, pro-
file moderately high (66% of length), posterior slope
concave. Juvenile shell and protoconch unknown.
Foramen oval, situated in posterior third of shell
length. Sculpture of concentric growth marks with
widely spaced pustules arranged randomly hyperdis-
persed. Interior of shell glossy, septum thin, slightly
curved, transverse, extending obliquely downward at
an angle of approximately 42° from the anterior slope
for almost half the height of the shell. Outline of
peristome flat.
Type Material: Holotype SBMNH 236523, 5.26 x
3.77 x 3.47 mm (LxWxH); paratype SBMNH 236524,
2.66 x 1.71 x 1.55 mm (LxWxH).
Type Locality: Concepcion Bay methane seep area,
off Concepcion, Chile, Southeast Pacific Ocean, BV
Agor Vidal Gormaz, 36° 22' S; 73° 43' W, 846 m depth,
coll. & leg. Guillermo Guzman.
Etymology: Named in honor of Guillermo Guzman,
M useo del Mar, Universidad Arturo Prat, Iquique, Chile,
who donated the type material of this new species.
Remarks: This species lives in or near methane
hydrate rubble. The holotype had a portion of the body
remaining in the shell, although the head with radula
unfortunately was not available.
DISCUSSION
Seven species assigned to the genus Cornisepta have
been recorded in the Pacific Ocean; C. acuminata
Figures 1-7. Cornisepta guzmani new species 1. Holotype SBMNH 236523, 5.26 mm, lateral view. 2. Holotype, dorsal view.
3. Holotype ventral view. 4. Paratype SBMNH 236524, 2.66 mm, lateral view. 5. Paratype, dorsal view. 6. Paratype, ventral view.
7. Paratype, detail of sculpture. Scale bars = 1 mm.
J.-F. Araya and D.L. Geiger, 2013
Page 117
Table 1. Summary of shell characters of Comisepta species from the Pacific Ocean based on Chino (2009), Cowan (1969),
McLean and Geiger (1998), and this study. PF: position of foramen.
Watson, 1883 from the Western Atlantic (290-710 m)
is also included (Table 1); C. antarctica from the
Weddell sea, Antarctica, in 280-700 m depth and
the Bellinghausen Sea, Antarctica in a depth range
of 400-500 m (Aldea et ah, 2008), C f estiva from
northwest of the Three Kings Islands, New Zealand,
in 805 m, C. levinae from the summit of Volcano 6,
Eastern Pacific Rise (12° 44' N, 102° 33' W) in 1775 m
depth, C. monsfuji from the SE China Sea, south-
western Japan in 240-270 m, and from off Olango
Island, Philippines, in 60 m, C pacifica from Kiska,
Aleutian Islands, Alaska to southern California, in a
depth range of 440-880 m (McLean and Geiger, 1998),
C. soyoae from Sagami Bay, Japan in 120-170 m depth
and C. verenae from Axial Seamount, Juan de Fuca
Ridge (45° 56' S, 130° 04' W) in 1530 m depth. All
of the species have shelf to bathyal distributions, and
two of them (C. levinae and C. verenae) occur only at
hydrothermal vents (McLean and Geiger, 1998; Sasaki
et ah, 2010).
Comisepta guzmani has a shell similar to C. pacifica-
the new species differs in having a lower profile (66 vs.
76% L/H), a more concave posterior profile, a larger
angle between the septum and the anterior slope and
sparse pustules over the shell. Moreover, C. guzmani
has been found at cold seeps while C. pacifica has
not been associated with such habitats. Comisepta
antarctica and C. festiva differ from the new species
in their larger shells with height almost equal or even
greater than length (96%, 1 13% L/H, respectively), have
different sculpture of pustules (with t-shaped pus-
tules in C antarctica) that are more delicate than in
C. guzmani , and straighter anterior and posterior pro-
files. Comisepta levinae and C. verenae differ from
C. guzmani in the higher profile (67%, 81% L/H,
respectively), the more densely ornamented shell, and
the comparatively smaller and more neatly ordered
pustules, with C. verenae exhibiting a distinct sculp-
ture of chained pustules encircling the shell. Comisepta
monsfuji and C soyoae differ from the new species in
having smaller shells, higher profiles (C monsfuji has
a taller than wider shell), and more strongly orna-
mented shells, with pustules aligned in vertical rows
in C monsfuji and in oblique prosocline rows in
C soyoae. They also have bathymetry ranges shallower
than C. guzmani. Comisej)ta acuminata has much denser
pustules and a much taller shell.
So far, C. guzmani has the lowest shell profile, with
the most posteriorly placed foramen of all the Pacific
Comisepta species. It is also the second species to be
recognized living at a cold seep, after C. acuminata
(Watson, 1883), reported for the Sinn River delta meth-
ane seep (09° 02' N, 76° 02' W), Colombia in 500 m
depth (Gracia et al., 2011), and the first such fissurellid
species for Chile and the southeastern Pacific coast.
ACKNOWLEDGMENTS
Guillermo Guzman, Museo del Mar, Universidad
Arturo Prat, Chile, who collected the first specimen
of the new species. Arnold Muller, Leipzig University,
Germany, Dai Herbert, KwaZulu-Natal Museum, South
Africa, and Lindsey Groves, Los Angeles County
Museum of Natural History, California, USA, provided
literature and commented on the genus. Bruce Marshall,
Museum of New Zealand Te Papa Tongarewa, Wellington,
New Zealand and an anonymous reviewer critically
reviewed the manuscript.
LITERATURE CITED
Aldea, C., C. Olabarria, and |. Troneoso. 2008. Bathymetric
zonation and diversity gradient of gastropods and
bivalves in West Antarctica from the South Shetland
Islands to the Bellingshausen Sea Deep-Sea Research
Part I Oceanographic Research Papers 55(3): 350-368.
http://dx.doi.Org/10.1016/j.dsr.2007.12.002
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THE NAUTILUS, Vol. 127, No. 3
Beck, T, T. Metzger, and A. Freiwald. 2006. BIAS - Biodiversity
inventorial atlas of macrobenthic seamount animals. OASIS
Deliverable 25 Final Report: 1-124.
Chino, M. 2009. A new species of the genus Comisepta
McLean, 1998 [sic] (Gastropoda: Fissurellidae) from Japan.
Venus 68: 63-66.
Cowan, I.M. 1969. A new species of gastropod (Fissurellidae,
Fissurisepta) from the eastern North Pacific Ocean. The
Veliger 12: 24-26.
Crozier, M.A. 1966. New species and records of Mollusca
from off Three Kings Islands, New Zealand. Transac-
tions of the Royal Society of New Zealand, Zoology 8:
39-49.
Gracia, A., N. Rangel-Buitrago, and J. Sellanes. 2011. Methane
seep molluscs from the Sinu-San Jacinto fold belt in the
Caribbean Sea of Colombia. Journal of the Marine Bio-
logical Association of the United Kingdom 92: 1367-1377.
doi: 10.101 7/S00253 15411 00 1421
Habe, T. 1951. Fissurellidae in Japan. Illustrated catalogue of
die shells of Japan 17: 109-120, pi. 17.
Habe, T. 1964. Shells of the Western Pacific in color, Vol. II.
Osaka: Hoikusha, 233 pp., 66 pis.
McLean, J.H. and D.L. Geiger. 1998. New genera and spe-
cies having the Fissurisepta shell form, with a generic-
level phylogenetic analysis (Gastropoda: Fissurellidae).
Contributions in Science, Natural History Museum of
Los Angeles County 475: 1-32.
Miiller, A. 2011. First record of fossil Comisepta McLean 1998
[sic] from the North Sea Basin (Early Oligocene, Central
Germany) (Gastropoda Vetigastropoda: Fissurellidae).
Arehiv fur Molluskenkunde: International Journal of
Malacology, 140: 239-244.
Sasaki, T., A. Waren, Y. Kano, T. Okutani, and K. Fujikura.
2010. Gastropods lrom Recent hot vents and cold
seeps: systematic, morphology and life strategies. Topics
in Geobiology 33: 169-254.
THE NAUTILUS 127(3):119-124, 2013
Page 1 19
A new species of Lithasia (Gastropoda: Pleuroceridae)
from the Buffalo River, Tennessee, USA
Kussell L. Minton
Department ol Biology
University of Louisiana at Monroe
700 University Avenue
Monroe, LA 71209-0520 USA
[email protected]
ABSTRACT
A new species of pleurocerid snail, Lithasia buhala new spe-
cies, is described from the Buffalo River, Tennessee, USA. The
species is sister to Lithasia geniculata from the Duck River in
Tennessee, and can he separated from it both morphologically
and using mitochondrial DNA sequences. Lithasia buhala rep-
resents the first snail endemic to the Buffalo River watershed.
Additional Keywords: Lithasia geniculata. Duck River, COI
sequences
INTRODUCTION
Pleuroceridae is a family of gill-breathing, opereulate
snails found in North America that reaches its greatest
diversity in the southeastern United States (Burch and
Tottenham, 1980; Lydeard and Mayden, 1995; Lydeard
et ah, 1997; Strong and Frest, 2007). Stabilizing the
taxonomy of pleurocerids in general is desirable given
the state of freshwater mollusks taxonomy in North
America. Like many other non-marine invertebrate groups
(Lydeard et ah, 2004), pleurocerids are experiencing
declines in the number of species and individuals because
of anthropogenic activities (Bogan et ah, 1995; Lydeard
et ah, 1997), and because tax a with small geographical
ranges tend to be more highly imperiled (Jablonsky,
1982). For example, in the Mobile Basin, all species in
one genus ( Gi/rotoma ) and approximately 31 species of
other genera are presumed extinct (Bogan et ah, 1995).
Recent efforts have improved our understanding of the
family and its systematic^ (Lee et ah, 2006; Dillon and
Robinson, 2009; Strong and Kohler, 2009), though more
work is needed in terms of species delineation and
conservation (Perez and Minton, 2008).
The genus Lithasia is one of six extant genera in
Pleuroceridae. The majority of work treating the genus
is concentrated on the taxonomy (based on shell char-
acters) and synonymy of the nominal species that com-
prise the modern Lithasia (Tryon 1873; Goodrich, 1940;
Burch and Tottenham, 1980) using shell characters. One
group in need of revision includes those individuals cur-
rently recognized as Lithasia genicidata Haldeman, 1840.
In modern treatments (e.g. Burch, 1982), L. geniculata
is comprised of three subspecies, genicidata, fidiginosa
(Lea, 1841), and pinguis (Lea, 1852), occurring in the
Cumberland and Duck River drainages in Kentucky and
Tennessee. Each subspecies was originally described as
its own species (Tryon, 1873) and later synonymized.
Differences between taxa were based on conchological
features and where they occurred within a river system.
Lithasia g. pinguis was considered the headwaters
form, transitioning into the L. g. fidiginosa form mid-
river, then into the L g. geniculata form in the deeper
main stem. Minton and Lydeard (2003) suggested that
L genicidata represented multiple species based on
analyses of morphology and mitochondrial COI DNA
sequences, while Minton et al. (2008) showed that much
of the shell variation in this group was due to phenotypic
plasticity. This paper aims to clarify the taxonomy of
L. genicidata by describing a new species from within
tire species putative range.
MATERIALS AND METHODS
Specimens of the new species described here were
found in the collections of the Florida Museum of
Natural History, Gainesville, Florida (FLMNH), Museum
of Comparative Zoology, Cambridge, Massachusetts
(MCZ), and the Museum of Zoology, Ann Arbor,
Michigan (UMMZ). Vouchers were deposited at the
North Carolina Museum of Natural Sciences, Raleigh,
North Carolina (NCMS) and the University of Alabama
gastropod collection, Tuscaloosa, Alabama (UAG).
Taxonomy follows Turgeon et al. (1998; but see Dillon,
2011). Radulae were prepared according to Holznagel
(1998) and visualized on a Hitachi S2500 scanning elec-
tron microscope. The distribution map was generated in
DIVA-GIS 7.1.6 ( Hi j mans, 2009).
Cytochrome oxidase subunit I sequences from Minton
and Lydeard (2003) were combined with selected
Page 120
THE NAUTILUS, Vol. 127, No, 3
non -Lithasia sequences from Sides (2005) and an
unpublished Juga orickensis sequence from Gen Bank
(Appendix 1) as the outgroup. Eight additional
sequences from the new species were generated by
the author and were identical to those generated
previously (AF435747-8). All sequences were aligned
by eye and analyzed under maximum likelihood using
the CIPRES 2.0 version of RAxML 7.2.5 (Stamatakis
et ah, 2008) under the GTR+I+G model as suggested
by jModelTest (Posada, 2008). Branch support was
assessed using 100 non-parametric bootstrap replicates
in RAxML. To determine the degree of exclusive
ancestry (i.e. complete lineage divergence), the genea-
logical sorting index (g si; Cummings et ah, 2008) was
calculated using 10,000 replicate generations. Values lor
gsi range from zero to one, with one indicating mono-
pliyly and complete separation from other lineages;
a p-value indicates significance of the gsi. To date the
divergence of clades from one another, BEAST 1 .54
(Drummond and Rambaut, 2007) was used to esti-
mate times to most recent common ancestors (TMRCA)
for Lithasia geniculata. The GTR+I+G model was
used and anchored with the estimated pleurocerid-
semisulcospirid split at 90 MYA (Strong and Kohler,
2009). A Yule process tree prior was used for compar-
ing different species with an uncorrelated log-normal
relaxed clock hypothesis to allow for rate heterogeneity
in the tree (Drummond et ah, 2006). The analysis ran
for ten million states, logging every ten thousandth state.
BEAST results were visualized in Tracer 1.5 (Rambaut
and Drummond, 2009) and recorded as mean time ±
standard error.
RESULTS
Maximum likelihood analysis yielded a single tree
( — 6178.75, a=1.24, pinvar=0.55) with well-supported
relationships between species of Lithasia (Figure 1).
Snails identified as L. geniculata from the “Cumberland”
Duck 1
Duck2 (n = 3)
Duck3 (n = 3)
Duck4
duttonianal (n = 3)
duttoniana2
Duck 5 (n = 6)
Duck6
Duck7 (n=6)
c.f. jayana
Buffalol (n = 5)
Buffalo2 (n = 5)
armigeral (n = 3)
armigera2 (n = 3)
armigcra3 (n = 3)
armigera4 (n=6)
armigeraS
Red 1
Red2 (n = 6)
Garrison Fork
lima 1 (n = 3)
salebrosa 1 (n = 3)
verrucosal (n = 3)
Iima2 (n=3)
salebrosa
verrucosa2 (n = 3)
verrucosa3 (n = 2)
verrucosa4 (n = 2)
verrucosa5 (n = 2)
verrucosa6 (n = 2)
"E." obovata2
1 E. laqueata
* E. striatula
nthonyi 1
A. anthonyi2
A. anihonyi3
Le. praerosa 1
Le. praerosa2
Duck
} Buffalo
{Cumberland
Le. praerosaS
• E. bullula
1 E. foremani
^ E. hydeil
1 E. hydei2
1 Juga orickensis
Figure 1. Maximum likelihood tree showing relationships of Lithasia geniculata populations to other pleuroeerids. One indi-
vidual was sequenced unless otherwise noted. Values above branches are non-parametric bootstrap values >50%.
R.L. Minton, 2013
Page 121
123333444789999911111122233344444444455555556666666777778888888888
82579368024567812566935614801134899915778990011279034660113555578
81436927200891235105910476481306806367038348456970
Buf TCACACTTTGCTACTAGCAATCCCGGTTTTGTACCTACCTTCGTGGCCAAATTTQAAGTCAGTTGA
Duckl GT. . .T C.TA.T. .C.T. .GA. .TA.GTA.AT
Duck2 T GT . . . T C . TG . T . . C . T . . GA . . TA . GTA . AT
Jay . TA.TTCGG. .ACTAC GT . . . T C . TG . T . . C . T . . GA . .TA.GTA.AT
GarFk . T . T AT ATGGCT . TAACCCGTCGTT . G . TCCTTGTTTTCGTCCG . GTTATGTAC . T
Redl . T . T AT ATGGCT . . AACCCGT . GTTGG . TC . TTGTTTTCGTCCG . GTTATGTAC . T
Red2 AT . T AT ATGGCT . T AACCCGT . GTT . G . TCCTTGTTTTCGTCCG . GTTATGTAC . T
Figure 2. Nucleotide substitution patterns tor nominal Lithasia species from Tennessee used in this study. The single haplotype
found in the Buffalo River is used for reference. Position numbers (based on 890 bp sequence) are above the nucleotides. Duckl
and 2 represent two haplotypes shared by L. duttoniana and L geniculate from the Duck River. Redl and Red2 represent
two haplotypes shared by L. g eniculata from the Red River The single specimen of L cf jaijana from the Duck River possessed
a unique haplotype (Jay), as did the single L. g eniculata from Garrison Fork (GarFk). Complete locality information can be found
in Minton and Lydeard (2003).
(Red River plus Garrison Fork), Duck, and Buffalo
drainages each formed reciprocally monophyletic clades;
‘Cumberland’ samples were sister to L. armigera , while
Duck and Buffalo samples were sister to one another.
All individuals from the Buffalo River shared a single
unique haplotype (Figure 2). Analysis using g si showed
that each clade is genealogically separate from the others
(gsi = 0.91-0.99, p«0.001). The TMRCA for Duck
and Buffalo River Lithasia was dated at 4 million ±
28.000 years ago, with the new species coalescing at
274.000 ± 5,000 years ago.
SYSTEMATICS
Class Gastropoda Cuvier, 1791
Family Pleuroceridae Fischer, 1885
Genus Lithasia Haldeman, 1840
Lithasia bubala new species
Buffalo Rocksnail
(Figures 2-6)
Diagnosis: Farge shell compared to sympatric pleu-
rocerids, ovately conic, golden-brown in color, fre-
quently with posterior tubercles on body whorl, and
posterior callus on columella. Body reddish-orange with
black mottling. All sequenced individuals to date possess
a single unique COI mitotype (Figure 2).
Description: Shell large, thick, ovately conic, height
to 40 mm (Figure 3). Spire short and tapered, often
eroded. Body whorl smooth, often plicate toward the
suture, varying frequently with angular ridge on lower
half. Transverse growth lines apparent. Sutures par-
tially impressed, but regular and distinct. Aperture
fusiform, one-half shell length. Columella white, with
thick anterior and posterior calluses on parietal wall.
Shell golden brown, frequently with purple- brown bands.
Animal reddish-orange mottled with black. Tentacles
bluish, wider at base, shorter than snout length. Radular
rachidian tooth with nine triangular cusps, central cusp
broader and longer than others (Figure 4). Fateral tooth
broad, top concave, with three cusps. Interior cusp
sharp and narrow. Central cusp large, broad, and lance-
olate. External cusp short, pointed, incurved, one-fifth
length of central cusp (Figure 4). Inner marginal teeth
with six to eight denticles of equal size except for
outer denticle on each side. Outer marginal teeth with
Figures 3-5. 3. Lithasia bubala new species holotype
FMNH 297357. 16.9 mm. 4. Lithasia bubala new species
rachidian and lateral teeth 5. Lithasia bubala new species
marginal teeth.
Page 122
THE NAUTILUS, Vol. 127, No. 3
11 to 13 denticles, of equal size except for outer denticle
on each side (Figure 5).
Type Locality: Buffalo River at takeout for Heath’s
Canoe Rental, 1076 Highway 13 N, Lobelville, Perry
County, Tennessee, USA. Approximate coordinates:
35.772° N 87.784° W.
Type Material: Holotype FMNH 297357; Paratypes
UAG 395.
Other Material Examined (all from Buffalo River,
Tennessee): ANSP 178961, north of Riverside, Lewis
County; FLMNH 35957, 12 miles north of Waynesboro,
Wayne County; FLMNH 82131, above Linden, Perry
County; FLMNH 82169, 5 mi. N of Lobelville, Perry
County; FLMNH 82182, Beardstown, Perry County;
FLMNH 232610, 1.1 miles north of Napier, Lewis
County; FLMNH 232615, 0.7 mi. NW of Flat Woods,
Perry County; FLMNH 232616, 2.0 mi. SSW of
Lobelville, Perry County; MCZ 93695, above Linden,
Periy County; MCZ 93701, 5 mi. N of Lobelville,
Perry County; MCZ 93711, Beardstown, Perry County;
UAG 406, Buffalo River Camping and Canoeing,
0.7 miles from Highway 13, south of Interstate 40,
Humphreys County; UMMZ 53215, 5 mi. N of
Lobelville, Perry County; UMMZ 53216, 5 mi. N of
Lobelville, Perry County; UMMZ 53219, Beardstown,
Perry County; UMMZ 53224, above Linden, Perry
County; UMMZ 53225, Topsy Bridge, Wayne County;
UMMZ 63873, 0.5 miles above mouth, Humphreys
County; UMMZ 154173, north of Riverside, Lewis
County; UMMZ 154174, 7 mi. S of Linden, Perry County.
Comparison with Duck River L. geniculata: Given
their reciprocal monophyly, body color is the easiest
way to distinguish between the two species. In addi-
tion, there is no overlap in their geographic ranges. In
life, individuals of L. bubala had orange-red bodies,
while the Duck River L duttoniana , L. geniculata ,
and L. jayana used in this study had greenish-yellow
bodies. While useful in this distinction, no study has
documented variations in body color in pleurocerids,
so its systematic utility remains unknown. Also, the
tentacles of L bubala tend to be darker blue-green
and wider at the base than those of L. geniculata.
Again, the systematic utility of tentacle characters
remains unknown.
Etymology: From Latin bubalus , an antelope, buffalo,
or ox, in reference to its endemism in the Buffalo River.
Distribution: Limited to the Buffalo River, Tennessee
(Figure 6).
DISCUSSION
The analysis of Minton and Lydeard (2003) suggested that
Lithasia geniculata from the Buffalo River, Tennessee,
represented a species separate from individuals from the
Figure 6. Distribution map of Lithasia bubala in the
Buffalo River based on museum collections. Type locality is
indicated with a star.
Duck River. Eight additional sequences and subsequent
analysis confirmed this finding and highlighted the
need to describe Buffalo River L. geniculata as a unique
species. The delineation of Lithasia bubala satisfies mul-
tiple species concepts with its diagnostic body color
compared to sister species (morphological), mono-
phyly (phylogenetic [Baum and Donoghue, 1995]),
and complete genealogical separation as measured
by gsi (genealogical [Avise and Ball, 1990]). Taken
together, L bubala additionally satisfies the criteria
for the encompassing, non-diagnostic evolutionary
(Wiley and Mayden, 2000) and unified (de Queiroz,
2005) species concepts.
The sister relationship of Lithasia bubala to all
Duck River Lithasia (L. duttoniana, L. geniculata,
and L. jayana) was not entirely unexpected, as the
relationship was shown previously (Minton and Lydeard,
2003), and given that the Buffalo River is the largest
tributary to the Duck River. Additionally, the two rivers
partially share freshwater faunal assemblages (Ortmann,
1924; van der Schalie, 1973). The Red River and
Garrison Fork grouping supports the notion that fauna
from the Upper Duck drainage has a Cumberlandian
origin (van der Schalie, 1973). The present analysis
suggested a split between L. bubala and Duck River
Lithasia occurring around four million years ago, in
the middle of the Pliocene. Lithasia bubala appears to
have coalesced as a species 274,000 years ago in the
Pleistocene. The Pleistocene is often considered a period
of freshwater species diversification in the east-central
United States, and examples can be found in fish (Near
and Keck, 2005), crayfish (Crandall and Templeton,
1999), and mussels (Watters et al., 2009).
The Buffalo River belongs to the Western Highland
Rim eeoregion in the Eastern Highlands (USEPA,
2011), an area recognized as having a high diversity
of flora and fauna. The Heritage Program recognizes
R.L. Minton, 2013
Page 123
48 rare plant and animal species in the Buffalo River
drainage, including L. g eniculata (TDEC, 2005). While
the Buffalo River itself is not seen as an area of
high aquatic endemism, two endemic terrestrial bee-
tles ( Pseudanophthalmus Hesperus Barr, 1959 and
P. occidentalis Barr, 1959) and one endemic terrestrial
annelid ( Cambarincola leptadenus Holt, 1973) are
found in the drainage (NatureServe, 2013). Additionally,
one species in the freshwater mussel genus Villosa may
also be endemic to the Buffalo River (G.T. Watters, pers.
comm.). Based on NatureServe conservation status defi-
nitions, I recommend assigning L bubala a G3/S3 rank-
ing of vulnerable, given its restricted range and the
ongoing plight of freshwater taxa in the United States
( Johnson et ah, 2013).
ACKNOWLEDGMENTS
The molecular data was generated at the Johnson
Laboratory for Molecular Systematic^ at The University
of Alabama and the molecular genetics laboratory at
ULM. W. Holznagel and J. Nunley at UA assisted with
radulae preparation and micrographs, and R. Gundersen
provided some shell photographs. S. Ahlstedt, K.
Cummings, J. Garner, P. Johnson, D. Neely and others
provided specimens. Thanks to the curators and staff at
the various museums for access to and assistance with
their respective collections, and to A. Bogan, K. Perez,
and anonymous reviewers for helpful comments. Portions
of this work were supported by the National Science
Foundation, the Conchologists of America, The University
of Alabama, and a Howard Hughes Medical Institute
faculty development grant awarded through ULM.
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APPENDIX 1
NCBI accession numbers for sequences used in this
study. Minton and Lydeard (2003): AF435739 through
AF435786. Sides (2005): EU106485, EU106517, EU106524,
EU 106525, EU 106526, EU106543, EU106544, EU106545,
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EU106570, EU 106575. Campbell et al. (unpublished):
EF587013.
THE NAUTILUS 127(3): 125-129, 2013
Page 125
Description of Fusilaria garciai new genus, new species
( Gastropoda: F asciol ari i dae : F asciolari i n ae )
from the western Caribbean Sea
Martin Avery Snyder
Academy of Natural Sciences of Drexel University1 2
Philadelphia, USA
and
Museum National d’Histoire Naturelle"
Paris, FRANCE
[email protected]
ABSTRACT
A new fasciolariine species is described from the western
Caribbean Sea, and assigned to a new genus. Four specimens
of this large species were collected from lobster traps set at
200 m depth off Roatan Island, Honduras.
Additional Keywords: Fasciolaria tephrina, Honduras
INTRODUCTION
This paper describes a new fasciolariine species col-
lected by lobster fishermen working from Roatan Island,
Honduras. Crabbed shells of the new species were col-
lected in lobster traps set at 200 m depth off Punta
Cameron, at roughly 16° N, 84° W. A new genus is pro-
posed for this species. Specimens of Fasciolaria tephrina
de Souza, 2002, another deepwater fasciolariine, were
collected concurrently in traps set in the same area at
500 m, and color forms of this species are discussed.
MATERIALS AND METHODS
The study material was obtained from a fisherman at
Roatan Island, Honduras by Dr. Emilio F. Garcia in
December, 2012. All four shells were occupied by hermit
crabs and collected in lobster traps. Measurements were
made with a manual vernier caliper and recorded to the
nearest 0.1 mm. Examinations were carried out with a
Nikon 70594 stereomicroscope. The abbreviation ANSP
is used for the Academy of Natural Sciences of Drexel
University of Philadelphia.
1 Research Associate
2 Research Associate
SYSTEM ATICS
Family Fasciolariidae Gray, 1853
Subfamily Fasciolariinae Gray, 1853
Fusilaria new genus
Type Species: Fusilaria garciai new species
Diagnosis: Shell moderately large for subfamily;
protoconch large; siphonal process long. Prominent sharp
columellar folds; prominent parietal ridge. Continuous
and interrupted lirae present on inner side of outer lip.
Description: Adult shell moderately large for subfam-
ily, with large protoconch and high spire. Axial sculpture
of low ribs forming nodes on peripheral keel adapical
to the axial midpoint of each whorl. Spiral sculpture of
numerous low, fine cords. Two or three prominent col-
umellar folds present at entrance to siphonal canal with
as many as four weaker folds adapically, not always vis-
ible from outside. Siphonal canal somewhat twisted.
Parietal ridge prominent with sharp edge. Inner side of
outer lip with fine lirae, both continuous and inter-
rupted (beaded).
Remarks: Snyder, Vermeij and Lyons (2012) recog-
nized 14 genera in the subfamily Fasciolariinae. Fusilaria
differs from all these genera; we discuss in detail those
genera with Caribbean members. Three genera, all con-
fined to the southeastern United States, contain only
fossil members. Liochlamys Dali, 1889 and Terebraspira
Conrad, 1862 are quite unlike Fusilaria , with species
differing markedly in general shell morphology, but
Pliculofusus Snyder, Vermeij and Lyons, 2012 requires
comparison because that genus shares several charac-
ters with Fusilaria. Of the other 1 1 genera, only four
( Au rant ilaria Snyder, Vermeij and Lyons, 2012; Cinctura
Hollister, 1957; Fasciolaria Lamarck, 1799; and Triplofusus
Olsson and Harbison, 1953) have Recent members
Page 126
THE NAUTILUS, Vol. 127, No. 3
occurring in the Caribbean, and of these only Fasciolaria
has an amphi-Caribbean distribution. The other three
genera barely satisfy' the Caribbean criterion, Aurantilaria
extending northward only to the southeastern corner and
Cinctura and Triplofusus extending southward only to the
northwestern edge of the Caribbean. One additional genus,
Granolaria , has two living members in the eastern Pacific
but has five Caribbean fossil members ranging in age from
Early Pliocene to Early Pleistocene. These six genera, their
Recent Caribbean members, and differences between
each of those genera and Fusilaria are as follows:
Aurantilaria Snyder, Vermeij, and Lyons, 2012. Included
species: A. aurantiaca (Lamarck, 1816). Principally a
Brazilian species but ranges northward to the Gren-
adines with fossil records in Florida and the Domin-
ican Republic. Fusilaria differs by having spiral
cords neither prominent nor nodular, and by having
a sharply edged parietal ridge.
Cinctura Hollister, 1957. Included species: C. lilium
(Fischer von Waldheim, 1807). Principally found in
the western Gulf of Mexico but occurs across the
Yucatan Platform as far south as Cancun. Fusilaria
differs by having axial sculpture on early whorls and
body whorl, spiral cords on all whorls except upper
part of body whorl, and some beaded lirae on inner
side of outer lip. It also has a larger size.
Fasciolaria Lamarck, 1799. Included species: F. bullisi
Lyons, 1972; F hollisteri Weisbord, 1962; F. tephrina
Souza, 2002; F. tulipa (Linnaeus, 1758). Fusilaria
differs by having axial sculpture on early whorls and
body whorl, discontinuous or beaded lirae on inner
side of outer lip, a prominent sharply edged parietal
ridge, and sharp columellar folds.
Granolaria Snyder, Vermeij, and Lyons, 2012. Two
Recent species in tropical eastern Pacific but only
fossil members in Caribbean region. Fusilaria differs
by having a high spire, some beaded lirae on inner
side of outer lip, and a sharply edged parietal ridge.
Pliculofusus Snyder, Vermeij and Lyons, 2012. Contains
only fossil members from the late Oligocene to
Pleistocene of the Atlantic sector of the southeastern
United States. Fusilaria differs by having axial ribs
that are not prominent, a relatively much longer
canal, a thinner shell, spirally elongate nodes, and
two or three rather than four columellar folds.
Triplofusus Olsson and Harbison, 1953. Included spe-
cies: Triplofusus giganteus (Kiener, 1840). Princi-
pally southeastern United States and Gulf of
Mexico, barely intruding in Caribbean southward
to Cozumel; Neogene record of genus at Punta
Cavil an, northern Venezuela (Rutseh, 1934). Fusilaria
differs by having a very thin, lightweight shell,
spiral cords that are not prominent, and some
beaded lirae on inner side of outer lip.
Etymology: The name is Latinized, combining Fusinus
and Fasciolaria ■ the gender is feminine.
Remarks: Although the columellar plicae reveal that
this new genus is fasciolariine rather than fusinine, the
profile of the whorls is reminiscent of the genus Fusinus
Rafinesque, 1815, and it is this combination of features
of two subfamilies that justifies creation of the new
genus-level taxon. Two fusinine species that correspond
in appearance to Fusilaria are Fusinus galatheae Powell,
1967 from the Kermadec Islands and Fusinus genticus
(Iredale, 1936) from New Zealand (and possibly eastern
Australia). Fusinus galatheae has a similar geneml appear-
ance, but differs in that the canal is straight, the outer lip
is angled at the periphery, and the spiral sculpture is
more pronounced, spaced more widely, and proceeds
up the neck. F. genticus also appears similar but the early
axial ribs are more prominent and not suppressed on the
first 4-5 teleoconch whorls.
Comparison is also invited by other fasciolariine spe-
cies, namely Kilbumia scholvieni (Strebel, 1911) from
South Africa and Australaria tenuitesta Snyder, Vermeij
and Lyons, 2012. Kilbumia has a larger protoconch (up
to 5 mm) and short, smooth lirae on the inner side of the
outer lip. The entrance fold to the siphonal canal is much
more strongly expressed than the two columellar folds.
Australaria differs in several ways, but in others is
quite similar. The lirae on the inner side of the outer lip
are short and smooth. The most anterior fold of the
siphonal canal is much more firmly expressed than
the two columellar folds. In the remarks concerning the
genus Australaria Snyder, Vermeij and Lyons (2012: 53)
observed, “Although we cannot unambiguously sepa-
rate the South African Kilbumia from the Australian
Australaria on shell characters, we provisionally main-
tain them as distinct genera because of their likely
long separate histories.” A similar statement is probably
appropriate for Fusilaria and Australaria.
Fusilaria garciai new species
(Figures 1-13)
Description: Shell moderately large for subfamily,
elongate, fusiform, with high spire and long siphonal
process. Protoconch large, maximum diameter 3 mm,
height 2.5-3. 0 mm, with two bulbous cream-colored
whorls, additional quarter whorl with sharp thin unor-
namented axial riblets, joining teleoconch smoothly.
Teleoconch of 6-7 angulate whorls separated by suture,
overlaid by spiral cords at the base of the whorl, with
spiral keel connecting 11-12 spirally elongate, axially
compressed nodules; with half whorl of wider unorna-
mented ribs, extremely short, rounded axial ribs extend-
ing adapically from nodes on later whorls, weakening
abapically; fine, very low spiral cords on all early whorls,
evanescent on upper part of body whorl, with a few
similar cords at end of siphonal process, oblique to pro-
cess axis; about 19 cords below keel and 18 above on
penultimate whorl. Outer lip thin and damaged in all
M. A. Snyder, 2013
Page 127
Figures 1-9. Fusilaria garciai new species. 1-3. Holotype, 161.7 mm, ANSP 450735. 4-6. Paratype, 183.0 mm, ANSP 450736.
7-9. Paratype, 119.5 mm, Garcia collection.
Page 128
THE NAUTILUS, Vol. 127, No. 3
Figures 10-13. Fusilaria garciai new species. 10. Protoconch,
holotype, ANSP 450735. 11. Principal columellar folds, liolo-
type, ANSP 450735. 12. Parietal ridge, holotype, ANSP 450735.
13. Lirae on inner side of outer lip, paratype, ANSP 450736.
examined material; inner side of outer lip with about
60 lirae, a mix of continuous, discontinuous, and beaded,
stopping short of the lip by about 3 mm; abapical lirae
stronger and more coarsely beaded. Anterior-most fold
to siphonal canal keel -like, with one or two additional
similar folds adapical to it. In some specimens as many
as four additional weak folds present, some not visible
from outside. Shell color ivory with peach overtones;
nodes lighter on peripheral keel with light pumpkin
splotch on both sides (axially) of node, with pale orange-
brown axial color stripes emanating from the nodes
adapically and abapically on the body whorl. Operculum
and soft parts not available for study.
Type Material: Holotype ANSP 450735, 161.7 mm, in
lobster traps, depth 200 m. Paratypes: 183.0 mm, ANSP
450736; 119.5 mm (Emilio Garcia collection), all from
lobster traps at type locality.
Type Locality: Off Punta Cameron, Honduras, approxi-
mately 16° N, 84° W, depth 200 m.
Other Material Examined: 181.1 mm, same data as
holotype (Emilio Garcia collection).
Distribution: Known only from the type locality.
Etymology: The species name honors Emilio F. Garcia
of Lafayette, Louisiana. Long a student of the molluscan
fauna of the Gulf of Mexico and western Caribbean, he
obtained these shells, recognized them as undescribed,
and was kind enough to donate the holotype and a
paratype to ANSP.
DISCUSSION
Fusilaria garciai new species was collected in traps
set at depths of approximately 200 m. Another large
fasciolariine, Fasciolaria tephrina, was collected in the
same area in traps set at 500 m. Although thin, the shell
of F. garciai is strong. Repaired breaks on the last whorl
suggest some sort of predation. Fasciolaria tephrina has
a fairly wide distribution from Belize to Honduras,
Nicaragua, and the northern Colombian Islands south-
ward to San Andres (see Miloslavich et ah: 14, table S6).
It is not unreasonable to speculate that both of these
species are representative of a larger widespread deep-
water fauna now beginning to be explored. We expect
that Fusilaria garciai may have a wider range in the
western Caribbean than indicated by this material, and
it is possible that additional material will be discovered in
other parts of the Caribbean at similar depths.
Fasciolaria tephrina apparently occurs in two color
forms: finely banded (Figure 14) and splotched (Figure 15).
The specimen depicted in Figure 15 was collected
in the same area as Fusilaria garciai, but in traps set
at 500 m. This specimen is a heavier shell than the
specimen in Figure 14, and shows signs of predation.
It is remarkable that species this large are still being
described from the Caribbean.
Figures 14-15. Fasciolaria tephrina. 14. “Banded form,”
277 mm, ANSP 450737). 15. “Splotched form”, 224.75 mm,
Garcia collection.
M. A. Snyder, 2013
Page 129
ACKNOWLEDGMENTS
Geerat Vermeij (University of California, Davis) read
an early version of this paper and offered helpful
remarks. Pan! Callomon (ANSP) also offered a helpful
conversation. Amanda Lawless (ANSP) produced the
figures and composed the plates. William G. Lyons of
St. Petersburg, Florida and an anonymous reviewer offered
careful, insightful and helpful suggestions. Mr. Lyons
also generously offered additional pertinent information.
Finally, Jose H. Leal (The Bailey-Matthews Shell
Museum, Sanibel) proffered helpful advice.
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Taxonomy and phylogeny of western Atlantic Lucinidae:
new genus for Lucina costata d’Orbigny, 1846, a new species
of Ferrocina and neotype designation for Venus orbiculata
(Montagu, 1808)
John D. Taylor
Emily A. Glover
Suzanne T. Williams
Department ol Life Sciences
The Natural History Museum
London SWT 5BD
UNITED KINGDOM
[email protected]
[email protected]
[email protected]
ABSTRACT
A new genus, Clathrolucina Taylor and Glover, is proposed lor
the widely distributed western Atlantic species Lucina costata
d’Orbigny, 1846, which has previously been placed in Codakia ,
Ctena, and Parvilucina . Molecular evidence indicates a closer
relationship to Radiolucina and Lucinisca. A new species,
Ferrocina garciai Taylor and Glover, is described from sub-
merged pinnacles oil the coasts ol Louisiana and Alabama.
It is similar to Ferrocina species described from the western
Pacific and, in molecular analysis clusters, with Parvilucina
and Bathyaustriella . Unusual features of this species include
the large ctenidia and the lateral coiled pouches of the vis-
ceral mass. A neotype is designated for Lucina orbiculata
Montagu, 1808 aka Ctena orbiculata, an abundant shallow
water lucinid of the western Atlantic. The original type mate-
rial is missing, the type locality is erroneous and considerable
doubt exists about the identity of this species.
INTRODUCTION
M ore than 40 species of Lucinidae, well known for
their chemosymbiosis with sulphide-oxidizing bacteria
housed in the ctenidia (Taylor and Glover 2006; Heide
et ah, 2012), are recorded from the tropical— subtropical
western Atlantic, including deeper water species. Many
of the shallow water species are relatively well known
and have been the subject of important studies of the
symbiosis (e.g. Giere, 1985; Fisher and Hand, 1984;
Distel and Felbeck, 1987; Frenkiel and Moueza, 1995;
Frenkiel et ah, 1996; Gros, Frenkiel and Moueza,
1998; Gros et ah, 1998, 2012; Gros Liberge and Felbeck,
2003, Brissac et ah, 2009). Nevertheless, there are
unresolved taxonomic and nomenclatural problems even
amongst abundant lucinids of the region including their
phylogenetic relationships. Greater taxonomic refinement
may also be needed since molecular analyses of other
bivalves such as Arcidae and Mytilidae have revealed
that species previously regarded as widespread in the
western Atlantic are in fact complexes of genetically
distinct taxa (Lee and 6 Foighil, 2005; Marko and
Moran, 2009). From morphological evidence, species
complexes are suspected within Lucinidae in the Lucina
pensijlvanica group (Gibson Smith and Gibson Smith,
1982) and Ctena orbiculata group (Taylor et ah, 2011).
Additionally, offshore exploration at hydrocarbon seeps
has recovered several new Lucinidae for the region
(Taylor and Glover, 2009).
In this paper we have three objectives. Firstly, we
clarify, using molecular evidence, the phylogenetic
position of “ Parvilucina ” costata (d’Orbigny, 1846), an
abundant species that has been variously and confus-
ingly classified in five different genera. A new genus
is proposed for this species. Secondly, we describe
an unexpected new species, dredged from depths of
58—86 m off Alabama and Louisiana. This lucinid
is unlike any other from the western Atlantic and is
most similar to species of Ferrocina from the western
Pacific (Glover and Taylor, 2007; in press). Thirdly, we
designate a neotype for Venus orbiculata Montagu,
1808 (i.e. Ctena orbiculata) perhaps the most abun-
dant lucinid of the subtropical-tropical western Atlantic.
This is necessary because the type material is missing,
the original figure is equivocal, the type locality is in
eastern Scotland and there is likely a complex of mor-
phologically similar but genetically distinct species living
in the western Atlantic.
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THE NAUTILUS, Vol, 127, No. 4
MATERIALS AND METHODS
Molecular Methods: Two new sequences from the
28S rRNA nuclear gene and mitochondrial cytB genes
were obtained from a single specimen of Ferrocina
garciai (GenBank numbers: KF793275, KF793276).
Extraction of DNA, PCR methods, and sequence editing
were as in Taylor et al. (2011). The new sequences were
analyzed along with a subset of previously published
sequences primarily from Taylor et al. (2011), although
distant outgroups and multiple exemplars of species
were removed. The alignment of cytB was unambiguous
but 2SS required alignment with MAFFT (v 6.864; Katoh
et al. 2002; online: http://mafff.cbrc.jp/alignment/server/).
The G-INS-I option was used with 1PAM / k=2’
because the species were from two closely related sub-
families. Gap opening penalty was 1.5 with an offset
value of 0.1 as long gaps were not expected based
on previous analyses. A total of 1453 bp remained
in the alignment after Gblocks was used to remove
ambiguously aligned regions (97% of the original
1497 positions) (0.91b, Castresana, 2000; http://molevol
.cmima.csic.es/castresana/Gblocks_server.html; using
options for a less stringent selection).
A species tree was constructed using concatenated
sequences from both genes and Bayesian inference as
implemented in Mr Bayes (v. 3.2.1, Huelsenbeck and
Ronquist, 2001). Nucleotide substitution models for both
genes were GTR+I+G as determined by MrModelTest
(v2.1, |. Nylander, http://www.abc.se/~nylander). Bayesian
analyses and tests for stationarity were performed follow-
ing protocols in Williams (2012).
Abbreviations: NHMUK: The Natural History Museum,
London; MNHN: Museum National d’Histoire Naturelle,
Paris; PRI: Paleontological Research Institution, Ithaca,
New York; USNM: National Museum of Natural His-
tory, Washington; YPM: Peabody Museum, Yale Univer-
sity, New Haven; H: shell height; L: shell length; PI:
Protoconch I; PI I: Protoconch 2; SEM: scanning elec-
tron microscopy; T: tumidity.
RESULTS
1 . New Genus for Lucina costata
The lueinid species often called Ctena or Parvilucina
costata (d’Orbigny, 1846) is widely distributed in the
western Atlantic, ranging from Bermuda, North Carolina,
through the Caribbean and Central America to Rio de
Janeiro (Britton 1970; Sehweimanns and Felbeck, 1985;
Rios, 1994; Vokes and Vokes, 1984; Redfern, 2001;
Hauser et ah, 2007; Mikkelsen and Bieler, 2007). It is
often abundant in shallow-water seagrass beds (Jackson,
1972; 1973). Despite being a common and widely dis-
tributed species, its systematic position is uncertain. It
has been variously placed in the genera Codakia , Ctena ,
Jagonia , Parvilucina , and Lucina in taxonomic and bio-
logical literature. In the last major taxonomic revision
of western Atlantic species, Bretsky (1976) listed some
shell characters to differentiate L. costata from Ctena
but placed it questionably in Parvilucina .
In common with all other investigated Lueinidae,
Lucina costata possesses symbiotic sulphide-oxidizing
bacteria housed in the ctenidia. Details of ctenidial
structure and location of symbionts in the species were
given by Giere (1985) and the bacteria characterized
molecularly by Distel et al. (1994) (as Codakia costata
GenBank L25712). Several other anatomical features
of the species were described by Allen (1958).
Samples of L. costata from Guadeloupe and Boeas
del Toro, Panama, were included in a molecular analysis
of a wide range of lueinid species with broad taxonomic
coverage (Taylor et ah, 2011). The results (Figure 1)
indicate that it does not group with Codakia , Ctena ,
Lucina, or Parvilucina but instead forms a clade with
Radiolucina and Lucinisca species within the subfamily
Lueininae. We are therefore introducing a new genus
to accommodate Lucina costata and two fossil species.
SYSTEM ATICS
Family Lueinidae Fleming, 1828
Subfamily Lueininae Fleming, 1828
Clathrolucina new genus Taylor and Glover
Type Species: Lucina costata d’Orbigny, 1846 here
designated.
Diagnosis: Shell small, L to 15 mm, subcircular,
inflated. Sculpture of numerous radial ribs of varying
thickness, prominent ribs often bifurcated, crossed by
fine commarginal lamellae. Lunule heart shaped, short.
Hinge with two cardinal teeth and large anterior and
posterior lateral teeth in each valve. Anterior adductor
scar broad, medium long, ventrally detached from
pallial line for about !4 length. Inner shell margin
finely dentate.
Etymology: clathrus , Latin for basket and Lucina,
in reference to the basket weave appearance of the
external sculpture.
Remarks: Molecular analysis (Figure 1) shows that
Clathrolucina costata falls within the subfamily Lueininae
and forms a subclade with Radiolucina and Lucinisca
(Taylor et al., 2011) rather than its previous placements
within Ctena and Codakia based on morphology. There
are superficial shell characters in common with Ctena
Moreh, 1861 ( type species: Codakia (Jagonia) mexicana
Dali, 1901) including the bifurcating radial ribs and
posterior dorsal areas that lack radial sculpture. The
anterior adductor scar of Ctena is longer and more
widely detached from the pallia] line (Figure 25). Jagonia
Recluz, 1869 (type species: Venus ehumea Gmelin, 1791)
has been used for C. costata by earlier authors such as
Dali (1901) and Chavan (1937) but it is now considered a
junior synonym of Ctena (Bretsky, 1976). In more recent
publications, Clathrolucina costata has been classified
J. D. Taylor et ah, 2013
Page 133
100
100
100
Lucinoma borealis
Lucinoma aequizonata
98 1 r Lucinoma myriamae ANG
10(? Lucinoma myriamae NIG
100
Codakia orbicularis
Codakia rugifera
Codakia tigenna
1 00 I — Codakia interrupta
Codakia paytenorum LH
Codakia oavtenorum OK
Codakia
76
100[
100
H 1
ooL-j —
ioo1 —
100
jm
100
TJfena~or51culalFBU
Ctena orbiculata FK
Ctena chiquita
Ctena imbricatula
Ctena mexicana
Ctena eburnea
Ctena decussata CR
100' Ctena decussata FR
Epicodakia tatei
Ctena delicatula KK
Ctena sp. PAN
1 00U — Ctena bella OK
691 — Ctena bella M B
r:; TunafufIFlevukana~
CODAKIINAE
on*
Ctena
100
Divalinga bardwelli
Divalmga quadrisutcata
Discolucina virgmea
— Lepidolucina venusta
Lucina adansoni
r Lucina pensylvamca FK
100
Lucina pensylvamca BOC
Lucina pensylvamca GDL
Lucina pensylvamca VEN
1 Qc\ Clathrolucina restate GDI '
P *£lathwlucimM0s.talaB,O.£„
100
9pj — Radiolucina amianta
100
100
H '
5oi_r
ioo1—
06
99
100
Radiolucina cancellaris
Lucinisca fenestrata
Lucmisca centrifuga
Lucinisca nassula
Rasta lamyi
Stewartia floridana
Pillucina australis
Pillucma sp PAN
Wallucina assimilis
— Chavania sp. MB
Chavania striata
Lucina
Clathrolucina
Radiolucina
Lucinisca
82
oo
umr:
1001-
LUCININAE
53
Lucinella divancata
Loripes clausus
100 r Loripes lucmahs UK
93
78
1 8S, 28S & cytB
100
1 Loripes lucinalis FR
— Troendleina cf musculator
Divaricella irpex
■ Bathyaustriella thionipta
100
0.1
Parvilucina creneila
— —Eamiucina pectinella
Austriella corrugata DMP
Parvilucina
■ Austriella corrugata SIN
1 0Oi — Indoaustriella plicifera
Indoaustriella lamprelli
■ Indoaustriella scartatoi
1 Indoaustriella dalli
(3Qj Cardiolucina quadrata
n_J Cardiolucina australopilula
■w* Cardiolucina semperiana
Cardiolucina new species
Cardiolucina pisiformis
Cardiolucina siquijorensis
Figure 1. Molecular phylogeny of Lucinidae produced by Bayesian analysis of concatenated sequences of 18S rRNA, 28S rRNA,
and cytochrome b genes (adapted from inset in figure 4 from Taylor et ah, 2011). The position of Clathrolucina costata is
highlighted, as are the genera where it has previously been assigned. Subfamily names are indicated on the right. Further details
given by Taylor et al. (201 1).
as a species of Parvilucina Dali, 1901 (type species:
Lucina tenuisculpta Carpenter, 1864, from the north-
eastern Pacific). However, in the molecular analysis,
although C. costata and two western Atlantic Parvilucina
species occur in the same major lucinid clade (subfamily
Lucininae; Taylor et ah, 201 1), they are not closely
related. Parvilucina species have fine radial ribs that
do not bifurcate, an extremely short anterior adductor
Page 134
THE NAUTILUS, Vol. 127, No. 4
sear detached for only about Va of length (Figure 26),
and the shell margin is finely denticulate. Clathrolucina
groups with Lucinisca Dali, 1901 (type species: Lucina
nassula Conrad, 1846) and Radiolucina Britton, 1972
(type species: Phacoides (Bellucina) amiantus Dali, 1901)
from the western Atlantic. Lucinisca has fine, non-
birfurcating radial ribs, crossed by low, thin commar-
ginal lamellae forming raised scales where they cross
the radial ribs. Radial ribs project beyond the ventral
shell margin as denticles. The lunule in the type species
is short and deeply inset and the anterior adductor
scar is ventrally detached for about Vz of its length
(Figure 24). The three living species of Radiolucina were
recently reviewed by Garfinkle (2012); Radiolucina
amianta has about 9— 1 1 prominent radial ribs with
deep interspaces, a short heart-shaped lunule with the
anterior adductor scar detached for about Vz of length
(Figure 23). The inner shell margin is coarsely den-
ticulate. Dali (1901) originally placed R. amianta in a
subgenus Bellucina (= Cardiolucina Sacco, 1901) but
Britton’s (1972) recognition of Radiolucina is cor-
roborated by molecular results in which Radiolucina
amianta and R. cancellaris are distantly separated from
Cardiolucina species (Taylor et al , 2011).
Included Species: Codakia (Jagonia) vendryesi Dali,
1903 from the Pliocene, Bowden Formation, Jamaica
(3 syntypes USNM 135720; see Woodring 1925:
109 pi. 14, figs. 1-4.) and Codakia (Jagonia)
umhonicostata Weisbord (1964 : 234, pi. 31, figs 7-8)
from the Pliocene of Venezuela (holotype PRI 1519a).
Clathrolucina costata (d’Orbigny, 1846)
(Figures 2-22)
Lucina costata d’Orbigny, 1846: 586; 1853: 296, pi. 27,
figs 40-42.
Lucina antillarum Reeve, 1850: pi. 10, fig. 37.
Lucina textilis Philippi, 1850: 104, pi. 2, fig. 7.
Lucina nux Verrill and Bush, 1900: 518, pi. 58, figs 12, 13.
Codakia ( Jagonia ) costata. — Dali, 1901: 800; Lamy, 1920:
262-263).
Jagonia (Jagonia) costata. — Chavan, 1937: 260.
Codakia (Codakia) costata. — Abbott, 1954: 390; Vokes
and Vokes, 1984: 38, pi. 38, fig. 5.
Codakia costata. — Allen, 1958: 426, fig. 64; Warmke
and Abbott, 1961:178; Abbott, 1974: 460, fig. 5298;
Rios, 1994: 253, pi. 87, fig. 1236.
Parvilucina ( Parvilucina ) costata. — Britton, 1972: fig. 5.
Lucina (Parvilucina?) costata. — Bretsky, 1976: 263, pi. 28,
figs 1-5.
Parvilucina costata. — Redfern, 2001: pi. 91, figs. 887
A, B; Jensen and Pearce, 2009: 330; Turgeon et ah,
2009: 727.
“Parvilucina” costata. — Mikkelsen and Bieler, 2007: 234.
Ctena orbiculata . — Tunnell et ah, 2010: 341 and figure,
non Montagu, 1808.
Figures 2-9. Type material ol Clathrolucina costata. 2-3. Exterior and interior of the holotype of Lucina costata. Rio de Janeiro,
Brazil, NHMUK 1854.12.4.765, L= 12.8 mm. 4-5. Paratype of L costata, exterior and interior of left valve, St. Thomas, NHMUK
1854.10.4.564 L = 8.9 mm. 6-8. Syntype of Lucina antillarum NHMUK1963192-3, exterior ol left valve and interior of left and
right valves L = 12.0 mm. 9. Holotype Lucina nux Bermuda, YPM 8760, exterior of right valve. L = 7 mm.
J. D. Taylor et al„ 2013
Page 135
Figures 10-21. Clathrolucina costata from Boeas del Toro, Panama, NHMUK 20130562 10. Exterior of left valve, L — 8.1 mm.
11. Dorsal view ol shell in Figure 10. 12—13. Exterior and interior of right valve, L = 8.3 mm. 14—15. Exterior and interior
of left valve, L = 8.3 mm. 16. Detail of sculpture. Scale bar = 1.0 mm. 17-18. Detail ol hinge teeth of left and right valves.
Scale bar = 1.0 mm. 19. Detail of anterior adductor muscle scar Scale bar = 2 mm 20. Protoconch, note mieropunctae of
post-settlement shell. Scale bar = 50 pm. 21. Microsculpture. Scale bar = 10 pm.
Page 136
THE NAUTILUS, Vol. 127, No. 4
22 .Clathrolucina
costata
23. Radiolucina
ami ant a
24. Lucinisca
nassula
25. Ctena
mexicana
26. Parvilucina
tenuisculpta
Figures 22-26. Outline drawings of anterior adductor muscle scars. Traced from digital images.
Types: Lucina costata , holotype, NHMUK1854. 12.4.765,
one left valve, L = 12.8 min, H 12.7 mm, T = 4.5
(Figures 2-3); paratypes NIIMUK 1854.10.4.564,
St Thomas, US Virgin Islands, 8 valves (4R, 4L); Gray
Cat No. 497 (Figures 4-5); Lucina ant ilia mm , 3 whole
syntypes, NHMUK 1963192-3 (Figures 6-8), L = 12.0 mm,
15.1 mm, 12.4 mm, St. Johns, Antigua; Lucina nux
holotype, YPM 8760, single right valve, L = 7 mm,
H = 8 mm, T = 3 mm (Figure 9).
Type Locality': d’Orbigny (1846: 586) cited Baie de Rio
Janeiro near St Christophe and gave other localities -
Antilles and Cuba.
Description: Shell small (L to 15.2, H to 14.3 mm),
subcircular (H/L=0.95, n = 10) moderately inflated,
posterior margin slightly truncated, umbones promi-
nent. Sculpture of numerous (ca. 35) rounded radial
ribs, varying in width, that occur in bundles of two
to three ribs, sometimes bifurcated ventral to umbo
(Figure 16). Ribs crossed by low, regularly spaced corn-
marginal lamellae. Posterior dorsal area without radial
ribs but with more prominent commarginal lamellae,
posterior dorsal margin with two low radial ribs with
raised scales. Anterior dorsal area marked by lower
and broader radial ribs. Lunule short, heart shaped,
smooth, slightly asymmetrical with right side larger.
Ligament, external, short, set in shallow groove. Micro-
sculpture of rows of shallow pits (Figure 21) or micro-
punctate in juvenile shell (Figure 20). Protoconch
(Figure 20): PI = 121 pm, PI + PII = 146 pm, PII
with 5-8 growth increments. Hinge (Figures 17-18);
LV with 2 cardinal teeth, the anterior larger, promi-
nent anterior and posterior lateral teeth; RV with
2 cardinal teeth, the posterior much larger and slightly
bifid, anterior and posterior laterals prominent. Anterior
adductor muscle scar broad (Figures 19, 22), medium-
long, with rounded ventral tip, ventrally detached from
pallia] line for about Vi of length and diverging at an
angle of 5-10°. Posterior adductor scar reniform. Pallial
blood vessel track visible. Pallial line entire. Inner shell
margin finely denticulate. Colour yellowish-white exter-
nally, white internally.
Distribution: Western Atlantic and throughout
Caribbean from North Carolina to Santa Catarina, Brazil
(Britton, 1970; Rios, 1994).
Remarks: The holotype is a worn shell in poor condi-
tion but has the features of the species. Other d’Orbigny
specimens in NHMUK from St. Thomas anti regarded
as paratypes are worn but in better condition. No local-
ity was given for Lucina textilis Philippi, 1850 and the
type has not been located. Syntypes of Lucina antillarum
Reeve, 1850 are in good condition anti one is figured
(Figures 6-8). Lucina nux Verrill and Bush, 1900
described from Bermuda (Figure 9) is regarded as a
higher than long shape variant of C. costata.
2. A New Species from the Northern Gulf of Mexico
This unusual species, a surprising addition to the shal-
low water bivalve fauna of the Gulf of Mexico sent
to us by Emilio Garcia, was dredged from submarine
pinnacles on the Alabama and Louisiana shelf.
SYSTEMATICS
Subfamily Lucininae Fleming, 1828
Ferrocina Glover and Taylor, 2007
Type species: Ferrocina multiradiata Glover and
Taylor, 2007.
Diagnosis: Shell to 20 mm, thin, subovate, posteriorly
truncate, sculpture of numerous fine to indistinct radial
ribs crossed by fine commarginal threads. Hinge plate
thin, small single cardinal tooth in RV, two in LV, lateral
teeth small or absent. Anterior adductor scar short,
detached for 1/3 of length. Interior shell margin coarsely
to finely dentate. Color bright to pale orange or rusty
J. D. Taylor et al., 2013
Page 137
red-brown, external color paler, often blotchy, stronger
in umbonal area.
Included species: Ferrocirw multiradiata, (Vanuatu,
Fiji, New Caledonia): Ferrocina new species Glover
and Taylor (in press), Philippines; and F. brunei from
off Brunei (Taylor and Glover, 2013).
Remarks: This distinctive, rare, and highly colored
genus with line radial ribs was originally described from
Fiji, New Caledonia, and Vanuatu with other species
known from the Philippines and Brunei.
Ferrocina garciai Taylor and Glover new species
(Figures 27-44)
Type Material: Holotype (Figures 27-34), USNM
1227857, L = 19.7 mm, II - 19.0 mm, T = 5.2 mm,
from type locality; Paratype 1 (Figures 35-37), USNM
1227858, L = 13.9 mm, H = 11.7 mm, T = 3.5, off
Alabama, 29° 15.681" N, 88° 20.237" W, 78-86 m,
dredged by R/V Pelican, 27 August 2012; Paratype 2
(Figures 38-39), Emilio Garcia collection 30783,
L = 11.7 mm, from type locality.
Type Locality: USA, northern Gull of Mexico, off
Louisiana, 27°56.835' N, 92°01.464' W, 58 m, rubble,
dredged by R/V Pelican, 26 August 2012.
Description: Small, L = 14.0 mm, II = 11.7 mm,
subcircular, slightly longer than high, less mature
paratypes more elongate with posterior margin trun-
cate; thin-shelled, moderately inflated, T/L = 0.26.
Shallow posterior sinus and posterior dorsal area.
Sculpture of numerous fine, low, radial ribs that
become obscure on ventral parts of shell, commarginal
sculpture of growth lines and halts. Posterior dorsal area
without radials. Protoconch (Figure 32): PI =113 pm,
PI+PII 160 pm, PII with many growth increments.
Lunule long, slightly asymmetric, larger in RV. Ligament
long, set in shallow groove. Hinge narrow: RV with
single cardinal tooth and irregular anterior folds; tiny
posterior lateral teeth present on smaller paratype; LV
with two cardinal teeth and anterior folds but poste-
rior laterals present on paratype. Anterior adductor
scar medium long, ventrally detached from pallia! line
for Vi of length at angle of ca. 10° (Figure 40). Pallial
line entire, dorsally lobate. Posterior adductor scar
ovoid. Impression of pallial blood vessel prominent,
broad, terminating ventral to anterior adductor scar
(Figures 40-41). Shell margin finely dentate and
linearly grooved. Exterior color pale pink-orange to
off white, umbonal area darker. Interior colour uni-
formly bright apricot-orange, extreme shell margin
creamy white.
Anatomy: Veiy large, thick ctenidal demibranchs
occupying most of mantle cavity (Figure 42). Mantle
gills absent. Foot large, fat, with short heel. Body wall
immediately anterior to the foot extended on each side
into posteriorly directed “ramshorn-like” coiled projec-
tion (Figures 43-44). Labial palps as short ridges.
Rectum passing dorsally and posterior to posterior
adductor muscle. Posterior exhalant aperture with an
inverted tube, inhalant aperture edged with papillae on
middle mantle fold. Posterior mantle fusion short.
Phylogenetic Position: The results of a molecular
analysis using sequences from two genes, 28S rRNA
and cytochrome B (Figure 45), show the position of
Ferrocina garciai (tissue from holotype) in relation to
a diverse range of lucinid species from the subfamilies
Codakiinae and Lucininae. Other more distantly related
lucinid subfamilies of Pegophvseminae, Leueosphaerinae,
and Myrteinae (see Taylor et al., 2011) were excluded
from this analysis. The results show that Ferrocina
parciai belongs in the Lucininae and that it is sister to
two species of Parvilucina from the western Atlantic
(PP=100%). This subclade is sister to Bretskya sp and
Bathyaustriella thionipta (PP=10()%). The four genera
are in turn a sister clade to Troendleina and Epicodakia
falklandica although support is lower (PP=82%).
Bretskya (type species: B scapula Glover and Taylor,
2007) is an Indo-Pacific genus of irregularly shaped
lucinids that are strongly associated with sunken wood
(Glover and Taylor, 2007; in press). Bathyaustriella
thionipta is recorded from a 500 m deep hydrothermal
vent on the Kermadec Ridge off New Zealand (Glover
et al., 2004). Troendleina (type species: T rnarquesana
Cosel and Bouehet, 2008) includes several deeper
water species (150-800 m) from the Pacific Ocean
(Cosel and Bouehet, 2008, Glover and Taylor, in
press). “ Epicodakia ” falklandica Dell, 1964 from the
southern Atlantic occurs at depths of 100 — 400 m but the
molecular results show that it should not be classified
in the Codakiinae as Epicodakia but in the Lucininae
with uncertain generic placement. The two Parvilucina
species analyzed are known from shallow water habitats
(1-100 m) in the western Atlantic.
Distribution and Habitat: The three individuals of
Ferrocina garciai were dredged from submerged cal-
careous pinnacles and banks at depths of 58-86 m off
Louisiana and Alabama. These pinnacles occur along
the northern Gulf of Mexico shelf from Alabama to
Texas and form islands of hard substrate surrounded
by mud (Parker and Curray, 1956). The unusual mol-
luscan faunas of the pinnacles, including many endemic
species and new records for the Gulf, have been docu-
mented by Garcia (2000; 2008) and Garcia and Lee
(2002; 2013).
Remarks: Although presently known from only three
specimens, Ferrocina garciai is a highly distinctive
species and cannot be confused with any other lucinid
from the western Atlantic. It differs from the type
species Ferrocina multiradiata (Vanuatu, Fiji, New
Caledonia) (Figures 46-47), depth range 80-400 m,
in having finer, less prominent radial ribs and differs
in shape and ribbing from a new Ferrocina species
from the Philippines (Glover and Taylor, in press). The
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THE NAUTILUS, Vol. 127, No. 4
Figures 27-39. Ferrocina garciai. 27-28. Exterior of right and left valves of holotype, L = 19.7 mm. 29. Dorsal view of holotype.
30-31. Interior of right and left valves of holotype. 32. Protoconch of holotype, boundary between PI and PII indicated with
arrow. Scale bar = 50 pin. 33-34. Details of hinge of left and right valves of holotype (SEM). Scale bar = 2 mm. 35. Exterior
of left valve of paratype 1. L = 13.9 mm. 36-37. Interior of left and right valves of paratype 1. 38-39. Exterior and interior
of left valve of paratype 2, I, = 11.7 mm.
J. D. Taylor et al., 2013
Page 139
Figures 40-41. Outline drawings of shell interiors of Ferrocina 40. Holotype. 41. Paratype 1.
Figures 42-44. Ferrocina garciai, body of holotype. 42. Left side with mantle removed showing large, thick, left ctenidial
demibranch. 43. bight side with mantle and part of right demibranch removed showing foot and coiled lateral visceral extension.
44. Detail of ramshorn visceral extension Abbreviations: aa, anterior adductor muscle; ex, exhalant aperture; f, foot; h, heel of
foot; hg> hindgut; ia, inhalant aperture; it, invertible tube of posterior exhalant aperture; k, kidney; Id, left demibranch; Ip, labial
palps; m, mantle margin; pa, posterior adductor muscle; r, rectum; rd, right demibranch; ve, visceral ‘ramshorn’ extension.
Page 140
THE NAUTILUS, Vol. 127, No. 4
100
97
100
28S & cytB
n=65
0.2
rLu
K
qqI —
99
100
Lucinoma borealis
Lucinoma kazani
Lucinoma aequizonata
Lucinoma myriamae
Codakia rugifera
I 99 Q
100
Codakia tigerina
i oo | — Codakia interrupta
l — Codakia paytenorum OK
I00r Ctena orbiculata FK
Ctena orbiculata BMD
Ctena chiquita
oslIOOr Ctena imbricatula
100
Ctena mexicana
□ Ctena eburnea
Ctena decussata CR
- Epicodakia tatei
Ctena delicatula KK
Ctena sp PAN
Ctena bella MB
Ctena bella OK
■ Funafutia levukana
59
51
97. Lepidolucina venusta
' Discolucina virginea
77 | Divalinga bardwelli
Divalinga quadnsulcata
Lucina adansoni
100 r Lucina pensylvanica FK
' Lucina pensylvanica GD
98
100
76
381 L
rt-r'
100*—
59l4 —
931 —
100
82
100
Lucinisca fenestrata
Lucinisca centrifuga
Lucinisca nassula
Clathrolucina costata BOC
Radiolucina amianta
Radiolucina cancellaris
— Troendleina of musculator
Epicodakia falklandica
100
67
100
— Bretskya sp VAN
Bathyaustriella thiompta
Ferrocina garciai
100 i — Parvilucina crenella
■cr
68
91
93
100
Parvilucina pectmella
■ Rasta lamyi
Stewartia floridana
1 00 i Pillucma australis
Pillucma pisidium
— Wallucina assimilis
Chavania sp. MB
Chavania striata
100 | Pillucma vietnamica
' Pillucina sp PAN
100
63
90
99
Loripes clausus
Lucinella divaricata
■ Loripes lucinalis UK
89
100
Divaricella irpex
Austriella corrugata DMP
100
r~A
1 100
-Q'
100*-
100
fTT
100*-
Indoaustriella scarlatoi
Indoaustriella plicifera
Indoaustriella lamprelli
Cardiolucina new species
Cardiolucina pisiformis
Cardiolucina siquijorensis
70 _
64
100
_| — Cardiolucina quadrata PH
Cardiolucina quadrata MAD
— Cardiolucina australopilula
Cardiolucina semperiana
Figure 45. Molecular phylogeny of Lucinidae produced by Bayesian analysis of concatenated sequences of 28S rRNA, and
cytochrome h genes including new sequences for Ferrocina garciai. Only lucinids from subfamilies Codakiinae and Lucininae
included in the analysis. Locality and GenBank details of other lucinid taxa given in Taylor et al. (2011). GenBank numbers for
Ferrocina garciai : 28S rRNA KF793276, cytochrome b KF793275.
J. D. Taylor et ah, 2013
Page 141
Figures 46-49. Other species discussed under the “Remarks” for Ferrolucina garciai. 46—47. Ferrocina multiradiata Glover and
Taylor, 2007, holotype Vanuatu, 90-200 m, MNHN, exterior and interior of left valve, L = 11.2 mm. 48-49. Syntype of Codakia
(Jagonia) pertenera Dali, 1903, exterior and interior of left valve, L = 35 mm, USNM 135716, Pliocene, Bowden Formation, Jamaica.
juvenile paratype shells of F. garciai have stronger
radial ribs and a shape more similar to F. multiradiata .
The curled body extension of F garciai is similar to
that described from Bathyastriella thionipta, a species
from a hydrothermal vent on the Kermadec Ridge,
New Zealand (Glover et al., 2004). Bathyaustriella
thionipta also has very large enveloping ctenidial
demibranchs and a papillate inhalant aperture. We
have no anatomical data for the type1 species Ferrocina
multiradiata but the recently described, small, Ferrocina
Brunei from off Brunei, Borneo (Taylor and Glover,
2013) also has the curled body extension but the
demibranchs are relatively small. Troendleina cf.
musculator from the Philippines has the visceral mass
extended into a ventral lobe and Parvilucina crenella has
prominent lateral pouches to the visceral mass (unpub-
lished observations) that may be homologous with the
curled body extensions of Ferrocina. Visceral extensions
may be an apomorphy of this subclade of Lucininae.
Ferrocina garciai has many shell features similar to
the Ferrocina species known from the tropical Pacific,
but it is a surprising addition to the western Atlantic
fauna because the western Pacific species are also
rare and known from only a few specimens. However,
diere is a probable fossil antecedent in the Caribbean
(Figures 48—49). This is Codakia (Jagonia) pertenera
(Dali, 1903: 1347 pi. 51, fig. 4., four syntypes, USNM
135716) from the Late Pliocene, Bowden Formation
of Jamaica, later reclassified as Myrtaea ( Myrteopsis )
pertenera by Woodring (1925: 113, pi. 14, figs 13-16).
Subsequently, Bretsky (1976: 284) discussed the spe-
cies, apparently not recorded from mainland North
America and supported an assignment to Ctena. It
is larger than F garciai (L = 35 mm) and the life
color is unknown but it is similar in shape, denti-
tion, and anterior adductor scar, with weak radial
ribs on the dorsal part of the shell, and (accord-
ing to Woodring) the inner shell margin is sometimes
slightly fluted.
Etymology: Named for Emilio Garcia who collected
this distinctive species.
Page 142
THE NAUTILUS, Vol. 127, No. 4
3. Neotype for Venus orbiculata Montage, 1808
(= Ctena orbiculata)
Ctena orbiculata (Montagu, 180S) is one of the most
abundant shallow water bivalves of the western Atlantic
(Britton, 1970; Bretsky, 1976; Jackson, 1972, 1973) with
a reported geographical range from Bermuda to Brazil.
Venus orbiculata was described by Montagu (1808: 42)
from a locality on the east coast of Scotland: “Found
on the shore near Dunbar bv Mr Laskey.” No Ctena spe-
cies are known to live around Scotland or the British Isles
and the record is regarded as adventitious despite find-
ings of two shells of the Mediterranean Ctena decussata
(da Costa) from the Seilly Islands and Cornwall - see
McMillan, 1971 How the species came to be recorded
from Dunbar is a mystery. Dunbar is on the eastern
coast of Scotland, to the east of Edinburgh and in
the late 18th and 19th centuries was a busy fishing
port and for a while, a whaling port. Western Atlantic
mollusc species occasionally wash up on British shores
but they usually occur on the west coast (Oliver et al.,
2009). The specimen could have been dumped from
ship ballast or there was an accidental confusion of
localities. Cleevely (1995: 391) noted that collectors
such as Laskey from whom Montagu obtained shells
are known to have been less than scrupulous over the
provenance of their specimens.
Original Description from Montagu (1808 p. 42):
“ Shell white, orbicular depressed , and cancellated: the
umbo remarkably small , beneath which is a minute cor-
diform depression. Inside white; margin plain, teeth,
two primary approximate, and one remote standing
transverse; the margin where the lateral tooth is placed
projects into an angle. Diameter five eighths of an inch
(approx. 16 mm). The shell has probably been con-
founded with Venus tigerina, but it differs somewhat
in contour, is not so flat, more orbicular, and more
coarsely decussated, and the lateral tooth is nwre
remote than in tigerina. Found on the shore near
Dunbar, by Mr Laskey.’'
Since Dali (1901) the name has been accepted for
the western Atlantic species and the type locality of
Dunbar, Scotland, was regarded as erroneous. No type
material exists among Montagu specimens either at the
Boyal Albert Memorial Museum, Exeter, or The Natu-
ral History Museum, London. Furthermore, Montagus
figure (see Figure 50) is equivocal and cannot with
confidence be assigned to any Ctena species. Never-
theless, Lamy (1920) listed Venus orbiculata Montagu
as a synonym of the western African species now
known as Ctena ebumea (Gmelin, 1791). Other names
introduced for western Atlantic Ctena species ( Lucina
occidentals Beeve, 1850; Lucina imbricatula C. B.
Adams, 1845 and variety names for C. orbiculata
var. filiata and var. recurvata introduced by Dali
(1901)), have usually been placed in synonymy with
C. orbiculata (Britton, 1971; Bretsky, 1976). The name
Ctena orbiculata has become firmly entrenched in
publications concerning western Atlantic mollusks (e.g.
Warmke and Abbott, 1961; Abbott, 1954, 1974; Bretsky,
1976; Yokes and Vokes, 1984; Bedfern, 2001; Mikkelsen
anil Bieler, 2007) and variously classified under other
generic and subgeneric combinations of Codakia
( Jagonia ), Codakia (Ctena), or Codakia (Codakia).
An additional problem arising from ongoing molecu-
lar analyses, including our own results (Barnes and
Weigt. 1998; Taylor et al., 201 1), is that Ctena orbiculata
is likely a complex of morphologically similar species
as have been documented for other western Atlantic
bivalves. Certainly, our own results separate Ctena
orbiculata from the Florida Keys and specimens from
Bocas, Panama that on shell characters resemble
the type of Lucina imbricatula C. B. Adams, 1845, a
species that is usually synonymized with C. orbiculata
(e.g. Bretsky, 1976).
Ctena orbiculata in its varying name combinations
is the subject of continuing investigations into the
biology of the bacterial symbiosis, host-symbiont rela-
tionships (e.g. Giere, 1985; Schweimanns and Felbeck,
1985; Gros et al., 1998; Gros and Felbeck, 2003;
Brissac et al., 2009, Gros et al., 2012) and reproductive
biology (Bigatti et al., 2004). Because of the biological
interest in Ctena orbiculata, its abundance in the
western Atlantic and the possible existence of sibling
species, it has become important to stabilise this well-
established name by the designation of a neotype from
an appropriate western Atlantic type locality and for
which genetic data is available. The International Code
of Zoological Nomenclature (ICZN, 1999) specifies
(Article 75.3.6) that the neotype should come from a
place as near as practicable to the original type locality.
In this case, the geographical range of the species lies
several thousand kilometres west of the original (likely
erroneous) type locality and we have chosen the locality
in the Florida Keys because we have published molecu-
lar data from a co-occurring, morphologically similar
specimen to the neotype.
SYSTEMATICS
Subfamily Codakiinae Iredale, 1937
Genus Ctena Morch, 1861
Type Species: Codakia (Jagonia) mexicana Dali, 1901,
subsequent designation Dali, Bartsch and Rehder, 1938.
Ctena orbiculata (Montagu, 1808)
(Figures 50-59)
Venus orbiculata Montagu, 1808: 42, pi. 29, fig. 7.
Neotype: Whole shell, live-collected, FMNH 339457,
collected by G. Bigatti, M. Perhada, J. Taylor and
E. Glover during the International Marine Bivalve
Workshop 2002, stn. IMBW-FK-622A, 22nd July 2002
(Mikkelsen and Bieler, 2004).
J. D. Taylor et al., 2013
Page 143
Figures 50-59. Ctena orbiculata. 50. Venus orbiculata, copy of original figure from Montagu (1808). 51-55. Neotype, Venus
orbiculata, Long Key, FMNH 339457, L=13.6 mm. 51-52. Exterior of right and left valves 53. Dorsal view of conjoined
valves. 54-55. Interior of left and right valves. 56-59. Ctena orbiculata, SEM details of coated specimen from same sample
and location as neotype, NHMUK 20130563. 56-57. Hinge teeth of left and right valves. Scale bar = 2 mm. 58. Microsculpture.
Scale bar = 10 pm. 59. Protoconch. Scale bar = 50 pm.
Page 144
THE NAUTILUS, Vol. 127, No. 4
Type Locality: USA, Florida Keys, Long Key, bayside,
24°49.5' N, 80° 48.9' W. 0.5-1 .0 m, thin sand over rock
platform with Halodule wrightii , Thalassia testudinum ,
and St/ringodium filiforme (Mikkelsen and Bieler, 2004).
Description of Neotype: Shell small, L = 13.6 mm,
H = 12.5 mm, T (single valve) = 4.2, longer than high,
H/L = 0.92, moderately inflated, T/L = 0.31, exterior
and interior pale, yellowish white. Sculpture of many
radial ribs (55 at periphery), crossed by fine commarginal
lamellae producing scales on summits of ribs. Ribs both
divide and intercalate. Prominent major growth halts
present. Lunule lanceolate. Ligament short, set in shal-
low groove. Hinge robust; RV with two cardinal teeth,
anteriormost very small, anterior and posterior lateral
teeth present, LV with two cardinal teeth, posterior tooth
thin, anterior and posterior lateral teeth present. Anterior
adductor scar short, broad, detached from pallial line
for Vz length at an angle of 15°. Posterior adductor scar
reniform. Shell interior within pallial line with radial
grooves. Pallia! line entire. Inner shell margin denticulate.
Remarks: Details of surface microsculpture and
protoconch were examined by SEM on another coated
specimen collected with the neotype (NHMUK 20130563).
The microsculpture consists of a meshwork of irregular,
shallow punetations, ea. 3-6 pm in size (Figure 58) simi-
lar to structures seen in other Codakiinae (Glover and
Taylor, in press). The protoconch (Figure 59) PI+PII =
165 pm, PI — 142 pm is irregularly lumpy, PII a narrow
rim with fine growth increments.
A specimen (NHMUK 20100254) included in molec-
ular analyses (Taylor et ah, 2011) was collected at
the same locality and time as the neotype and is very
similar in shell characters. Gen Bank numbers for
sequences of three genes; 18S rRNA (A| 581853), 28S
rRNA (AJ581887), cytochrome B (FR6S6627).
ACKNOWLEDGMENTS
We are extremely grateful to Emilio Garcia for allow-
ing us to work on the specimens of Ferrocina garciai
from his personal collection. We thank Lisa Smith
(NHMUK) for help with sequencing of F garciai.
Olivier Gros kindly sent us material of Clathrolucina
costata from Guadeloupe. The neotype and associated
specimens of Ctena orbiculata were collected during
the 2002 PEET workshop in the Florida Keys
organized by Rudiger Bieler and Paula Mikkelsen.
We thank Harry Taylor for macroimages and Alex Ball
for help with SEM. Reviewers Gene Coan and Paula
Mikkelsen made useful suggestions. We appreciate the
continuing support from Phil Rainbow and Depart-
ment of Life Sciences, NHMUK.
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THE NAUTILUS 127(4): 147-152, 2013
Page 147
Extractrix dockeryi , a new species from the Eocene
of the southeastern United States, with notes on open coiling
in the Cancellariidae (Gastropoda: Neogastropoda)
M. G. Harasewych
Department of Invertebrate Zoology
National Museum of Natural History
Smithsonian Institution
P.O. Box 37012
Washington, DC 20013-7012, USA
[email protected]
Richard E. Petit
806 St. Charles Road
North Myrtle Beach, SC 29582-2846 USA
[email protected]
ABSTRACT
Extractrix clockert/i is described from beds of Middle Eocene
(Bartonian) age in the Gosport Sand Formation at Little Stave
Creek, Alabama and the contemporaneous McBean Formation
at Orangeburg, South Carolina. This new species represents
the earliest occurrence of open coiling in the family
Cancellariidae. Other records of open coiling in the
Cancellariidae are reviewed, and the taxonomic status and
composition of the genus Extractrix is discussed.
Additional Keywords: Cancellariidae, Extractrix , open coiling,
Eocene
INTRODUCTION
We report the discovery of a new species of open-coiled
cancellariid from two different Middle Eocene (Bartonian)
deposits in the southeastern United States, one, tire
Gosport Sand Formation at Little Stave Creek, Alabama,
the other, the McBean Formation at Orangeburg, South
Carolina. This new species represents tire earliest known
occurrence of open coiling in the family Cancellariidae. It
is assigned to the genus Extractrix Korobkov, 1955, based
on similarities to its type species, Extractrix extractrix
(Boettger, 1906), from Middle Miocene (Badenian)
deposits of the Transylvanian Basin in Romania (Lapugiu
de Sus). We review the literature for other records of
whorl detachment in the Cancellariidae, and discuss the
taxonomic status and composition of the genus Extractrix.
OCCURRENCE OF OPEN COILING IN
CANCELLARIIDAE
Whorl detachment, the lack of contact between succes-
sive whorls of a coiled shell, is rare among gastropods.
The degree and regularity of whorl detachment may vary
considerably, from simple detachment of portions of the
final whorl, as in Valvata sincere ontariensis Baker, 1931
(see Clarke, 1973: 225, pi. 20, figs. 8,9) to the occurrence
of multiple detached and loosely or irregularly coiled
whorls, as in Tenagodus squamatus (Blainville, 1827) (see
Abbott and Dance, 1982: 61. as Siliquaria squamata).
We follow Yochelson, (1971: 236) in applying the term
“disjunct” to forms that have a detached last whorl;
“uncoiled” to forms in which coiling is very irregular;
and “open-coiled" to forms in which the whorls are
detached, yet coiling conforms closely to a logarithmic spiral.
The prevalence of whorl detachment within particular
gastropod taxa may also vary'. Disjunct, uncoiled and
open-coiled forms of whorl detachment may occur as
unique or rare events in normally coiled species, either
as teratalogical or gerontie (Yochelson, 1971: 236) condi-
tions, or as a malformation following unsuccessful preda-
tion involving slit'll breakage. Whorl detachment may
also occur more widely in predominantly tightly coiled
taxa [e.g., in Spirolaxis rotulacatherinea (Melvill and
Standen, 1903), see Bieler, 1993:325, figs. 268-270; or in
Valvata jtdiae Scholz and Glaubreeht, 2010, (their fig. 2
documents frequencies of open coiling of up to 25% in
various populations)]. Detached whorls may also become
a fixed character of a species or subspecies. Rex and Boss
(1976) reviewed the incidence of such open-coiling in
Recent gastropods and reported that it occurs at very'
low frequency (~7.5xl0 4). Among the examples they
documented, 26.7% were planispiral species in which
open coiling was due primarily to increases in the dis-
tance of successive whorls from the axis of coiling [high
values of D, according to the terminology of Raup’s
(1961, 1966) model of shell coiling]. For the majority of
taxa (60.0%), open coiling was a consequence of elonga-
tion of the shell along the axis of coiling [high whorl
translation rate, T (Raup, 1961; 1966)], while 13.3% of
the taxa were open-coiled according to both criteria. In
some instances, open coiling appears to be sufficiently
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THE NAUTILUS, Vol. 127, No. 4
adaptive to have given rise to diversified open-coiled
lineages [e.g., Cycloscala Dali, 1889 (Epitoniidae);
Eccliseogyra Dali, 1892 (Nystiellidae) ; Extractrix
Korobkov, 1955 (Cancellariidae)].
Known instances of whorl detachment witliin the family
Cancellariidae are listed in Table 1 . Extractrix extractrix ,
E. dockeryi new species, and E.milleri (Burch, 1949) all
show a marked increase in the degree of elongation
(increase in the whorl translation rate, T) at the
protoconch / teleoconch transition, resulting in open
coiling for all teleoconch whorls (only the first 0-14
teleoconch whorls are in contact with prior whorls).
Although Janssen (1984a: 21) considered Trigonostoma
protrigonostoma Sacco, 1894 to he closely related to
Extractrix extractrix , the former has a shell (Sacco, 1894:
pi. 1 , figs. 3a,3b) that more closely resembles Trigonostoma
umlnlicare (Brocehi, 1814), differing from that species in
having a portion of the body whorl (~ 14 whorl) disjunct.
In contrast, the early teleoconch whorls of Trigonostoma
(“Extractrix”) hoerlei Olsson, 1967 are nearly planispiral,
with the increase in the elongation rate occurring in the
second teleoconch whorl (Figure 23, arrow), resulting in a
shell in which only the last 1-114 whorls are disjunct.
There is a report (Coucom, 1975) of a single disjunct
specimen of Trigonostoma scalare , a species that is nor-
mally coiled, in which the last 14 whorl is detached.
SYSTEM ATICS
Family Cancellariidae Forbes and Hanley, 1851
Subfamily Cancellariinae Forbes and Hanley, 1851
Genus Extractrix Korobkov, 1955
Pseudomalaxis ( Extractrix ) Korobkov, 1955:138. Type
species: Pseudomalaxis extractrix (Boettger, 1906) by
original designation.
Trigonostoma ( Extractrix ) Olsson, 1967:23-24.
Diagnosis: Shell small to medium-sized (to 27 mm),
with tall, conispiral, deeply umbilicate, open-coiled shell.
Protoconch of smooth, evenly rounded whorls. Transi-
tion to teleoconch marked by major increase in degree
of elongation along the axis of shell coiling, resulting in
loss of contact between whorls within first !4 teleoconch
whorl. Teleoconch of up to 4 whorls, sharply triangular in
cross-section. Carinae along shoulder and siphonal canal
with open spines (long to barely discernible).
Remarks: The type species of Extractrix was
described as Discohelix ( Pseudomalaxis ) extractrix, in
the family Architectonicidae by Boettger (1906: 138),
and retained in this family by subsequent authors (e.g.,
Cossmann, 1916; Zilch, 1934), including Korobkov
(1955), who proposed Extractrix as a subgenus of
Pseudomalaxis. Citing Boettger’s (1906: 138) reference
to two small columellar plaits, Olsson (1967:23) trans-
ferred Extractrix to the family Cancellariidae, as a sub-
genus of Trigonostoma , but regarded it as a “form genus”
and questioned the relationship of its type species to
other loosely coiled cancellariids. Janssen (1984a: 20)
treated the type species as belonging to Trigonostoma
sensu stricto, and considered it to be closely related to
Trigonostoma protrigonostoma Sacco, 1894, a species in
which only a portion of the body whorl is disjunct.
Harzhauser and Landau (2012: 56) regarded the status
of the genus Extractrix as questionable. They considered
the species assigned to this genus to form a morphologi-
cal group based on open coiling that was not necessarily
monophyletic, but possibly an assemblage of indepen-
dently evolved, uncoiled species of Trigonostoma.
Prior authors have assigned cancellariid species to the
genus Extractrix primarily, if not exclusively on the basis
of the presence of either disjunct whorls or open coiling,
an approach that has rendered this genus paraphyletie.
The updated diagnosis of Extractrix is based on more
precisely defined shell characters that are shared by the
type species and some, but not all cancellariid species
with disjunct or open coiled shells. We conclude that
these characters are sufficient to define a clade that is
related to Trigonostoma, but lias persisted from the
Table 1. Records of whorl detachment witliin the family Cancellariidae.
Harzhauser and Landau (2012:56) suggested that the specimens from the Netherlands reported by Janssen ( 1984a, b) represent a
species different from E. extractrix.
3 Landau et al. (2012:223)
1 Jung (1977) reported this species from the early Pliocene Punta Gavilan Formation of coastal Venezuela.
5 Coucom (1975)
M.G. Harasewych and R.E. Petit, 2013
Page 149
Eocene to the Recent, with records in the eastern
Pacific [Pleistocene-Recent], southeastern United States
[Eocene] and northern and eastern Europe [Miocene].
Extractrix dockeryi new species
(Figures 1 -13)
Description: Shell (Figures 1-13) small (to 10.7 nun)
thin, with tall, narrow (spire angle 51-66°), conispiral.
deeply umbilicate, open-coiled shell. Protoconch
(Figures 6, 7) increasing in diameter from 83 pm to
1 .09 mm in 2]A smooth, evenly rounded whorls, deviated
from teleoconch axis by ~9°. Transition to teleoeonch
distinct, marked by slight change in surface texture and
curvature (Figures 6, 7, arrows), as well as a major
increase in the degree of elongation along the coiling axis
of the shell [rate of whorl translation, T (Raup, 1966)
increasing from ~1.5 for the protoconch to ~3.25 for
Figures 1-13. Extractrix dockeryi new species. 1-8. Holotype, UF 242210, Little Stave Creek, Alabama. 1. Apertural. 2. Lateral,
3. Dorsal, 4. Apical, and 5. Umbilical views of the shell. 6. Apical and 7. Lateral views ot the protoconch. Arrows indicate transition
from protoconch to teleoconch. 8. Columella. Arrows indicate vestiges of columellar folds. 9-1 1. Paratype 1, Petit Collection 3488,
about 3 miles WNW of Orangeburg, Orangeburg County, South Carolina. 12-13. Paratypes 2 and 3. FMNH 328571, Edge of
Caw Caw Swamp, WNW of Orangeburg, Orangeburg County, South Carolina. 5 mm scale bar applies to Figures 1-5 and 9-1 L
500 pm scale bar applies to Figures 6, 7; 1 mm scale bar applies to Figures 8, 12, 13. Abbreviations: pc, peripheral carina; s, shoulder
spines; ssc, siphonal canal spines.
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THE NAUTILUS, Vol. 127, No. 4
teleoeonch] that results in none of the teleoconch whorls
being in contact with each other. Teleoconch with up to
2Vz sharply shouldered whorls that are strongly triangu-
lar in cross section. The adabical portion of each whorl
is flat, tabulate, nearly perpendicular to coiling axis
(forming an angle of 70-82°), the abaxial margin is
demarcated by a sharply angled shoulder with a weak
peripheral carina (Figure 4, pc) bearing broad, open
spines. Outer portion of each whorl weakly convex, sinu-
ate and recurved at juncture with short siphonal canal.
Columellar portion of each whorl straight, nearly parallel
to coiling axis in early whorls (Figures 12, 13), angle
increasing up to 22° in later whorls (Figure 1). Outer
portion of whorl and columella form an acute angle
(30-39°), bearing a carina along their juncture, while
the juncture of the columella and the adapical region is
evenly rounded. Spiral sculpture absent or limited to
barely perceptible, extremely fine spiral threads on por-
tions of the outer surface of the last whorl. Axial sculp-
ture of simple, weak, prosoeline, growth striae, raised,
open spines (Figure 1, s) along shoulder (8-11 on
last whorl), and weaker, adaxially deflected open spines
(Figure 1, sse) along the carina at the anterior edge of
the siphonal canal (0-7 on last whorl). Spines on shoul-
der and siphonal canal may be prominent or weak and
worn. Aperture triangular, with short, pronounced
siphonal canal. Outer lip smooth, columella with two
very weak columellar folds (Figure 8, arrows).
Type Material: Holotype, Florida Museum of Natural
History, University of Florida, UF 242210; Paratype 1,
Petit Collection 3488, About 3 miles WNW of
Orangeburg, Orangeburg County, South Carolina,
[McBean Formation, Middle Eocene (Bartonian)];
Paratypes 2-3. The Field Museum, FMNH 328571, Edge
of Caw Caw Swamp, WNW of Orangeburg, Orangeburg
County, South Carolina [McBean Formation, Middle
Eocene (Bartonian)].
Type Locality: Little Stave Creek, Clarke County,
Alabama [Gosport Sand Formation, Claiborne Group,
Middle Eocene (Bartonian)].
Distribution and Habitat: This species is presently
known from the type locality as well as from the McBean
Formation, WNW of Orangeburg, South Carolina. Both
deposits are of Middle Eocene (Bartonian) age. The sand
deposits at the type locality were “probably laid down in
less than 20 fathoms [36.6 m] of water” (Gardner, 1957:
584). The Orangeburg Sand of the McBean Formation
correlates with the Gosport Sand (Dockery et ah, 1992)
and was also deposited on a shallow marine shelf, mainly
in low energy conditions (Nystrom et ah, 1991).
Etymology: This new species honors Dr. David T.
Dockery III, of the Office of Geology, Mississippi
Department of Geology and Environmental Quality, in
recognition of his many contributions to the study of the
Tertiary paleontology of Alabama and Mississippi.
Comparative Remarks: This new species most
closely resembles Extract rix extractrix (Boettger, 1906),
the type species of the genus Extractrix Korobkov, 1955
(Figure 14), a Miocene species that has been reported
from the Burdigalian-Langhian of Romania (Boettger,
1906) and the Netherlands (Janssen, 1984a,b) as well as
from the Messinian of Italy (Davoli, 1995). Both species
have open-coiled shells in which teleoconch whorls
are not in contact due to high rates of whorl translation
(T = 3. 0-3. 8), sharply triangular whorl profiles, and lack
pronounced spiral sculptural elements. Extractrix dockeryi
new species differs from E. extractrix in having propor-
tionally broader, more inflated whorls, and pronounced
earinae along the shoulder, and at the end of the short,
constricted siphonal canal. The presence of open spines
distinguishes E. dockeryi from Romanian and Italian
specimens of E. extractrix, but specimens from the
Netherlands may have low spines along both earinae that
may be connected by weak axial ridges (Janssen, 1984a:
pi. 2, fig. 16; 1984b: pi. 66, figs. 6,7). Harzhauser and
Landau (2012: 56) suggested that the specimens from
the Netherlands represent a species different from
E. extractrix. Extractrix dockeryi is easily distinguished
from its Pleistocene-Recent congener, E. milleri (Burch,
1949) (Figures 15-21), a species from the tropical eastern
Pacific that has a much larger (to 26 mm), broader shell
with axial and spiral sculpture.
DISCUSSION
As more rigorously defined, the genus Extractrix con-
tains its type species Extractrix extractrix from the Mio-
cene of Europe, E. dockeryi new species from the
Eocene of the southeastern United States, and Extractrix
milleri from the Pleistocene- Recent of die tropical east-
ern Pacific. In the event that the specimens from the
Miocene of the Netherlands that had been attributed to
E. extractrix by Janssen (1984a, b) are segregated as
a separate species, as suggested by Harzhauser and
Landau (2012), this too should be included in Extractrix.
Both “ Trigonostoma ” hoerlei and “Trigonostoma”
protrigonostoma are excluded from Extractrix as both
have disjunct rather than open coiled shells. The rela-
tionship of T. hoerlei (Figures 22-26) appears closer to
some species of Ventrilia, suggesting an independent
origin of whorl detachment.
The genus Extractrix appears to have originated in the
western Atlantic during die Eocene and spread eastward
to Europe, where widespread populations (North Sea,
Paratethys and Tethys Basins) were present during the
Miocene. It survives in the Recent fauna as a paciphile
genus, which once inhabited the western Tertiary Carib-
bean Province, but is now presumably extinct there and
survives in eastern Pacific waters (Woodring, 1966: 426).
Open coiling is generally regarded as maladaptive as it
increases vulnerability to shell breaking predators and
impedes mobility, but may survive in situations where
such ecological limitations are relaxed (e.g., when
M.G. Harasewych and R E. Petit, 2013
Page 151
Figures 14-26. Extractrix species. 14. Extractrix extractrix (Boettger, 1906). Lectotype, Lapugiu de Sus, Romania (Specimen
lost, see Harzhauser and Landau, 2012: 56. Figures from Cossmann 1916: pi. 12, figs. 25-26). 15-21. Extractrix milleri (Burch,
1949). 15. Apertural view of Holotype USNM 600660, Near Punta Arenas, Tambor, Costa Rica. 16. Apertural view of USNM
679301, Guaymas, Mexico 17. Apertural, 18. Lateral, 19. Dorsal, 20. Apical and 21. Umbilical views oi shell Petit Collection 2397,
Arenas de Quebro, Panama, in 73 m. 22-26. Trigonostorna ("Extractrix”) hoerlei Olsson, 1967. Holotype, USNM 645162, Kissimmee,
Florida. 22. Apertural, 23. Lateral, 24. Dorsal, 25. Apical and 26. Umbilical views of shell. Scale bar = 3 mm for figure 14;= 1 cm for
Figures 15-21; = 1.5 cm for Figures 22-26.
populations expand following a calamity, or when a pre-
viously unoccupied or temporarily favorable environ-
ment is colonized) (Vermeij, 1987: 42).
Disjunct coiling in Valvata juliae , for example, was
limited to a short stratigraphic interval within the Pleis-
tocene, and believed to be a response to high levels of
environmental stress caused by lake level fluctuations in
the Lake Turkana (Scholz and Glaubrecht, 2010).
Rex and Boss (1973) conceded that the selective
significance of open coiling is unclear, and recognized
that it evolved independently in several unrelated lineages
to fill “dissimilar environmental roles." They noted the
success of some open-coiled taxa in terms of broad geo-
graphic distributions and/or their persistence through
geological time. Several lineages with open-coiled shells
(e.g., genera within Epitoniidae, Nystiellidae, Architec-
tonieidae) are known to be ectoparasites of cnidarians.
Cancellariids are characterized by an alimentary system
highly specialized for suctorial feeding on body fluids
(Harasewych and Petit, 1998). Depending on species,
prey organisms range from sharks and rays (O’Sullivan
et ah, 1987) to bivalves and gastropods (Loch, 1987).
Page 152
THE NAUTILUS, Vol. 127, No. 4
No eancellariid species have yet been reported to feed on
enidarians. When the habitat and diet of Extractrix milleri
is better known, it may shed insights on the advantages
open coiling bestows on species of the genus Extractrix, a
lineage that has persisted from the Eocene to the Recent.
ACKNOWLEDGMENTS
We are grateful to Dr. David T. Dockery III, of the
Office of Geology, Mississippi Department of Geology
and Environmental Quality, and to the late William |.
Fulton, of Chamblee, Georgia, for making available most
of the specimens in the type series.
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THE NAUTILUS 127(4): 153-155, 2013
Page 153
A new land snail of die genus Eucalodium from Belize
(Gastropoda: Pulmonata: Urocoptoidea: Euealodiidae)
Fred G. Thompson
Florida Museum of Natural History
University of Florida
Gainesville, Fl 32611 USA
[email protected]
Daniel C. Dourson
Belize Foundation for Research and Environmental Education
P.O. Box 129
Punta Gorda, BELIZE
[email protected]
ABSTRACT
Eucalodium belizensis new species is described from Belize It
is the first species of the genus reported from there.
Additional Keywords: Gastropods, land snails. Central America
INTRODUCTION
The subgenus Eucalodium is known from mountainous
terrains in a rather small area of Belize, Guatemala, and
part of Mexico (northern Chiapas and Tabasco.)
Eucalodium species of are seldom encountered, and
they are not common where they are found. This is the
first species of the genus and the subgenus reported
from Belize. The species is described prior to a review
of the genus Eucalodium by the senior author.
MEASUREMENTS
The length of adult shell is measured from the top of
the apical plug to the base of the aperture. The width
of the shell is perpendicular to the axis, but does not
include the aperture. The apex is the whorl at the
apical plug. The following abbreviations are used in the
text: Afl = height of aperture; AW = width of aperture;
ApexW = width of apical whorl; SL — standard length;
SW = standard width.
Eucalodium ( Eucalodium ) belizensis new species
Figures 1—4
Diagnosis: The shell is purple-brown. It is decollated,
solid, and thick-walled, about 51-57 mm long, and
includes 9.4-1 1.8 whorls below the apical plug. The
aperture is broadly ovate in shape. The columella is
strongly twisted, producing a flattened, crenulated
lamella. The uniformly tapered spire is nearly straight-
sided. The sculpture consists of distinct riblets on the
upper 4-5 whorls. The riblets are less distinct or obsolete
on the lower half of the shell, and then again become
distinct on last whorl.
Description: The solid thick-walled shell is purple-
brown with a lighter colored base and with a white peri-
stome (Figure 1). It is moderately large, attaining a
length of about 51-57 mm below the apical plug, 14.4-
15.0 mm standard width, 0.25-0.28 times as wide as long.
The aperture is about 0.71—0.85 times the shell width.
The shell is decollate and contains 9.4-11.8 whorls. The
apical whorl is 7.0-7. 8 mm wide, and is 0.48-0.52 times
the standard shell width. The whorls are nearly flat with a
shallow suture. The spire is slightly concave in profile
along the earlier whorls and is slightly convex below.
The apical plug is oblique and nearly flat. The base of
the last whorl is rounded with an obsolete circum-basal
cord. The imperforate umbilicus is marked by a narrow
indentation. The sculpture consists of numerous oblique,
slightly sigmoid, sharp thread-riblets. The riblets are
most distinct on the upper whorls where they are about
as wide as their interspaces, and are slightly higher than
wide (Figure 3). The riblets are less defined and distinct
on the lower part of the spire, and then again become
more distinct on the last whorl. In one paratype they
are distinct nearly through the length of the spire. The
holotype has 96 riblets on the penultimate whorl. A
paratype (UF 448722b) has 106 riblets. The aperture
projects forward slightly in lateral profile (Figure 2), and
is oblique, lying at an angle of about 20-21° to the
shell axis (21 in holotype). It is broadly ovate in shape
with an obtuse posterior corner. The parietal margin is
weakly convex. The peristome is narrowly reflected and
rounded. The peristome is widest along the palatal and
basal margins, and is narrowest along the parietal margin.
A columellar fold extends down and forward to the peri-
stome. The internal axis is strongly twisted, producing a
flattened spiral lamella that is crenulated along its edge
(Figure 4). The lamella extends for about 0.1 of the
distance across the cavity of the whorl. The crenulated
nodes on the lamella are most developed in the penulti-
mate and anti-penultimate whorls.
Page 154
THE NAUTILUS, Vol. 127, No. 4
Figures 1-4. Eucalodiurn belizensis new species. 1-3. Holotype (UF 449720). 4. Paratype (UF449722).
Type Material: Holotype: UF 449720, April 2010.
Paratypes: UF 449721 (one specimen), April 2010;
UF 449722 (2 specimens), April 2012. All type material
Iron) type locality anti collected by Valentino Tzub.
Three paratypes are available. Two (UF 449721, UF
449722b) are bleached and worn specimens in which
the sculpture is no longer clearly evident. The third
paratype (UF 444922b) retains the periostracum and
the sculpture. It differs from other paratypes and the
holotype by having a relatively wider aperture in which
the columellar lamella is stronger anti distorts the
columellar lip. Standard measurements of type material
are given in Table 1 .
Type Locality: Belize, Toledo District, 5.5 km north
of San Jose; 2 km northeast of Miramar Hill (16.335° N,
89.114° W), 650 m altitude.
The type locality is reached by the Southern Highway
about 8 km from the intersection of the road to Punta
Gorda, and right (north) on a paved road, past Crique
Jute for about 33 km to the village of San Jose. The type
Table 1. Measurements in nun of type material of Eucalodiurn
belizensis new species.
locality is approximately 5.5 km due north on a trail
from San Jose.
The collector, Valentino Tzub, is a Kek’chi Mayan
from the village of San Jose who occasionally works as
a guide and research assistant for scientific expeditions
and research projects in Belize.
Habitat: Specimens were found under leaf litter near
Cretaceous limestone outcrops. The landform surround-
ing the type locality includes hilly karst topography,
containing sinkholes and multiple cave formations. The
type locality is entirely forested with a tropical wet broad-
leaf evergreen forest with eohune palms ( Attalia cohune)
and occasional emergent tree ( Ceiha ) and with an under-
story layer dominated with shrubs, pteridophytes, and
Araeeae. Farming activity from San Jose is encroaching
into the near-by forest.
DISCUSSION
Eucalodiurn belizensis is the first record of the genus
Eucalodiurn in Belize. Eucalodiurn Fischer and Crosse,
1868 includes four subgenera: Eucalodiurn s.s.,
Oligostylus Pilsbry, 1895, Resupinata Martens, 1897,
and Ptijchocentmm Bartsch, 1943 (Thompson, 2011).
Oligostylus and Resupinata are distinguished from
Eucalodiurn s.s. by lacking an internal lamella on the
axis, and by having thin-walled shells with a brownish
periostracum. The generic affinity of Ptychocentrum is
problematic. It has an internal lamella similar to that in
species of the subgenus Eucalodiurn , but the lamella is
not crenulated, and the axis is hollow, approaching that
F.G. Thompson and D.C. Dourson, 2013
Page 155
of Coelocentrum species. Its shell is thick-walled and is
chestnut-brown in color.
The subgenus Eucalodium is known for certain from a
rather small area of northern Chiapas, Tabasco (both in
Mexico), Guatemala, and Belize. Formerly the subgenus
must have been generally distributed and common
within this area, but deforestation has extirpated it from
much of its former range. Records from there are few.
Species are seldom encountered, and they are not com-
mon where they are found. The paucity of specimens
in museum collections makes taxonomic study difficult.
All locality records for the subgenus Eucalodium lie north
of the North American-Central American Continental
Divide in the Polochfc Valley of Guatemala-Chiapas.
Oligostylus is more widespread, ranging from Jalisco,
M exico south to El Salvador (Thompson, 1963).
Resupinata is found from San Luis Potosi south the
Veracruz. Ptychocentrum is known from a single locality
in Chiapas.
The subgenus Eucalodium includes E. compaction
Pilsbry, 1893, E decollation decollation (Nyst, 1841),
E. d. guatemalensis (Bartseh, 1906, E otoides Thompson,
1968, and E. mexicanum (Pfeiffer, 1860). The species are
medium sized to large and have a strongly twisted axis that
forms a flattened spiral lamella, which may be crenulated
along it edge (as in Figure 4). Eucalodium d decollation
is known only from small area near Teapa, Tabasco
(Thompson, 201 1). Eucalodium decollation guatemalensis
is reported from an unspecified locality in Guatemala
(Bartseh, 1906). Eucalodium otoides is known from a
single locality in northern Chiapas, and E compaction is
known from an unspecified locality in Tabasco. Pfeiffer
(1860) described E mexicanum from an unspecified
locality in Mexico. Fischer and Crosse (1873) record a
variety of E ( Eucalodium ) mexicanum from between
Tactic and Tamahu, Dept. Alta Verapaz, Guatemala, and
they record two forms of E mexicanum from Juquila,
Oaxaca. Their records concerning the distribution of
E mexicanum appear unlikely because of the wide geo-
graphic separations.
Eucalodium belizensis is similar in size to E compaction ,
which differs from £. belizensis by having weaker, more
widely spaced sculpture, a concave parietal margin in the
aperture, and a thinner, light-brown shell Eucalodium
otoides differs from E. bezensis by having fewer whorls
below the apical plug (7. 6-7. 9) and by having an auriform
aperture with a straight parietal margin, a strongly
reflected and twisted eolumellar margin that forms an
obtuse angle with the basil margin, and a more robust
shell that is 0.31-0.32 times as wide as long. Eucalodium
decollation is a large, robust species. Shells are 64—74 mm
in length, and 0.28-0.31 times as wide as long. The sculp-
ture consists of poorly defined oblique riblets that are
nearly obsolete over most of the shell. There are about
75-86 obsolete riblets discernible along he suture on the
penultimate whorl. Eucalodium decollation guatemalensis
is similar in length and whorl count to E. belizensis , except
that it is more robust, being 17.5 mm wide at the pen-
ultimate whorl, and it has a smooth, not crenulated, eolu-
mellar lamella. Eucalodium mexicanum differs from
E. belizensis by having a thin-walled shell that is light
brown in color.
ACKNOWLEDGMENTS
We thank Steven Brewer, Belize Foundation for
Research and Environmental Education, Punta Gorda,
Belize for the habitat description.
LITERATURE CITED
Bartseh, P. 1906. The urocoptid mollusks from the mainland of
America in the collection of the United States National
Museum. Proceedings of the United States National
Museum 31: 100-160.
Bartseh, P 1943. A new subgenus and species of Coelocenrmn.
The Nautilus 56: 91-92.
Fischer, P. and H. Crosse. 1870-1878. Mission Scientifique au
Mexique et dans L’Amerique Centrale. Etudes sur les
mollusques terrestres et fluviatiles du Mexique et du
Guatemala, I Paris, 702 pp.
Martens, E. v. 1890-1901. Biologia Centrali-Americana.
Mollusca. British Museum (Natural History), 706 pp.
Nyst, H. 1841. Notices sur deux Coehilles Mexicaines
appurtenant aur genres Pupa et Helix. Buletin de
f Academic Royal de Bruxelles 8: 343.
Pfeiffer, L. 1860. Descriptions of thirty-six new species ol land
shells from Mr. 11 Cuming’s collection. Proceedings of
the Zoological Society of London 1860: 1 313— 141.
Pilsbry, H.A. 1893a. Notes on a collection of shells from the
state ol Tabasco, Mexico. Proceedings ol the Academy ot
Natural Science of Philadelphia 44: 338-341.
Pilsbry, H.A. 1902-1903. Manual of Conchology: Uroeoptidae.
Ser. 2, Vol. 15. Academy of Natural Sciences, Philadelphia,
323 pp.
Strebel, H. and G. Pfeffer. 1880. Beitrag zur Kenntniss der
Fauna mexikaniseher Land- und Susswasser-Conchylien.
Teil IV. Hamburg, J. J. Kerbst, 122 pp., pis. 15 pis
Thompson, F’.G. 1963. New land snails from El Salvador.
Proceedings ot the Biological Society ol Washington 76:
19-32.
Thompson, F’.G. 1968a. Some Mexican land snails of the family
Uroeoptidae. Bulletin of the Florida State Museum 12:
125-183; fig. 1-29.
Thompson F.G. 201 1. An annotated checklist and bibliography
ol the land and freshwater snails of Mexico and Central
America. Bulletin of the Florida Museum of Natural
History 50: 1-299.
THE NAUTILUS 127(4): 156-159, 2013
Page 156
Determination of EC50 for normal oyster larval development
in extracts from bloom-forming green seaweeds
Timothy A. Nelson
Department of Biology, Suite 205
Seattle Pacific University
Seattle, WA 98119-1950 USA
[email protected]
Bridget C. Gregg
NewFields Northwest, UPC.
Port Gamble Environmental Laboratory
PO. Box 216
4729 NE View Drive
Port Gamble, WA 98364 USA
[email protected]
ABSTRACT
Blooms of ulvoid algae can decimate marine ecosystems, out-
competing seagrasses and other habitat-forming species and
directly killing animals due either to the production ol allelo-
chemicals or various effects (e.g., anoxia) associated with bloom
mortality. Oyster larvae ( Crassostrea g igas) are susceptible to
extracts from dried ulvoids at relatively low concentrations.
Here we determine a specific EC50 for extracts from the
bloom-forming species Ulva lactuca and Vivaria obscura.
The EC50 for Ulva is equivalent to 0.13 gfm'L-1 while the
value for Vivaria is 0.21 gfm'L 1 Assuming moderate bloom
conditions, these data suggest that an EC50 could be achieved
in 5,200 to 8,500 L of seawater for each m“ of algid bloom.
Possible mechanisms for developmental toxicity include the
breakdown products of DMSP (in both species), dopamine
quinones (in Vivaria only), or as yet unidentified toxins.
Additional Keywords: Crassostrea gigas , Ulvaria obscura, Ulva
lactuca. Allelopathy, Harmful Algal Blooms
INTRODUCTION
So-called “green tides” (blooms of green macroalgae) have
been increasing worldwide causing problems ranging
from anoxia to direct overgrowth of valuable marine plant
communities (Valiela et al., 1997, Hauxwell et al., 2000).
The organisms causing green tides are predominantly
ulvoid macroalgae (Family Ulvaceae), though other spe-
cies may be responsible (e.g., Lapointe et al. 2005).
Mass mortality of fish and bivalves coincident with
macroalgal blooms have been reported in Sweden
(Rosenberg et al., 1990). Their effects have directly
altered human activities as well, e.g., causing concerns
with sailing events during the 2008 Olympic Games and
the death of domestic animals (Anonymous, 2009). In
Washington State, USA, the public has been alarmed
by bloom-induced anoxia creating potentially dangerous
H^S concentrations in residential neighborhoods. Local
blooms are sufficient to negatively impact eelgrass growth
(Nelson and Lee, 2001).
Specifically allelopathic impacts of ulvoid algae have
been noted or suggested occasionally in the past.
Extracts from Ulva spp. are thought to inhibit larval
development in barnacles, crab, flounder, and mosqui-
toes (Magre, 1974; Johnson, 1980; Johnson and Welsh,
1985; Thangam and Kathiresan, 1991, Walters et al.,
1996, Borowsky et al., 1997). Nelson et al. (2003a) have
demonstrated that crude, water-soluble extracts from
two bloom forming species ( Ulva lactuca L., 1753 and
Ulvaria obscura [Kiitzing] Gayral, 1969) inhibit normal
development of oyster ( Crassostrea gigas Thunberg,
1793) zygotes. Here we determine the concentration
of extracts from two common “green tide” seaweeds
(U. lactuca and U. obscura) that result in a 50% decline
in normal oyster zygote development through the D-
hinge stage (EC50).
MATERIALS AND METHODS
Algae were collected from Seattle, Washington, USA
(47°32.4' N, 122° 23. 8' W). Algal disks (9 cm diameter,
approximately 0.7 gfm per disk) were dried for 30 min
outdoors in the shade, effectively killing the alga. Dried
disks were placed in microwave-sterilized seawater for
90 min at a ratio of one disk per 50 mL of seawater
(or ~14 gfm'L-1). This yielded what we estimated to be
the maximum concentration of extract that organisms
encounter in the field and is scaled to 100% hereafter.
The concentration is equivalent to a volume of seawater
5 cm deep covering the densest portion of a heavy bloom
of ulvoid algae (i.e. ~six algal thalli thick, as observed by
the authors and reported in terms of mass by Nelson
et al., 2003b).
EC50 for oyster embryo larval development was deter-
mined following standard techniques (ASTM, 1989,
described fully in Nelson et al., 2003a). The study was
conducted during late July to correspond with the
natural gamete maturation in oysters. Oysters were
T.A. Nelson and B.C. Gregg, 2013
Page 157
hand-spawned and aliquots of sperm were added to
egg suspension to achieve a ratio of 10-20 sperm
per egg. Within 1 h of fertilization, the embryo suspen-
sion was transferred into test chambers (polystyrene
Coulter counter vials).
For Ulvaria , we tested concentrations of 0%, 0.1%,
0.3%, 0.6%, 1.6%, and 3.9%. For Ulva , we tested con-
centrations of 0.5%, 0.9%, 1.5%, 2.7%, and 4.7%. Each
treatment dilution had 7 (Ulva, at 2.7%) or 8 (all others)
replicates. Filtered seawater was used for seawater con-
trols, stock solutions, and dilutions. The test chambers
were maintained at 25°C for 24 h. Temperature and [CL]
of a separate container of seawater were checked every'
4-6 h as a proxy for the incubation vials containing oyster
larvae. Larvae were fixed with I mL of formaldehyde
added to each vial after 24 hours. All larvae in each test
chamber were identified as normal or abnormal based
on their development to the D-hinge stage.
EC.50 of Ulva and Ulvaria extracts were determined by
fitting a non-linear regression as follows:
Y = B + (T — B)/(l + (X/EC50)(H)) (Equation 1),
where X is the concentration tested, Y is the fraction
of the larvae that developed normally, B represents the
lowest possible proportion surviving, T represents the
Percent of Maximum Likely Concentration
Figure 1 . The fraction of oyster larvae developing to the D-hinge stage in various concentrations of extracts from Ulva lactuca and
Ulvaria obscura. Best-fit curves are shown for U. lactuca (solid line) and U. obscura (dashed line) based on the Equation 1 where
Y = (0+ (0.867— 0))/( 1 + ([Ulvaria extract]/!. 494%)°'906) and Y = (0+(0.835-0))/(l + ([[/. lactuca extractJ/0.910%)3^654).
Page 158
THE NAUTILUS, Vol. 127, No. 4
highest possible proportion surviving, and H is the Hill
coefficient. Constraints were placed on T and B (0< =
B<=0.5 and 0.5<=T<=1.0).
RESULTS
Regressions for both Ulva and Ulvaria extract concentra-
tion versus oyster larval success were significant (Figure,
F| 43=2.956, p=0.0304; and F4 44=4.982, p =0.002; respec-
tively) and explained over 90% of the variation in larval
success (R2=0.911 and 0.942, respectively. Figure). EC50
for Ulva was 0.91±0.054% and for Ulvaria 1.5±0.70%
(means±SE). The means are equivalent to 0.13 and
0.21 gfrn L ', respectively for Ulva and Ulvaria tissue.
Given a moderately dense bloom has 200 gdmm-
(Nelson et al., 2003b), the fm:dm ratio in these algae is
approximately 1:0.182, and the EC50 values reported
here, the death of the bloom would release sufficient
toxin to create EC50 concentrations in 5,200 (for Ulvaria)
to 8,500 (for Ulva) Lnf of bloom area.
DISCUSSION
Oyster developmental failure may be attributable to
one of several toxins found in ulvoid algae. Both Ulva
and Ulvaria spp. have been shown to contain nontoxic
dimethylsulfoniopropionate which breaks down sponta-
neously or enzymatically to form the toxins dimethyl
sulfide and acrylate (Van Alstyne et al., 2001; Van Alstyne
and Houser, 2003; Erickson et al., 2006; Lyons et al.,
2007). Ulvaria obscura contains dopamine, which oxi-
dizes in solution (Tocher and Craigie, 1966, Graham
et ah, 1978) to form reactive oxygen species (ROS) and
dopamine quinone. The quinone has both cytotoxic and
genotoxic effects (Stokes et ah, 1999). ROS can damage
lipids, proteins, and DNA (Halliwell and Gutteridge,
1989; Lesser, 2006). Larval crab ( Cancer spp.) survival
is significantly reduced when zoeae are exposed to
dopamine at environmentally relevant concentrations
(Harvey, 2007).
Further, it is possible that an unidentified toxin or other
cause is at play. Some authors suggest that fattv acids may
be involved. Ulva linza L., 1753 produces an approxi-
mately 400 Da lipid that inhibits Funis distichus L., 1767
germination (K. Van Alstyne, pens. comm.). Wang et al.
(2011) cite the work of Fusetani et al. (1976) on the
iethyotoxic properties of lipids produced by the green alga
Chaetomorpha minima F.S. Collins and Hervey, 1917 and
suggest that Ulva spp. may be similarly toxic. Wang et al.
(201 1) found that juvenile abafone ( Haliotis discus hannai
I no, 1953) were susceptible to toxins from media in which
living or decomposing Ulva prolifera O. F. Muller, 1778
had been steeped. Bodi were toxic with LC50 between
12-15 gfrrrL 1 algal tissue, with a slight but significantly
greater effect of decomposing tissues. The authors sug-
gest that allelochemicals produced by either the living or
decomposing algae contribute to mortality, with increased
mortality caused by decomposition-induced hypoxia as
well as release of sulfides and ammonia.
Release of these compounds and the toxic impacts of
crude extracts are associated with death of the algal tissue,
particularly when induced by desiccation (Nelson et al.,
2003a, Van Alstyne et al., 2011). Ulvaria is less desiccation
tolerant than Ulva, thus likely to release toxins following a
shorter drying period (Nelson et al., 2010). A flooding tide
on a hot day would create EC50 or higher conditions in
1,000's of L of seawater per m , potentially impacting
organisms on nearby shorelines.
Toxicity of green algal extracts to oyster larvae is partic-
ularly problematic in an era of increasing bloom intensity
and frequency. Losses in the larval stages could limit
the success of commercially grown and wild oyster
populations. Negative consequences are magnified by the
greater sensitivity of oyster larvae: The EC50 determined
here is between 1 and 2 orders of magnitude lower than
most previously tested algal and animal model systems
(Nelson et al., 2003a, Harvey, 2007, Wang et al., 201 1 ). If
toxic impacts are occurring primarily at cell surfaces, algal
cell w;dls and multicellular animal shells or exoskeletons
may limit the impacts and could account for differences
in toxic impacts noted for various species.
Algal bloom management practices should be informed
by the possible negative impact on oyster larvae. In
Washington State, USA, past practices have included sein-
ing of green algae off beaches followed by their release in
mid-channel. While appreciated bv beachfront property
owners, this practice simply moves toxins to another loca-
tion. In China, dried green algal tissue has been proposed
as a remedy for red tides (Jin et al., 2005). While effective
in limiting dinoflagellate population growth, the potential
for this “cure” harming species it is intended to protect
should be evaluated.
ACKNOWLEDGMENTS
Taylor Seafoods, Inc., provided laboratory space and
oysters for this research. This project was funded by
grants from the United States’ National Science
Foundation (IOS-0718341) to TAN, R.L. Ridgway and
K. Van Alstyne and from the National Oceanic and
Atmospheric Administration to K. Van Alstyne and T.
Nelson (NA05NOS4781192) as well as matching fund-
ing from Seattle Pacific University. This is publication
EC0763 from the Ecology and Oceanography of
Harmful Algal Blooms program.
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Intaglicollonia, a new name for the Eocene
gastropod Nehalemia Hickman, 1974
(Gastropoda: Coloniidae)
Intaglicollonia is proposed as a replacement name for
the monotypic Eocene gastropod genus Nehalemia
Hickman, 1974, family Colloniidae Cossmann, 1916.
The type species, Nehalemia hieroghjphica Hickman,
1974, was based on 11 specimens from a siltstone unit at
the top of the Cowlitz Formation in Washington County,
northwestern Oregon (early late Eocene). The siltstone
is immediately below the uneonformable contact with
the basal member of the overlying latest Eocene to earli-
est Oligocene Keasey Formation. The genus name is
preoccupied by Nehalemia Blake, 1973, an asteroid eelh-
noderm from the middle member of the Keasey Forma-
tion in Columbia County, Oregon. The type localities for
both species are on the Nehalem River, approximately
25 km apart and with a vertical stratigraphic separation
of approximately 450 m. Although the taxa are in sepa-
rate phyla, the geographic and stratigraphic proximities
of the type localities support the decision to avoid an
orthographically similar replacement name.
Intaglicollonia is monotypic and its type species name
retains the specific epithet originally proposed under
Nehalemia by Hickman (1974). Although the genus is
monotypic, it is the basal member of a putative evolu-
tionary lineage in which intraspecific sculptural polymor-
phism has been studied, and named patterns of incised
lines have an ontogenetic order of appearance in individ-
uals (Noda and Ogasawara, 1976). Further study of vari-
ation in the onset of timing of the incised patterns has led
to a model of heterochronic evolution in the lineage
(Majima and Murata, 1992). The genus also has figured
in the identification of the late Eocene origins of north
Pacific deep-water communities (Hickman, 1984), in
the characterization of biotic response to the Eocene-
Oligocene climatic transition (Hickman, 2003), and in
the reconstruction of patterns of trans-Paeifie migration
(Hickman, 2003; Amano, 2005).
Blake’s (1973) genus is also monotypic and of consid-
erable interest as a component of the famous Mist fauna
in northwest Oregon in the upper part of tire middle
member of the Keasey Formation. The Mist locality is
noteworthy as a Tertiary erinoid Lagerstiitte (Moore and
Vokes, 1953) as well as for its well-preserved asteroids,
ophiuroids, and echinoids (Burns and Mooi, 2003). The
carbonate layers at the Mist Locality are also noteworthy
for providing petrographic and isotopic evidence of meth-
anogenesis, possibly contemporaneous with the fauna and
a potential cold seep environment (Burns et al., 2005).
New illustrations of the holotype of Intaglicollonia
hieroglyphica are presented (Figures 1, 2) along with a
detail of the fine engraved lines (Figure 3), which were
interpreted originally as an unusual form of discor-
dant ( non -coll abral) secretory activity by the mantle
(Hickman, 1974). Discussion with Jonathan Todd (The
Natural History Museum, London) raises the interesting
possibility that the lines are “pseudoetchings” ( sensu
Palmer and Plewes, 1993; Todd and Palmer, 2002) gen-
erated as an active secretory response to the presence of
a thin, tubular commensal cnidarian. In this instance the
shallowly-incised lines would represent systematic and
continuous growth response by the snail to deformation
of the periostracal sheet at the growing shell margin.
Similar active deformation responses have been reported
in fossil bivalves (Todd, 1993).
Intaglicollonia , Phanerolepida Dali, 1907 and
Cantrainea Jeffreys, 1883 are known only from deep-
water faunas and sedimentary facies (>200 m) and are
distinguished from the shallow-water “dwarf turbans’ of
the colloniine genus Homalopoma Carpenter, 1864 by
their larger size and lack of nodulose spiral ribs sepa-
rated by deep grooves. Shells of the Mediterranean type
species, Homalopoma sanguineum (Linnaeus, 1758) are
6-7 mm in diameter and red or pink in color, in contrast
to the unpigmented shells of the larger-shelled species
in the deep-water genera.
The replacement name Intaglicollonia is chosen to call
attention to the finely-incised ( intagliare ) markings on
the shell surface as well as emphasizing the taxonomic
position of the genus in Colloniidae Cossmann, 1916, a
basal gastropod family group with a concentration of
morphological diversity in the Cenozoie fossil record
(Hickman and McLean, 1990). Publication of the replace-
ment name is prerequisite to a forthcoming reinterpreta-
tion of the taxonomy, paleoeeology, and unusual features
that Intagliocollonia shares with fossil and living species
of Cantrainea and Phanerolepida.
ACKNOWLEDGMENTS
I am grateful to Richard Squires and an anonymous
reviewer for their suggestions for improvement of the
manuscript. I thank Jon Todd for introducing me to
pseudoetchings and criteria for recognizing periostracal
deformation structures. Thanks also go to D. Smith
for assistance in preparing the Figures. This is con-
tribution 2047 of the University of California Museum
of Paleontology.
LITERATURE CITED
Amano, K. 2005. Migration and adaptation ol Late Cenozoie
cold-water molluscs in the North Pacific. In: A.M.T
Elewa (ed.) Migration of Organisms. Springer, Berlin
pp. 128-150.
C.S. Hickman, 2013
Page 161
Figures 1-3. Holotype (USNM 647339) of Intaglicollonia hieroglijphica Hickman, 1979; height = II mm, diameter = 16 mm
1. Apical and 2. Ahapertural view, 3. Detail of incised sculpture on body whorl.
Blake, D.B. 1973. Ossicle morphology of some recent asteroids
and description of some West American fossil asteroids.
University of California Publications in Geological Sci-
ences, 104: 1-59.
Burns, C. and B. Mooi. 2003. An overview of Eocene-
Oligoeene echinoderm faunas from the Pacific Northwest.
In: D.R. Prothero, L.C. Ivany and E.A. Nesbitt (eds.)
From Greenhouse to Icehouse: The Marine Eoeene-
Oligocene Transition. Columbia University Press, New
York. pp. 88-106.
Burns, C., K.A. Campbell, and H. Mooi 2005. Exceptional
erinoid occurrences and associated carbonates of the
Keasey Formation (Early Oligocene) at Mist, Oregon,
USA. Palaegeography, Palaeoclimatology, Palaeoecology
227:210-231.
Carpenter, P. P. 1864. Supplementary report on the present
status of our knowledge with regard to the Mollusca of
the West Coast of North America. Report of the 33rd
Meeting of the British Association for the Advancement
of Science (Newcastle-upon-Tyne), pp. 517-686.
Cossmann, M. 1916. [“1915”]. Essais de Paleoconchologie
Comparee. Vol. 10. Paris: Privately published, 292 pp.
Dali, W. H. 1907. Descriptions of new species of shells, chiefly
Buccinidae, from the dredgings of the U.S. “Albatross”
during 1906, in the northwestern Pacific, Bering, and
Japanese Sea. Smithsonian Miscellaneous Collections
50: 139-173.
Hickman, C.S. 1974. Nehalemia hieroglijphica , a new
genus and species of archaeogastropod (Turbinidae:
Homalopomatinae) from the Eocene of Oregon. The Veli-
ger 17: 89-91.
Hickman, C.S. 2003. Evidence for abrupt Eocene-Oligocene
molluscan faunal change in the Pacific northwest. In: D.R
Prothero, L.C. Ivany and E.A. Nesbitt (eds.) From Green-
house to Icehouse: The Marine Eocene-Oligocene Tran-
sition. Columbia University Press, New York. pp. 71-87.
Hickman, C.S. and | H. McLean. 1990. Systematic revision
and suprageneric classification of trochacean gastropods.
Natural History Museum of Los Angeles County Science
Series 35: 1-169.
Jeffreys, J.C. 1883. On the Mollusca procured during the
Lightning’ and ‘Porcupine’ Expeditions, 1868-70. Part 6.
Proceedings of the Zoological Society, London 1883,
pp. 88-115.
Linnaeus, C. 1758. Systema Naturae per Regna tria Naturae 1.
Regnum Animale. (10dl Edition) Salvius, Stockholm, Sweden.
Majima, R. and A. Murata. 1992 Intraspeific variation and
heterochrony of Phanerolepida pseudotransenna Ozaki
(Gastropoda: Turbinidae) from the Pliocene Nobori
Formation, Pacific side of southwestern Japan. Transac-
tions of the Paleontological Society ol Japan, N.S. 165:
1024-1039.
Moore, R.C. and HE Vokes. 1953. Lower Tertiary erinoids
form northwestern Oregon. U.S. Geological Survey Pro-
fessional Paper 233-E: 113-148.
Noda, H. and K Ogasawara 1976. On the variation of
Phanerolepida rehderi MacNeil Contributions from the
Department of Geology and Mineralogy, Niigata Univer-
sity 4: 431-436. (in Japanese with English abstract)
Palmer, T.J. and C. Plewes. 1993. Borings and bioerosion in
fossils. Geology Today 9: 138-142.
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illustrated by the Late Jurassic gryphaeoid, Deltoideum
delta (Smith). Scripta Geologica, Special Issue 2: 417-433.
Todd, J A. and T.J. Palmer. 2002. Pattern versus process or
informative versus ininformative Ichnotaxonomy: com-
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Carole S. Hickman
University of California
Department of Integrative Biology
and
Museum of Paleontology
Berkeley, California 94720-3140 USA
[email protected]
u
THE0NAUTILU S
Volume 127
2013
AUTHOR INDEX
Angulo-Campillo, 0 85
Araya, J.-F 115
Bertsch, 1 1 85
Bolton, M | 65
Chang, Y.-W. 19
Chen, T.-C 19
Dourson, D.C 153
Fernandes, MR 1
Geiger, D.L 115
Glover, E. A 101,131
Gregg, B.C 156
Grego, J 78
Harasewych, M.G 43, 147
Hossack, B.R 40
Kantor, Yu. 1 43,57
Kosyan, A 57
Kronenberg, G.C 36
Leal, J 11 1
Minton, R.L 119
Mok, II K 19
Nekola, | C 107
Nelson, T.A 156
Paustian, M.E 90
Pearce, T.A 90
PlMENTA, A.D 1
Petit, R.E 147
Portell, RAY 65
Quinta, M.H 29
Rosenberg, G 107
Sasaki, T. 93
Sato, K 93
Snyder, M.A 125
Steffek, | 78
Taylor, J.D 101, 131
Thompson, F.G 153
Watanabe, 11 93
Watters, G.T. 78
Wl LEAN, R.C 19
Williams, S.T. 131
Yu, M il 19
NEWTAXA PROPOSED IN VOLUME 127
GASTROPODA
Aidernofusus Kosyan and Kantor, 2013, new genus (Buccinidae)
Aidemofusus ignotus Kosyan and Kantor, 2013, new species (Buccinidae)
Comisepta guzmani Araya and Geiger, 2013, new species (Fissurellidae)
Eucalodiurn belizensis Thompson and Dourson, 2013 (Euealodiidae)
Extractrix dockeryi Harasewych and Petit, 2013, new species (Caneellariiidae)
Fusilaria Snyder, 2013, new genus (Fasciolariidae)
Fusilaria garciai Snyder, 2013, new species (Fasciolariidae)
Intaglicollonia Hickman, 2013, new name (Coloniidae)
Isotriphora onca Fernandes, Pimenta, and Leal, 2013, new species (Triphoridae)
Isotriphora tigrina Fernandes, Pimenta, and Leal, 2013, new species (Triphoridae)
Lithasia bubala Minton, 2013, new species (Pleuroceridae)
Marionia kinoi Angulo-Campillo and Bertsch, 2013, new species (Tritoniidae)
Omphalotropis ilapiryensis Pearce and Paustian, 2013, new name (Assimineidae)
Strebela Kantor and Harasewych, 2013, new genus (Buccinulidae)
Vertigo marciae Nekola and Rosenberg, 2013, new species (Vertiginidae)
Weilandipoma attduboni Watters, Grego, and Steffek, 2013, new species (Annulariidae) .
Weilandiporna inacaijaense Watters, Grego, and Steffek, 2013, new species (Annulariidae)
, 58
. 58
116
153
149
125
126
160
. 10
. . 8
121
. 85
. 90
, 53
109
. 81
, 82
BIVALVIA
Clathrolucina Taylor and Glover, 2013, new genus (Lucinidae) 132
Ferrocina garciai Taylor and Glover, 2013, new species (Lucinidae) 137
Nucinella giribeti Glover and Taylor, 2013, new species (Nucinellidae) 102
Solemya (solemt/a) flava Sato, Watanabe, and Sasaki, 2013, new species (Solemyidae) 94
Striostrea paucichomata Bolton in Bolton and Portell, 2013, new species (Flamingostreidae) 68
REVIEWERS FOR VOLUME 127
Amano, Kazutaka
Bieler, Rudiger
Bogan, Arthur E.
Breure, Brain
Cameron, Robert
Coan, E.V.
Cowie, Robert
Cuezzo, M.G.
Cummings, Kevin
Dillon Jr., Robert T
Harasewych, M.G.
Healy, John
Kiel, Steffen
Herbert, Gregory S.
Kano, Yasunori
Krylova, Elena M.
Landau, Bernard
Lee, Harry G.
Lima, Silvio F.B.
Lyons, Williams G.
Marshall, Bruce A.
Mikkelsen, Paula M
Nekola, Jeffrey C.
Pernet, Bruno
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CULTURE
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DIVISION of CULTURAL AFFAIRS
Seddon, Mary
Slapcinsky, John
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Von Proschwitz, Ted
Zardus, John
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